Plant Production Science

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Growth and Physiological Characteristics of Rice

Seedlings Raised with Long Mat by Hydroponics:
—Comparison with young seedlings raised in
soil—

Yingdian Wang, Kohei Tasaka, Akio Ogura & Sachio Maruyama

To cite this article: Yingdian Wang, Kohei Tasaka, Akio Ogura & Sachio Maruyama (1999)
Growth and Physiological Characteristics of Rice Seedlings Raised with Long Mat by
Hydroponics: —Comparison with young seedlings raised in soil—, Plant Production Science,
2:2, 115-120, DOI: 10.1626/pps.2.115

To link to this article: https://doi.org/10.1626/pps.2.115

© 1999 Crop Science Society of Japan

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 Plant Prod. Sci. 2 (2) : 115-120 (1999)

Growth and Physiological Characteristics of Rice Seedlings

Raised with Long Mat by Hydroponics
-Comparison with young seedlings raised in soil-

Yingdian Wang, Kohei Tasaka *, Akio Ogura ** and Sachio Maruyama **

(Beijing Normal University, Beijing 100875, China;

* Kyushu National Agricultural Experiment Station, Chikugo, Fukuoka 833-0041, Japan;
**National Agriculture Research Center, Tsukuba Science City, Ibaraki 305-8666, Japan)

Abstract: Growth and physiological characteristics of rice seedlings raised with long mat by hydroponics (SLH) were
compared with those of young seedlings raised in soil (YSS). When SLH and YSS were raised under the similar
environmental conditions, the dry weight of shoot in SLH was significantly lighter than that in YSS at the time of
seedling harvest, whereas the shoot in SLH was as long as that in YSS. The increase of dry weight of shoot and the
decrease of dry weight of grain were smaller in SLH than in YSS during the raising of seedlings. In addition, the
content of soluble sugars in the grain of SLH was apparently lower than that of YSS, indicating that the supply of
carbohydrates from endosperm to shoot was less in SLH than in YSS. Crop growth rate and mean leaf area index of
SLH were lower than those of YSS, while net assimilation rate of SLH was similar to that of YSS. The size of leaf blades
in SLH was markedly smaller than that in YSS, whereas no significant difference was observed in the rates of
photosynthesis and dark respiration in leaf blades between SLH and YSS. The results suggest that the leaf growth of
SLH is reduced by the insufficient supply of carbohydrates from endosperm, and this is the major cause of lower levels
of dry matter production in SLH as compared with that in YSS.

Key words: Growth, Hydroponics, Leaf area, Oryza sativa L., Raising of seedlings, Rice, Soluble sugars.

A system for the mechanization of rice cultivation
based on medium-sized machines was developed in the
1970's in Japan (Kushibuchi, 1988; Tasaka et a1.,
1996). In the medium-sized mechanization system, the
transplanting machine together with the head-feeding
combine contributed markedly to the saving of labor
(Kushibuchi, 1988). However, the labor for raising and
transplanting of seedlings still accounts for one fourth of
the total labor hours required for rice cultivation
(Ministry of Agriculture, Forestry and Fisheries, 1998).
Moreover, for machine transplanting, seedlings are
raised in a nursery box filled with 6 to 8 kg of soil, and
200 nursery boxes are required for transplanting in one
hectare of paddy field (Tasaka et a1., 1996). Since
manual handling of the nursery boxes is necessary sev-
eral times from sowing to transplanting, the handling of
thousands of nursery boxes is the heaviest labor in rice
cultivation, especially for large scale rice farms. Thus, it
is still necessary to save labor in raising and transplant-
ing of seedlings to enlarge farm size in the machine
transplanting system.
Yokoyama and Nakanishi (1990) investigated a rais-
ing and transplanting system for rice seedlings without
nursery boxes. In this system, seeds are sown in soil with
an aluminum frame that is 30 cm in width and 400 cm in
length, and special devices are used for soil filling, sowing
Received 21 July 1998. Accepted 25 December 1998. Corresponding author: S. Maruyama (samaru@narc.affrc.go.jp, fax +81-298-38-
8952) .
Abbreviations: CGR, crop growth rate; DW IL, ratio of dry weight of shoot to plant length; EC, electric conductivity; LAI, mean leaf
area index; SLH, seedlings raised with long mat by hydroponics; NAR, net assimilation rate; YSS, young seedlings raised in soil.
and covering with soil in the frame. Although the seed-
lings raised by this method were suitable for machine
transplanting, harvest and mounting of the seedlings on
a transplanting machine have remained as big problems,
because the mat of seedlings with soil was too heavy for
manual handling (Yokoyama and Nakanishi, 1990).
Tasaka et al. (1996) developed a new technique using
a long mat for raising and transplanting hydroponically
grown rice seedlings. Seedlings are raised by hydroponics
in a nursery bed that is 10 times longer than the nursery
box for the conventional transplanting system. After
raising of seedlings, the mat with seedlings is rolled up,
mounted on a transplanting machine, and transplanted
with special attachments (Tasaka et al., 1997). In this
technique, no manual handling of seedlings is necessary
before the harvest of seedlings. In addition, as the weight
of the mat of seedlings is only one fifth of the young
seedlings with soil, it is possible to mount a large number
of seedlings on the transplanting machine and to reduce
supply frequency of seedlings during transplanting
(Tasaka et al., 1996). Thus, the technique dramatically
reduced manual handling for raising and transplanting
of the seedlings as compared with the conventional
machine transplanting. However, it has been reported
that the dry weight of the seedlings at transplanting was
lighter and tillering after transplanting was inferior in the

