J. Japan. Soc. Hort. Sci. 82 (4): 322327. 2013.

Available online at www.jstage.jst.go.jp/browse/jjshs1

JSHS 2013

Root-zone Cooling at High Air Temperatures Enhances Physiological Activities

and Internal Structures of Roots in Young Tomato Plants

Yasushi Kawasaki1,2*, Satoshi Matsuo1, Katsumi Suzuki1, Yoshinori Kanayama2

and Koki Kanahama2
1
NARO Institute of Vegetable and Tea Science, Taketoyo, Aichi 470-2351, Japan
2
Graduate School of Agricultural Science, Tohoku University, Sendai 981-8555, Japan

Low-cost technology is needed to alleviate high-temperature injury for high-yield greenhouse tomato production.
To acquire information about the physiological and morphological effects of root-zone cooling, we grew young
tomato plants for 2 weeks in nutrient solution held at about 25C, considered to be the optimum temperature for
tomato plants. We investigated plant growth, nutrient uptake, root activity (xylem exudation and root respiration
rate), root indole-3-acetic acid (IAA) concentration, and internal root structure. The root-zone temperature was
maintained at 24.7C by cooling, while the air temperature and control temperatures were higher than optimum
(30.8 and 33.7C, respectively). Root-zone cooling increased the relative growth rate (RGR) of roots compared
with the control, followed by shoot RGR. Root IAA was positively correlated with root RGR. Root-zone cooling
increased Ca and Mg uptake as well as root xylem exudation and respiration. It also advanced the development
of the internal structure of the xylem near the root tip. Thus, possibly by increasing root activity and root IAA,
root-zone cooling promoted root growth and nutrient uptake mediated by the development of the root xylem,
and thus shoot growth. These results suggest a physiological and morphological mechanism of growth
enhancement by root-zone cooling under high air temperature conditions.

Key Words: IAA, nutrient uptake, relative growth rate, root structure, root-zone cooling.

One such system, the root-zone cooling technique, is

Introduction
In tomato cultivation in the greenhouse, avoidance of
high temperatures is important in summer in Japan
because summer temperatures exceed the optimum for
tomato production. High temperatures reduce yields by
reducing pollen viability and fruit set (Sasaki et al., 2005;
Sato et al., 2006), causing cuticle cracking (Dorais et al.,
2004) and decreasing root activity (Gosselin and Trudel,
1983a, b; Klock et al., 1997).
Cooling technologies have been developed to permit
year-round production to alleviate high temperature
injury to tomato. Heat exchanger (De Gelder et al., 2005),
pad-and-fan, and fog (Peet and Welles, 2005) cooling
systems are sometimes used, but they are expensive to
run or not particularly effective under high humidity.
Therefore, effective and economic cooling systems are
needed. These are also important for decreasing CO2
emission to prevent global warming.
Received; June 3, 2013. Accepted; August 21, 2013.
* Corresponding author (E-mail: yask@affrc.go.jp).
proposed. In the case of tomato, when the root-zone
temperature was cooled to 25C under high air
temperatures, vegetative growth was improved by
increasing leaf area and plant height compared with
uncooled plants, and fruit yield was also increased
(Fujishige et al., 1991; Nkansah and Ito, 1994). In
addition Sasaki and Itagi (1989) reported that fruit yield
was increased by root-zone cooling at 20C in summer
tomato production. These increases of growth and fruit
yield were explained by enhancing photosynthetic
activities (Nkansah and Ito, 1995a), improving water
and nutrient uptake (Klock et al., 1997; Nkansah and
Ito, 1995b), and increasing root respiration rate (Klock
et al., 1997). These parameters were maximized at
around 25C root-zone temperature. However, little
knowledge comparing the partial difference in growth
and the nutrient content by root-zone cooling is available,
and physiological or morphological information is also
insufficient.
It is important to obtain fundamental knowledge about
the effects of root-zone cooling on plant growth as well

