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To cite this article: Tohru Kobata & Murshidul Hoque (1999) Reduction in Leaf Water Potential
and Hydraulic Conductance of Young Rice Plants (Oryza Sativa L.) Grown in Wet Compacted
Soils, Plant Production Science, 2:1, 14-20, DOI: 10.1626/pps.2.14
Article views: 47
Crop Ecology
Reduction in Leaf Water Potential and Hydraulic Conductance
of Young Rice Plants (Oryza Sativa L.) Grown
in Wet Compacted Soils
Tohru Kobata and Md. Murshidul Hoque
Abstract: Lower leaf water potential (1/11) was observed in young rice plants grown in wet and highly compacted soil.
Our objectives were to establish why the reductions of 1/11 occurred and to find the effect of plant hydraulic
conductance on 1/11' One seedling of a lowland cultivar Nipponbare and an upland cultivar Senshou were grown in
plastic pipes with a diameter of 5.4 cm and a height of 45 cm filled with a soil mixture. Soil bulk density (SED) was
set at four levels and wet soil conditions were maintained. In a growth chamber 1/11' transpiration rate per unit leaf area
(T L), leaf area and root length were measured at 35 days after sowing. In both cUltivars, plant hydraulic conductance
(C p = - T L/ 1/11) decreased as SED increased and there was a positive correlation between C p and root length per unit
leaf area. The greater decreases in root length than in leaf area, in the soil with a high bulk density, were suggested
to reduce Cp , thus resulting in lower 1/11' In both cultivars, root hydraulic conductance per unit root length (C R ),
estimated using a pressure-flux method, increased with increase in SED. The increases in CR were accompanied by
the increases in T L per unit root length at a high SBD. We suggest that the suppression of root length in rice by highly
compacted soil causes lower C p which in turn reduces 1/11' even if the soil is wet. C R , water absorption rate per unit
root length and the diameter of the primary roots, clearly increased under highly compacted soil conditions, but this
might not be able to compensate for the greater reduction in root length than in leaf area, and so may not permit Cp
and 1/11 to be maintained.
Key words: Hydraulic conductance, Leaf water potential, Rice, Root length, Soil bulk density, Transpiration rate,
Wet soil conditions.
Lower leaf water potential (1/1'1) is observed under suppression of growth, such as reductions of leaf expan-
rhizosphere and aerial environmental stresses, such as sion rate, transpiration rate, stomatal conductance and
soil desiccation (Turner, 1997; Kramer and Boyer, biomass production in many plant species (Masle and
1995), soil salinity (Munns and Schachtman, 1992), Passioura, 1987 ; Smucker and Allmaras, 1992 ; Passiour-
toxicity by soil reduction under submerged soil (Hiras- a and Stirzaker, 1992). In rice plants, a reduction of 1/1'1
awa et al., 1992b), higher evaporative demand of atmo- was observed in the soil with high bulk density even if
sphere (Kramer and Boyer, 1995 ; Kobata et al., 1993a), the soil surrounding the roots was wetted (Hoque and
and strong winds of low temperature (Bolger et al., Kobata, 1998), while in other plants, such as wheat, no
1992 ; Fukuda et al., 1993) or of desiccated high temper- reduction of 1/1'1 in wet compacted soil was observed
ature air (Muramatu, 1982). The reduction of 1/1'1' (Masle and Passioura, 1987). Reduction of 1/1'1 may
caused particularly by higher evaporative demand under cause the suppression of growth in rice (Hoque and
moist soil conditions, seems to directly affect physiologi- Kobata, 1998), but the reason for the reduction of 1/1'1 is
cal processes such as stomatal conductance and not known.
photosynthetic rate, and consequently plant productivity Plant hydraulic conductance is one of the most impor-
(Boyer et al., 1980; Kobata et al., 1993b), although tant factors affecting 1/1'1' because 1/1'1 is a function of the
under desiccated soil conditions the physiological sup- transpiration rate if the hydraulic conductance is a
pression may not be caused by the change of 1/1'1 but constant (Gardner, 1968; Boyer, 1985). Under wet soil
predominantly by root dehydration (Davies and Zhang, conditions hydraulic conductance is a dominant factor in
1991) . determining 1/1'1' because the effect of soil water potential
Soil hardness is known to be an important factor in can be almost ignored.
Corresponding author: T. Kobata (kobata@life.shimane-u.ac.jp, fax + 81-852-32-6499) .
Abbreviations: fl' leaf water potential; Cp , plant hydraulic conductance; CR, root hydraulic conductance; SBD, soil bulk density; T L,
transpiration rate per unit leaf area.