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 116 Plant Production Science Vol. 2, 1999
seedlings raised by hydroponics than in the young seed-
lings raised in soil (Tasaka et aL, 1996; Tasaka et aL,
1997). These reports indicate that the method of raising
of seedlings by hydroponics has not yet been established
for rice cultivation.
In the present study, we investigated the growth
characteristics of rice seedlings raised with long mat by
hydroponics to develop a technique of raising of seed-
lings under various transplanting conditions. First of all,
we compared the characteristics of the seedlings raised
with long mat by hydroponics (SLH) with the young
seedlings raised in soil (YSS) at the time of seedling
harvest. Next, we compared SLH with YSS for their
growth, dry weight and soluble sugar contents. Finally,
we analyzed the difference between the two kinds of
seedlings in photosynthesis, dark respiration and the
contents of chlorophyll and soluble proteins in leaf
blades.
Materials and Methods
1. Plants and growth conditions
Experiments were conducted in the vinyl houses at
National Agriculture· Research Center in 1996 and 1997.
Rice (Oryza sativa L., cv. Kinuhikari or Akitakomachi)
seedlings with long mat were hydroponically grown with
the continuous-flow-solution system previously described
(Tasaka et al., 1996). Briefly, presoaked seeds were sown
at a rate of 1.25 kg dry seeds m- 2 on a nonwoven cloth
(Oikos AM20300275, 30 g m- 2, Nisshinbo Industries,
Tokyo, Japan) in a nursery bed (600X28X5 cm) and
germinated in tap water. After 5 days from sowing,
fertilizer for hydroponics (Ootsuka No.1 and 2, Ootsuka
Chemical, Osaka,japan) was applied to the tap water in
the nursery bed. This fertilizer was selected for the
reduction of labor in the present study. Nutrients in the
solution were 91.5 ppm N (9.0 ppm NH 4 -N and 82.5
ppm N0 3 -N), 110.0 ppm P20S' 195.1 ppm K 20, 66.6
ppm CaO, 39.8 ppm MgO and 0.7 ppm Fe-EDTA. The
preliminary experiments showed that these are the opti-
mum concentrations for the growth of shoot and root of
the seedlings. The nutrient concentration of the culture
solution was maintained by the addition of the fertilizer
by adjusting the EC value to the original level (1.35 mS
cm- I ) every day. The pH of tap water and the nutrient
solution was not adjusted, but the pH of tap water was
about 7.5 and that of the nutrient solution was in the
range of 6.5 -7.0. Tap water or the culture solution was
circulated in the nursery bed at a flow rate of 1 L min- 1 •
Temperature in the vinyl house was maintained at 18 to
33°C during the raising of seedlings.
The seedlings in the control group (YSS) were raised
in soil by the conventional method. Two-hundred grams
of dry seeds of the same cultivars as those used for SLH
were presoaked and sown in a nursery box (58 X 28 X 3
cm) filled with soil (Kumiai Ryujobaido K, Kureha
Chemical Industry, Tokyo, Japan). Nutrients in the soil
were 7.39 g m- 2 N, 14.78 g m- 2 P 20 S and 12.93 g m- 2 K 2
0. The sowing density was the same as that for SLH,
1.25 kg dry seeds m- 2. After sowing, the nursery box was
placed in a nursery chamber at 32°C and 100% relative
humidity in the dark for 2 days, and then transferred to
a nursery bed in the other vinyl house. The seeds for YSS
were sown earlier than those for SLH to make the growth
stage at harvest time uniform (Experiment I). In Experi-
ment II, the seeds for YSS were sown on the same day as
those for SLH, and the nursery box was placed at the
same height of the nursery bed for SLH in the same vinyl
house to maintain the similar environmental conditions.
Both experiments were repeated three times.
2. Measurement of growth and dry matter produc-
tion
Seedlings were sampled from five places in the nursery
bed for SLH and in the nursery box for YSS. Ten
seedlings were randomly selected from each place, and
the leaf age and the lengths of shoot and root were
recorded. The seedlings were washed with water and
separated into four parts : leaf blades, leaf sheaths, roots
and grains. After the area of the leaf blades was mea-
sured with photocopy, the separated plant parts were
dried at 90°C for 48 hr. The dry weight of the plant parts
was determined, and the values of the dry weight and the
leaf area were used for calculations of crop growth rate
(CGR) , mean leaf area index (LAI) and net assimila-
tion rate (NAR) .
3. Measurement of the rates of photosynthesis and
dark respiration
The rates of photosynthesis and dark respiration of
leaf blades in rice seedlings were measured by the
method previously described (Maruyama and Nakamu-
ra, 1997). The blades of the third leaves were sampled at
11 a.m. from SLH and YSS. Middle part of the leaf
blades (ca. 3 cm in length) was further excised with a
razor blade, and four leaf sections were placed in a
chamber of a leaf-disk oxygen electrode system (Model
LD-2, Hansatech Ltd. Norfolk, UK). The photosynthetic
rate was determined by measuring the rate of increase in
O 2 at a CO 2 concentration of approximately 10% under
a photon flux density of 600,umol m- 2 S-1. The dark
respiration rate was measured by the rate of decrease in
O 2 at a CO 2 concentration of 0.03% in the dark. Tem-
perature in the chamber was kept at 25°C.
4. Analysis of soluble sugars
SLH and YSS were separated into shoots, roots and
grains. The grains were hulled for analysis of soluble
sugars. After the fresh weight of each part was quickly
measured, the samples were stored at -40°C until fur-
ther analysis.
The samples were placed in 20 mM Hepes-NaOH
buffer (pH 7.4), heated at 90°C for 5 min, and then
ground on ice with fine quartz sand. After the
homogenate was centrifuged at 18,000 X g for 10 min at
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 Wang et al. --Rice Seedlings Raised with Long Mat by Hydroponics 117