322
 J. Japan. Soc. Hort. Sci. 82 (4): 322327. 2013.
as to develop a practical root-zone cooling system.
Therefore, we investigated physiological and morpho-
logical changes at the early developmental stage of
tomato plants.
Materials and Methods
Plant materials, growth condition, and temperature
management
Seeds of tomato (Solanum lycopersicum L.)
Momotaro-Yoku (Takii seed Co., Ltd., Kyoto, Japan)
were sown in 72-cell trays filled with vermiculite, and
seedlings were grown for 3 weeks in a growth chamber
with day/night air temperatures of 25/20C. The
seedlings were then transplanted into a nutrient-film-
technique hydroponic system in a glasshouse on 6 July
2012. The glasshouse was ventilated when the
temperature exceeded 25C. The nutrient solution,
renewed every 2 weeks, contained 82 (gg1) N, 15 P,
134 K, 55 Ca, 12 Mg, 1.3 Fe, 0.4 Mn, 0.3 B, 0.04 Zn,
0.01 Mo, and 0.01 Cu. The root-zone cooling treatment
was started 5 days after transplanting, with the solution
temperature maintained at around 25C, and continued
for 14 days. The root zone of half of the seedlings
remained uncooled as a control. The average air
temperature in the glasshouse during the treatment period
was 30.8C, and the maximum and minimum
temperatures were 38.3C and 24.6C, respectively. The
nutrient solution temperature averaged 24.7C in the
cooling treatment and 33.7C in the control.
Measurement of xylem exudation, root respiration, and
dry weight
Plants were assessed on days 0 (with 79 leaves), 7
(1012 leaves), and 14 (1316 leaves) of cooling
treatment. Plants (n = 6) were cut off below the first true
leaves, the cotyledons were removed, and we collected
the xylem exudate for 10 min in absorbent cotton for
later weighing (Yamaguchi et al., 1995). Some well-
developed lateral roots (0.1 to 0.6 g of total dry weight)
from base to tip (n = 6 plants) were immersed in 500 mL
stirred nutrient solution, which was the same solution
as for cultivation, and the rate of oxygen consumption
was measured with an oxygen electrode (D-55;
HORIBA, Ltd., Kyoto, Japan) at 30C to calculate the
root respiration rate. Xylem exudation and the root
respiration rate were measured between 9:00 and 15:00.
Shoots and roots (n = 8) were dried at 80C for at least
48 h and then weighed for calculation of the relative
growth rate (RGR), which was calculated from the
equation:
RGR = (ln W2 ln W1)/(t2 t1)
where W2 and W1 were the dry weight at time t2 and t1,
respectively.
Mineral nutrient analysis
Fifty milligrams of dried samples of shoots and roots
323
were pulverized and ashed in 10 mL nitric acid and 2 mL
hydrogen peroxide at 180C for 20 min in a microwave
ashing system (START D; Milestone-general Co., Ltd.,
Kanagawa, Japan). P, K, Ca, and Mg were measured by
inductively coupled plasma atomic emission spectrom-
etry (SPS7700; Hitachi High-Tech Science Corporation
Co., Ltd., Tokyo, Japan). N was measured using a CN
analyzer (JM1000CN; J-Science Lab Co., Ltd., Kyoto,
Japan).
Quantification of IAA
For analysis of indole-3-acetic acid (IAA), about 1 g
fresh roots was homogenized in liquid nitrogen and
placed in 10 mL methanol/water/formic acid (15 : 4 : 1,
v/v/v). 13C6-labeled IAA (Cambridge Isotope Laboratories,
Inc., Tewksbury, MA, USA) was added to the extracts
as an internal standard. After overnight extraction at
4C, solids were separated by centrifugation and re-
extracted for 30 min in 10 mL of the same extraction
solution. To remove interfering compounds, extracts
were first passed through an Oasis HLB column (200 mg;
Waters, Milford, MA, USA) equilibrated with 1 M
formic acid. After further washing with 5 mL extraction
solvent, the combined eluate was evaporated; the residue
was dissolved in 5 mL of 1 M formic acid, and the
solution applied to an Oasis MCX column (150 mg;
Waters) equilibrated with 1 M formic acid. After further
washing with 5 mL of 1 M formic acid, the IAA was
then eluted with 5 mL methanol. The eluate was
evaporated, and the residue was dissolved in 3 mL of
1% acetic acid. The solution was applied to an Oasis
WAX column (60 mg; Waters) equilibrated with 1%
acetic acid. After further washing with 3 mL of 1% acetic
acid and then with 2 mL methanol, IAA was eluted with
6 mL of 80% methanol containing 1% acetic acid. The
eluate was evaporated at 40C under vacuum. The
residues were then dissolved in water/methanol/acetic
acid (80 : 19.95 : 0.05, v/v/v) and analyzed by high-