Our objective was to establish why 1/1'1 of rice decreases beads to prevent soil evaporation. The transpiration rate
under wet compacted soil conditions, and to show the for each plant over 24 h was monitored using an electric
effect of plant hydraulic conductance on the reduction of balance. Transpiration rate scarcely changed during the
1/;'1 by using two cultivars which differ in their light period and was very small during the dark period
hydrological adaptation. (< 10% of the transpiration over 24 h), because evapor-
ative demand is determined mainly by radiation, humid-
Material and methods
ity and wind speed (Van Bavel, 1966; Kobata et al.,
1. Plant materials 1993a) and these are constant in a growth cabinet. The
The lowland improved nce cultivar Nipponbare (a transpiration rate thus was indicated by the rate per h
drought sensitive cultivar) (Kobata et al., 1996) and the during the 12 h of daylight.
upland local cultivar Senshou (a drought resistant Plant hydraulic conductance (C p ) was calculated
cultivar ) were used for the experiment. Polyvinyl chlo- from
ride pipes with an inside diameter of 5.4 cm and a height Cp =T L/ (1/I'soI1- 1/1'1) (1)
of 45 cm were filled with a mixture of top soil from where T L is the transpiration rate per unit leaf area
Andosols and granitic parent material of Cambisols, (g cm- 2 h- 1), and 1/I'soil is soil water potential, which is
which were sieved using a 0.5 cm mesh, in a volume ratio estimated to be approximately as zero under wet soil
of 1 : 1 on 23 August 1996; 0.11 g N as ammonium conditions (Gardner, 1968). We ignored the effects of
sulfate, 0.09 g P as superphoshate of lime and 0.09 g K 1/I'so11 on the estimation of C p , because bulk 1/I'so11 under
as potassium chloride were mixed with the soil in each the irrigated conditions, which was estimated from these
pipe. The bottom of the pipe was closed with a plastic soil water characteristic curves was at most within the
sheet and the outside wall of the pipe was covered by a range between 0.05 and 0.06 MPa at all SBD. Thus eq.
reflection sheet (Dry Sheet, Daiken Kogyo Co. Tokyo, (1) is rewritten as
Japan) to protect against heat gain from radiation. Soil C p =-T L/1/I'1 (2).
at a series of four soil bulk densities (SBD, 1.30, 1.35, 1. A direct relationship between T Land 1/1'1 was suggested,
40 and 1.50 dry soil g cm- 3 ) were set up by compacting because in our experiment TL was higher than 0.2
air dried soil with a steel stick after a small amount of g cm- 2 h- 1 at all SBD (Hirasawa and Ishihara, 1991).
water had been added. The actual SBD for the dry soil After measurement of 1/1'1' the plant shoots were cut
was calculated from soil volume and soil weight dried at near the ground surface and the pot was set intact into
lOSOC. Two seeds were sown and one plant was left to the stainless steel cylinder of a pressure chamber which
grow after establishment in each pipe and the SBD for was 50 cm high. The cut surface of the shoot was outside
each cultivar was replicated three times. Tillers appeared the cylinder and the base was closed with a silicon
were cut off weekly and only a main culm was grown. rubber stopper. The roots were pressurized with nitrogen
gas at the rates of 0.3 and 0.6 MPa . After the pressure
2. Measurements of leaf water potential and of 0.3 MPa was held for 30 sec to equilibrate the flow
hydraulic conductance from the root, the sap exuded from the cut shoot surface
All plants were grown in a non-temperature-controlled was gathered using a cotton pad for one minute, the
glasshouse for 25 days and transferred to a growth cotton was immediately placed in a plastic vial and
chamber (TGE-6H-4, Tabai Espec Co. Osaka, Japan) in weighed (Kobata and Kobayashi, 1993). Next, the
which room temperature was kept at 25 ± 1°C, relative pressure of 0.6 MPa was applied and the sap was
humidity at 75 ± 7%, light period at 12 h, and light gathered in the same manner. Flux from the root (F, mg
intensity at 300 ,!Lmol m- 2 S-1 at plant level. The amount min- 1 plane 1 ) was calculated from the differences in
of water lost over the previous 24 h was added every weight of the cotton pad before and after pressurization.