Table 1. Growth characteristics of YSS and SLH at the same leaf age (Experiment I).
Kind of Length of shoot Dry weight (mg planc l ) DW/L t
Leaf age
seedlings (cm) Shoot Root Grain (mg cm- l )
YSS 3.2±O.1 12.7±O.2 13.7±1.7 3.5±O.5 7.0±O.l 1.13±O.13
SLH 3.2±O.1 ns l1.4±O.1** 7.8±O.5** 3.l±O.2 ns 8.7±O.7** O.66±O.05**
t DW/L, Dry weight of shoot divided by shoot length. The values of leaf age, shoot length and DW/L are
expressed as means ± S.D. of three groups of plants, and those of dry weight as means ± S.D. of five groups
of plants (lO plants per each group). ** Significant at the 1% level; ns, not significant at the 5% level.

Table 2. Growth characteristics of YSS and SLH sown on the same day and grown under the similar
environmental conditons (Experiment II).
Kind of Length of shoot Dry weight (mg plant- l ) DW/Lt
Leaf age
seedlings (cm) Shoot Root Grain (mg cm- l )

YSS 3.3±O.2 12.9±O.6 11.0±O.3 2.8±O.l 5.9±O.5 O.84±O.04

SLH 3.7 ±O.l ** 13.6±O.9 ns 9.3±O.5** 3.6±O.2** 5.4±O.2 ns O.70±O.05*
t DW/L, Dry weight of shoot divided by shoot length. The values of leaf age, shoot length and DW/L are
expressed as means ± S.D. of three groups of plants, and those of dry weight as means ± S.D. of five groups
of plants (10 plants per each group). *, ** Significant at the 5% and the 1% level, respectively; ns not
significant at the 5% level.