performance liquid chromatography (HPLC)/tandem
quadrupole mass spectrometry (MS/MS). The HPLC-
MS/MS system consisted of a Prominence 20A Series
HPLC (Shimadzu Co., Ltd., Kyoto, Japan), a 3200 QTrap
LC/MS/MS System (AB Sciex, Framingham, MA,
USA), and an electrospray interface. The purified
samples were injected onto a Shim-pack XR-ODS
column (2.2 m, 100 mm 2.0 mm; Shimadzu) at 45C
and eluted at a flow rate of 0.2 mLmin1. For
chromatographic separation, mobile phase A was water/
methanol/acetic acid (80 : 19.95 : 0.05, v/v/v) and mobile
phase B was methanol. The initial conditions were 100%
A for 2 min; linear change to 40% A/60% B over 7 min;
to 100% B over 10 min; and held at 100% B for 3 min.
Before each analytical run, the column was equilibrated
with mobile phase A for 10 min. IAA was quantified by
multiple reaction monitoring of the protonated intact
precursor ion [M + H]+ and a specific product ion, using
the following mass transitions: [13C6]IAA, 182.1 > 136.2;
 324 Y. Kawasaki, S. Matsuo, K. Suzuki, Y. Kanayama and K. Kanahama
IAA, 176.1 > 130.1. Data were analyzed using Analyst
v. 1.4.2 software (AB Sciex). Concentrations were
calculated from the peak area of the endogenous
compound, relative to the area of the internal standard.
Three biological replicates were analyzed for all samples.
Root morphology
On day 14, four lateral root samples near the root tip
from each treatment were fixed in FAA (40%
formaldehyde/acetic acid/ethanol/water = 1 : 1 : 9 : 9).
Then samples were dehydrated in a graded ethanol series
(70%, 80%, 90%, 100%), and embedded in Technovit
7100 resin (Heraeus Kulzer, Wehrheim, Germany). The
embedded samples were sectioned on a microtome at
2 m and stained with toluidine blue. The cross sections
were then observed and photographed under an optical
microscope (CX41; Olympus Co., Ltd., Tokyo, Japan).
Results
Growth and root activity
Although the shoot dry weight in the root-zone cooling
treatment was significantly less than that in the control
on day 7, the difference had disappeared by day 14
(Fig. 1A). There was no difference in shoot RGR during
day 0 to 7, but RGR was significantly greater in the
cooling treatment during day 7 to 14 (Fig. 1B). Root
dry weights did not show significant differences between
treatments on days 7 or 14 (Fig. 1C). Root RGR was
significantly greater in the cooling treatment during day
0 to 7, but there was no difference during day 7 to 14
(Fig. 1D).
The rate of xylem exudation did not change over time
Fig. 1. Effects of root-zone cooling treatment and time on (A) shoot dry weight (DW), (B) shoot relative growth rate (RGR), (C) root DW, and
(D) root RGR in tomato plants. Data are the means (SE) of eight independent measurements. *: significant at P < 0.05 by t-test.
in the control, but it was doubled on day 7 and quadrupled
on day 14 in the cooling treatment (Fig. 2A). The root
respiration rate increased in both the control and
treatment groups on day 7 and then decreased in both
on day 14. That on day 14 was significantly higher in
the cooling treatment group (Fig. 2B).
Determination of IAA and mineral concentration
IAA concentration was determined in roots under root-
zone cooling treatment in order to show the relationship
between IAA concentration and root growth. Although
there was no significant change in root IAA
concentration over time in the control, the concentration
increased significantly on day 7 and decreased on day
14 in the cooling treatment group. There was a significant
positive correlation between IAA concentration and root
RGR (Fig. 3).
Effect of root-zone cooling on mineral concentration
was investigated in shoots and roots (Table 1). Shoot Ca
and Mg concentrations were significantly lower on days
7 and 14 than on day 0 in the control. In contrast, these
decreases were not observed with cooling treatment, and
thus were higher than in the control. Shoot N
concentration was also significantly lower on days 7 and
14 than on day 0 in the control but not in the cooling
treatment group. Shoot P and K concentrations were
significantly lower on days 7 and 14 than on day 0 in
both treatments. Root Ca and Mg concentrations showed
the opposite tendency: they were significantly higher on
days 7 and 14 than on day 0 in the control but unchanged
with the cooling treatment, and thus lower than in the
control. Root K concentration was significantly lower
 J. Japan. Soc. Hort. Sci. 82 (4): 322327. 2013. 325