morning to maintain water content near the maximum Root hydraulic conductance (C R ) was estimated from
water holding capacity of the soil (volumetric soil water CR=F/P.RL (3)
content differed between 0.29 g cm- 3 at the lowest SBD where P is an applied pressure to the root system (MPa)
and 0.32 g cm- 3 at the highest SBD). Wet soil conditions (Turner, 1988; Kobata and Kobayashi, 1994) and RL is
were maintained during the measurements, because root length (cm plane 1) . The pressure under the two
transpiration loss from a pot over 24 h was less than 1% rates of P (0.3 and 0.6 MPa) were used to calculate CR'
of the water contained in the pot: the loss caused a There was a linear relationship between F and P when
reduction of only 0.005 MPa in soil water potential at the the P under -1 MPa was applied to rice roots grown in
most. After the plants had adapted to the environment of moist soils (Kobata and Kobayashi, 1994). Sap could be
the growth chamber for 10 days, 1/1'1 of the fully expand- gathered by the pressures > 0.3 MPa, because the appar-
ed top leaves of three plants at each SBD under light ent root bulk water potential (- 0.2 MPa or less), which
conditions was measured using a pressure chamber (Soil was measured just after removing the shoot from a plant,
Moisture Console, Model 305, Soil-Moisture Equipment was lower than the pressure applied. Generally root
Co. CA., USA) (Kobata and Takami, 1984). The surface water potential may be much higher than leaf water
of the soil in the pots was covered with 1 cm of plastic potential, since there is a negative gradient in water
potential along the plant organs (Kramer and Boyer, SBD pots after measurements of 1/1'1 and T L was
1995) . maintained at around 0.3 g cm- 3 (Fig. 1). When bulk
Leaf area for cut shoots was measured using an soil impedance was calculated from SBD and volumetric
automatic leaf area meter. After the measurements of C R , soil water content (Hoque and Kobata, 1998), it was
roots were washed and stained with 0.25% Coomassie between 0.17 to 0.70 kg cm-s.
Brilliant Blue R, and RL was measured with a laser area Shoot and root dry matter of both cultivars decreased
meter (CI-203, CID, Inc., Cambridge, England). Diam- as SBD increased (Fig. 2). The root-to-shoot ratio, which is
eter of primary root at the center position of the five thick indicated by the slope of a line connecting the origin and
roots in each plant was measured with a slide caliper. each symbol, decreased with the increments of SBD
Shoots and roots were oven dried at 80°C for 48 hand (only the lines for the lowest and the highest SBD rates
then weighed. are indicated in Fig. 2) : the root-to-shoot ratio decreased
from 0.38 to 0.17 in Nipponbare and from 0.68 to 0.44 in
Results
Senshou. Thus the reduction of the ratio in Nipponbare
Volumetric soil water content for both cultivars in all was greater than in Senshou. Both leaf area and total
.;.0
root length were suppressed by higher SBD, although
1=1
Ql
.-.. suppression was significantly less in Senshou than in
.;.0
1=1 "" 0.8
0
u SU
!-< bll
<il
.;.0
~
~ 0.6
III
~ U
~,......, s:l
0"" ~
0.4
~
"'d
S Ql
~
.....
!-<
U
bll
.S 0.2 80
~'-'
s .....
.....
0
.E0 'ClJ.
> 0 40
1.3 1.4 1.5
Soil bulk density (g em-S)
o L - -_ _...L..-_ _---I
20
15
+"
10
,.Q
bJJ
'0; 0.2
~
5
>.
~ 0 ~=--....J.-_--'-_ _.L.-.-..._---' o L - -_ _...L..-_ _---I
Nipponbare (Fig. 3). In both cultivars, RL per unit leaf when the rate of P was increased from 0.3 to 0.6 MPa
area was similarly reduced as SBD increased. (Fig. 6). Therefore C R is indicated by the slope of the
At higher SBD 1/1'1 and T L decreased similarly in both regression line. In both cultivars, CR increased in
cultivars, although 1/1'1 in Senshou was slightly higher at response to higher levels of SBD (Fig. 7). Transpiration
low SBD, and the reduction in Senshou more clearly rate per unit root length was accelerated by soil compac-
defined than for Nipponbare (Fig. 4). In both cultivars tion in both cultivars (Fig. 8). The close relationship
C p was reduced in relation to increase in SBD. The between C R and T L per unit root length (Fig. 9) indicat-
suppression of C p was accompanied by the decrease of ed that the hydraulic conductance of the root system
root length per unit leaf area (Fig. 5). affected the rate at which water was absorbed by roots.
There was a close linear relationship between F and P,
Discussion
In several crops, the hydraulic conductance of plant or
~ 1.3 1.4 1.5 plant organs affects 1/1'1 in dry soil (Saliendra and
:c -0.5 .---------,-----, Meinzer, 1992), in reduced and anaerobic soil (Hiras-
~o -0.6 rl1
(I)
~ ~ -0.7
t.)
~ '"d ~
~
C\i
-0.9 0=
t.)
.-;
0.05
cd
j -1.0" t.)