4°C, the supernatant was used for the analysis of soluble
sugars. Glucose, fructose and sucrose in the supernatant
were determined by.the coupled enzymatic assay method
(Bergmeyer, 1974). In the present study, the amount of
soluble sugars was expressed as the sum of glucose,
fructose and sucrose, because they are major compo-
nents of soluble sugars in rice plant (Murayama et al.,
1955; Murata et al., 1968).
5. Analysis of chlorophyll and soluble proteins
Chlorophyll and·· soluble proteins in leaf blades of
seedlings were measured by the methods previously
described (Maruyama et al., 1990). The leaf blades were
cut into pieces and homogenized in a pre-chilled 50 mM
Tris-HCl buffer (pH 7.5) containing I mM Na 2 -EDTA,
8 mM MgCl 2 and 2% (v/v) 2-mercaptoethanol. An
aliquot of the homogenate was extracted with acetone for
determination of the chlorophyll content by the method
of Arnon (1949).
For the measurement of soluble protein content, the
sample of leaf blades was homogenized in the same
buffer, and the homogenate was centrifuged at 18,000 X
g for 10 min at 4~C. The supernatant was used to
measure soluble proteins by the method of Lowry. et al.
(1951), with bovine serum albumin as a standard.
Results
1. Dry weight of seedlings at harvest
In Experiment I, the seeds for YSS were sown 8 days
earlier than those for SLH. SLH at the 14th day and YSS
at the 22nd day were at the same leaf age, and their
growth characteristics were examined (Table I). The
length and the dry weight of shoot were smaller and the
dry weight of grain.· was larger in SLH than in YSS,
whereas little difference was observed in the dry weight
of root. The value of the dry weight of shoot divided by
shoot length (DW/L), which is an indicator of seedling
quality (Matsushima, 1980), was significantly smaller in
SLH than in YSS.
In Experiment II, the seeds for SLH and YSS were
sown on the same day and both seedlings were raised in
the same vinyl house for 13 days. The growth character-
istics of the seedlings are shown in Table 2. The leaf age
of SLH was older than that of YSS at the harvest time.
Although the shoot length was similar, the dry weight of
shoot in SLH was significantly lighter than that in YSS.
Thus, the value of DW/L in SLH was smaller than that
in YSS. The dry weight of root in SLH was heavier than
that in YSS, but no significant difference was observed in
the dry weight of grain between them.
2. Changes in growth of rice seedlings
The time courses of the changes in leaf age, shoot
length, root length and the number of roots in Experi-
ment II are shown in Fig.l. The leaf age of SLH was
slightly older than that of YSS during the first 13 days
(Fig. IA). However, the lengths of shoot and root of
SLH were apparently smaller than those of YSS until 9
days after sowing (Fig. IB and IC). SLH grew rapidly
thereafter, and the shoot and root became longer than
those of YSS at harvest time. The number of roots of
SLH was also higher than that of YSS in the late phase
of the raising period (Fig. ID).
3. Changes in dry weight and soluble sugars in the
shoot, root and grain
Changes in dry weights of shoot, root and grain
accompanied with the progress of leaf age in Experiment

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 118 Plant Production Science Vol. 2, 1999

16 24 . . . - - - - - - - - - - - - - - ,
5
,,-....
A S
4 ()
12 20
'-"
(!)
...c
......
bl)
~ 3 e
4-< ~ 8
ro
(!) ......
0
....:I 2
.2
r:/) 4

o ._---L.--.&.---'
o 5 10 15
Days after sowing Days after sowing

10 10
1.0 2.0 3.0 4.0
,-,
8
C rn.
...... 8
S 0 Leaf age
0 0
'-" l-<
...c 4-<
6 Fig. 2. Changes in dry weight of shoot, root and grain of rice
...... 6 0
b!) l-< seedlings with the progress of leaf age. 0 and e, Shoot; 0
e (!)