only on day 7 in the control and was unchanged in the
cooling treatment group. Root P concentration was
significantly lower only on day 14 with cooling
treatment. Root N concentration was unchanged. The
total uptake of N, Ca, and Mg, which were their content
per plant by multiplying each concentration by dry
weight, was significantly greater in the cooling treatment
than in the control group on day 14.
Internal root structures
We investigated not only dry weight or root activity,
but also internal root structures by observation of root
sections. Histological examinations revealed that xylem
vessel cells near the root tip were clearly larger in the
cooling treatment group (Fig. 4). That change was
observed among all sections and typical images are
shown in the figure.
Discussion
Photosynthetic activities are decreased at day/night
air temperatures of 40/23C in young tomato plants
(Nkansah and Ito, 1995a) and RGR is also decreased
over 30C air temperature (Gent, 1986). Pollen viability
and release of pollen grains were reduced at 32/26C
(Sato et al., 2006). Tomato fruit set was reduced at 34/
20C (Sasaki et al., 2005). Therefore, at an average of

Fig. 3. Correlation between root relative growth rate (RGR) and root
IAA concentration in control and root-zone cooled tomato plants
after 7 and 14 days of treatment. Data are means (SE) of three
(root IAA concentration) or eight (root RGR) independent
measurements. *: significant correlation at P < 0.05.

Fig. 2. Effects of root-zone cooling treatment and time on (A) xylem

exudation rate and (B) root respiration rate in tomato plants. Fig. 4. Cross-sections of root near the root tip of tomato plants from
Data are the means (SE) of six independent measurements. (A) the control and (B) root-zone cooling treatment on day 14.
*: significant at P < 0.05 by t-test. Scale bar = 50 m. Arrows indicate xylem vessels.

Table 1. Nutrient concentrations and total uptake in shoots and roots of tomato plants subjected to high (Control) and optimum (Cooling) root-
zone temperature.

Shoot conc. (mgg1 DW) Root conc. (mgg1 DW) Uptake (mg per plant)
Days
N P K Ca Mg N P K Ca Mg N P K Ca Mg
z
0 Control 39.4 a 5.9 a 51.2 a 16.5 a 5.6 a 41.8 a 8.7 a 32.2 a 5.2 b 7.4 b 143.2 a 22.0 a 178.5 a 56.4 a 20.8 a

7 Control 35.1 b 4.7 b 41.4 b 15.9 b 4.6 b 37.3 a 8.3 ab 23.4 b 12.0 a 9.7 a 354.9 b 48.9 b 406.1 b 157.3 b 49.5 b
Cooling 38.0 ab 5.0 b 40.4 b 18.6 a 5.6 a 43.4 a 7.2 bc 33.0 a 4.5 b 7.2 b 328.1 b 44.5 b 338.6 b 149.3 b 48.8 b