.I""'t ~
:: ~ 0.04
~
cd C'l
fo.t
0.04
'"d
~
S 0.03
t.)
,J:l
= --
btl
~
cd
P"""l 0.02
~ 6 10 14 18
Root length/leaf area (em em- 2)
Fig. 5. The relationship between plant hydraulic conductance
and root length per unit leaf area of Nipponbare(e) and
Senshou (.) under different soil bulk densities at 35 days
after sowing. The lines give the fitted regressions;
0'--------'------' C p =0.0024lRLjLA-0.0124 (r 2 =0.729, P<O.Ol).
Values are means ± one standard error of the mean of three
replications.
0.06
:::'
~
~~ 0.04 .I""'t
S 60
P-4 .-;
~ St.)
~ 0.02 bIJ
C,) S
<:.D
--
bJl
o
1.3
L - -_ _--L...-_ _----l
1.4 1.5
6
--
l"""'l
~
( I)
cd
fo.t
Soil bulk density (g cm- 3)
~
0
Fig. 4. Leaf water potential (-lfl) of a fully expanded top leaf, ~
~
0.2 0.4 0.6
transpiration rate (T L) and plant hydraulic conductance (C p ,
Pressure rate (MPa)
eq (2)) in Nipponbare (e) and Senshou (.) under various
soil bulk densities at 35 days after sowing. The curve lines give Fig. 6. Relationship between the flux of water per unit root
the fitted regressions ; length and applied hydrostatic pressure in the rice plants
1/;'1 = -0.0ISBD2+30.26SBD-20.78 (r 2 =0.679, P<0.05) grown under different soil bulk densities (e: 1.30, /:). : 1.35,
T L =0.I8exp( - 1.37SBD) (r 2 =0.72I, P<O.OI) D : 1.40 and.: 1.50 g cm- 3 ) in a pressure-flux experiment
Cp = -0.123SBD-0.21O (r 2 =0.827, P<O.OI). (eq (3)). Values are means and one standard errors of three
Values are means±one standard error of the mean of three replications. Root hydraulic conductance (C p ) was estimated
replications. from the slope of the linear regressions through the origin.
..cl
::' ~
b1l
.S100
-
l=l
Cl)
S 4
~ 80
~
0 .-.
-S
0 ~
(,) ~
~ 60 ..Cl 3
~
Cl)
Ca ..p.;I
~ 40 ~ (,)
~ ~
btl 2
S 20 l=l
0
'!"'l
M
6
o
..p.;I
,-i
~
L . . . -_ _--L..-_ _- - l '-'
~
.!"'l
~
1.3 1.4 1.5 rJJ
l=l
Soil bulk density (g cm- 3 ) ~
~
E-t 00 50
Fig. 7. The relationship between root hydraulic conductance (by 100
pressure-flux experiment) and soil bulk density in Nipponbar- Root hydraulic conductance
e (e) and Senshou (.&.) at 35 days after sowing. The lines
(10- 6 mg MPa- 1 cm- I min-I)
give the fitted regressions;
C R =0.000348SBD-0.000424 (r 2 =0.754, P<O.Ol). Fig. 9. Transpiration rate per root length and plant hydraulic
Values are means±one standard error of the mean of three conductance of Nipponbare (e) and Senshou (4.) under
replications. different soil bulk densities at 35 days after sowing. The lines
give the fitted regressions;
T L=15.4C R +0.0016 (r 2 =0.792, P<O.Ol).
,J:l Values are means ± one standard error of the mean of three
~
btl
l=l
4 replications.
Q,)
...-l
~
~
.-<
6 A
0 ::' 3 ..j.;,
-S
0 ~
~
..Cl ~
,.....,
~
P..
Cl)
C,) .-< 4
~
Q,)
(,) 2 ~ ;::
~
~ ~
.-<
~ btl C,) 03
;j p.,
l=l M '"d 2
0 6 ~ ~
.!"'l 0
~
,-i C,) ~
'-'
~ C,)
~
'!"'l ~~ 0
~
rJJ o'------'---------' ~ 0.06
0 400 800 1200 1600
l=l
~ 1.3 1.4 1.5 E ;:::;" B
~ ~ ;::
E-t ~
Soil bulk density (g cm- 3 ) ~
,.....,
.-<
03
p., ~ 0.04
Fig. 8. Transpiration rate per root length (TL/RL) and soil bulk
C'I
density of Nipponbare (e) and Senshou (.&.) at 35 days
after sowing. The lines give the fitted regressions;
S
~O. 02
T L/RL = 0.00066SBD - 0.00667 (r 2 = 0.915, P < 0.00 1) .
Values are means ± one standard error of the mean of three
replications. OL.--e---'----'------..J
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