~ 4
,.D
S 4 and +, root; 0 and _, grain; I:::. and A.., total. Open sym-
......
0 ;::l bols are for YSS, and closed symbols are for SLH. Bars
0 Z indicate standard deviations of results from four replicates,
~ 2 2
and they are not shown when smaller than symbols. Arrows
0 indicate the days after sowing (DAS).
0 5 10 15 5 10 15

Days after sowing Days after sowing

30 A
Fig. 1. Time course of the changes in leaf age (A), shoot length 25
(B), root length (C) and number of roots (D) of rice seed- ,,-....
,...., 20
lings. Open symbols are for YSS, and closed symbols are for I
15
SLH. Bars indicate standard deviations of the results from 30 ~
Q 10
plants, and they are not shown when smaller than symbols. Of)
Of)
5
8
'-"
Vl
I-<
0
II are shown in Fig. 2. In SLH, the dry weight of shoot ~ 60 B
;::l
Vl
increased and that of grain decreased more slowly than 50
~
in YSS throughout the raising period, whereas the dry .D
;::l 40
weight of root was similar to that in YSS. The total dry '0
C/)
30
weight of SLH was heavier than that of YSS at the early 20
stage, but no increase in total dry weight was detected in
10
SLH at the end of the raising period.
Changes in soluble sugars in shoot and grain were also o
0.0 1.0 2.0 3.0 4.0
examined (Fig. 3). The content of soluble sugars in the
grain of SLH was markedly lower than that of YSS, Leaf age
especially at the early stage in the raising of seedlings Fig. 3. Changes in soluble sugars in grain (A) and shoot (B) of
(Fig. 3A). In contrast, the content of soluble sugars in rice seedlings with the progress of leaf age. Open symbols are
the shoot of SLH was similar to that of YSS (Fig. 3B). for YSS, and closed symbols for SLH. Bars indicate standard
deviations of results from five replicates, and they are not
4. Growth analysis and physiological characteris- shown when smaller than symbols.
tics of leaf blade
CGR, NAR and LAI of SLH and YSS during the last
4 days of raising in Experiment II are shown in Fig. 4. Table 3, no significant difference was observed in the
CGR of SLH was 11 % lower than that of YSS. Although rates of photosynthesis and dark respiration between
NAR of SLH was 3% higher than that of YSS, LAI of SLH and YSS. The leaf areas and the contents of chloro-
SLH was 14% smaller than that of YSS. phyll and soluble proteins in the second- and the third-
The rates of photosynthesis and dark respiration of the leaf blades were also determined (Table 4). The leaf
third-leaf blades in SLH and YSS were measured at 25° area of SLH was markedly smaller than that of YSS,
C with a leaf-disk oxygen electrode system. As shown in while the chlorophyll content per mg fresh weight of leaf

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 Wang et al. --Rice Seedlings Raised with Long Mat by Hydroponics 119