14 Control 35.0 b 4.8 b 40.9 b 16.0 b 4.5 b 38.8 a 8.2 ab 32.4 a 13.0 a 9.2 a 672.9 c 95.8 c 766.2 c 300.8 c 90.9 c
Cooling 37.9 ab 5.4 ab 38.3 b 21.5 a 5.4 a 41.0 a 6.2 c 35.0 a 5.3 b 6.5 b 807.2 d 112.7 c 786.4 c 426.8 d 112.4 d
z
Values within a column followed by the same letter are not significantly different at P < 0.05 by Tukeys multiple comparison test (n = 8).
 326 Y. Kawasaki, S. Matsuo, K. Suzuki, Y. Kanayama and K. Kanahama
30.8C, the air temperature during the treatment period
in our study was high enough to injure tomato plants.
The rates of photosynthesis and growth were
decreased at a root-zone temperature of 30C under high
air temperature conditions (Nkansah and Ito, 1995a). In
other plants, they were also decreased at high root-zone
temperatures (Benoit and Ceustermans, 2001; Du and
Tachibana, 1994; He et al., 2001; Wang and Tachibana,
1996). Although there have been studied at optimum air
temperature, root-zone temperature exceeding 30C
reduced nutrient uptake (Tindall et al., 1990) and yield
(Gosselin and Trudel, 1983a). In our experiment, the
average temperature was 33.7C in the control solution,
which is supposed to cause high temperature injuries,
but was 24.7C in the cooling treatment, close to the
optimum (Klock et al., 1997; Nkansah and Ito, 1995a, b).
Cooling treatment increased xylem exudation and the
root respiration rate (Fig. 2), which are indicators of root
activity (Yamaguchi et al., 1995). These results indicate
improved transport of water and nutrients (Klock et al.,
1997; Yamaguchi et al., 1995) and support the report
that water uptake by roots was maximized at 25C
(Nkansah and Ito, 1995b).
Root IAA was high on day 7 of cooling treatment.
This result suggests a response of roots to the optimal
root-zone temperature. There was a positive correlation
between root IAA and RGR (Fig. 3). In Arabidopsis,
root IAA promotes root growth (Novickien et al., 2010;
Ohashi-Ito et al., 2013). Therefore, this result confirms
the promotion of root growth during day 0 to 7 of cooling
treatment and agrees with the previous study (Nkansah
and Ito, 1994). On the other hand, root IAA was
decreased on day 14. Pressman et al. (1997) showed that
root RGR was decreased with root growth and our result
agrees with this observation. Cooling treatment
promoted the development of xylem tissue near the root
tip (Fig. 4). Although we could find no information on
the effect of root-zone temperature on root morphology,
the promotion of xylem development by auxin previously
reported in Arabidopsis (Deng et al., 2012; Ohashi-Ito
et al., 2013) could explain that the transient increase in
IAA promoted root xylem development.
Cooling treatment promoted shoot Ca and Mg
concentrations and total uptake of Ca, Mg, and N
(Table 1). These results are supported by a previous study
(Nkansah and Ito, 1995b). Although there have been
studied at the optimum air temperature, similar
tendencies were reported (Kabu and Toop, 1970; Tindall
et al., 1990). On the other hand, root Ca and Mg
concentrations were reduced. Although we could find
no information on the effect of root-zone cooling on
mineral concentrations in roots, these opposite results in
shoots and roots suggest that the transport of Ca and Mg
within the plant might be promoted by increased root
activity and xylem development at the optimal root-zone
temperature.
Shoot dry weight on day 7 of cooling treatment was
lower than the control (Fig. 1A). Although we could
find no information about shoot growth suppression by
root-zone cooling, the change of root-zone temperature
in a short time might cause shoot growth to be suppressed
transiently. Shoot RGR was higher during day 7 to 14
of cooling treatment (Fig. 1B). This result suggests that
shoot growth was promoted as a consequence of
enhanced root activities and nutrient uptake, and agrees
with a previous study (Nkansah and Ito, 1994). In this
research, cooling treatment was continued for 14 days.
Therefore, further study is needed to reveal the effect of
cooling for more than 14 days.
Our results show that root-zone cooling increased root
activity and transiently enhanced root IAA, which in
turn promoted root growth and nutrient uptake mediated
by the development of the root xylem, and thus promoted
shoot growth. This study could add physiological and
morphological knowledge about the mechanism of
growth promotion by root cooling at high air
temperature, and provide scientific evidence for the
effectiveness of cooling.
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