blades in SLH was slightly lower than that in YSS, and also reported that the tillering in SLH after transplanting
no significant difference was observed in the content of was less active than that in YSS. Although the inferior
soluble proteins in the leaf blades between the two kinds growth of SLH might be partly caused by the mechanical
of seedlings. damages during transplanting, the inferior qualities of
SLH might affect the tillering ability after transplanting
Discussion
in their experiment. The increase in dry weight of shoot
Rice seedlings with high quality should be transplant- and the improvement of qualities in SLH are important
ed, especially in the area under unfavorable conditions for the growth at the early stage of tillering.
such as low temperature or low solar radiation. Matsu- Cereal seedlings grow heterotrophic only at an early
shima (1980) reported that healthy seedlings should be stage, depending entirely on the reserves in endosperm,
short and thick, and this characteristic of seedlings could and then enter a transition phase during which the
be expressed numerically as the ratio of the dry weight of growth is sustained both by the reserves in endosperm
shoot to the plant length (DW/L). In fact, the higher the and photosynthesis (Whalley et a1., 1966). Finally, seed-
DW/L of the seedlings, the more active the tillering after lings become autotrophic, depending totally on photo-
transplanting (Matsushima, 1980). Tasaka et a1. (1996) synthesis. Yoshida (1973) reported that the grain
investigated growth characteristics of SLH in comparison reserves mainly supported the growth of rice seedlings in
with YSS raised by the conventional method. According the early stage after sowing, and the seedlings became
to their experiments, the plant length was shorter and the autotrophic at a leaf age of 2.7. Therefore, the growth of
dry weight of shoot was lighter in SLH than in YSS. rice seedlings should be analyzed in relation to both
However, they failed to detect the difference in DW/L grain reserves and photosynthesis during the raising
between the two kinds of seedlings. In the present study, period. In the present study, the decrease in the dry
the DW/L in SLH was apparently smaller than that in weight of grain in SLH was slower than that in YSS, and
YSS in both Experiment I and II, indicating the qualities the levels of soluble sugars in the grain increased more
of SLH are inferior to that of YSS. Tasaka et a1. (1997) slowly in SLH than in YSS. The results suggest that the
supply of carbohydrates from the endosperm to shoot is
_____ 8 slower in SLH, and this causes the retarded growth of
40 8 100
----- 35
'( 100 '( 7 (;'7 86
SLH in the early phase of the raising period. On the
;>-, 89
30
<:1:1 ~6 I 6 other hand, growth analysis showed that CGR of SLH
"0 "0 100 103 8 5
C';l 25 N 5 N was lower than that of YSS, and the difference was not
8 20 S 4 S4
;:;-3 ascribed to NAR, but to LA!. These findings were
~ 15 ~3
0:: 10 0:: 2 j 2 confirmed by the measurements of leaf size and physio-
t? 5
U
<t:
1 1
logical parameters of leaf blades : the leaf size in SLH
0 Z 0 0
YSS SLH YSS SLH YSS SLH was smaller than that in YSS, while the photosynthetic
Fig. 4. Crop growth rate (CCR), net assimilation rate (NAR)
rate of leaf blades in SLH was similar to that in YSS. The
and mean leaf area index (LA!) of rice seedlings in the last 4 results indicate that the smaller leaf blades lowered the
days of the raising period. The numbers above bars indicate dry matter production of SLH in the late phase of raising
relative values to YSS. period. In conclusion, the leaf growth of SLH is reduced
by the insufficient supply of carbohydrates from en-
dosperm, and this is the major cause of lower levels of
Table 3. Photosynthesis and dark respiration of the third-
dry matter production in SLH than that in YSS.
leaf blades in YSS and SLH.
Although hydroponics for rice plant has been devel-
Kind of Photosynthetic rate Dark respiration rate oped based on the experiments on mineral nutrition after
seedlings (umol O 2 m- 2 S-I) (umol O 2 m- 2 S-I) transplanting (Kimura, 1931), few investigations have
YSS 12.3± 1.6 1.1 ±O.l been reported on hydroponics for raising of rice seed-
SLH 13.0± 1.7 ns 1.2±O.3 ns lings. In the present study, presoaked seeds were sown on
Values are expressed as means±S.D. of five independent a nonwoven cloth in tap water and the whole seedling
measurements. ns Not significant at the 5% level. was exposed to the light after sowing. Palmiano and

Table 4. Leaf size and contents of chlorophyll and soluble proteins of leaf blades in YSS and SLH.
Kind of Leaf area (cm 2 blade-I) Chlorophyll Soluble proteins
seedlings 2nd 3rd (flg mgFW-l) (flg mgFW-l)

YSS O.60±O.O8 2.39±O.l4 3.25±O.1O 1O.5±O.5

SLH O.l6±O.OI ** 1.08±O.O9** 2.9S±O.08* 9.5±O.9 ns
Values of leaf area are expressed as means ± S.D. of 10 replicates and those of chlorophyll and soluble
proteins as means ± S.D. of five replicates. *, ** Significant at the 5% and the 1% level, respectively;
ns not significant at the 5% level.

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 120 Plant Production Science Vol. 2, 1999
Juliano (1972) reported that the dry weight and the
starch content of grain decreased more slowly in the light
than in the dark. Therefore, less supply of grain reserves
to shoot in SLH may be ascribed to the exposure to light
of the seedlings. Nutrient concentrations also affect the
growth of rice plant (Kimura, 1931). In the present
study, rice seedlings were raised in the nutrient solution
developed for vegetable crops, because no commercial
fertilizer had been developed for raising of rice seedlings
with the continuous-flow-solution culture system. Since
the nutrient concentration was five to 10 times higher
than that for rice plant after transplanting (Kimura,
1931), the higher concentration of nutrients may also
affect the growth of SLH. Further investigations are
required to clarify the relationships between cultural or
environmental factors and growth of rice seedlings raised
by hydroponics.
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Kushibuchi, K 1988. Rice Cultivation in Japan. National Agricul-
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