J.

Agronomy & Crop Science (2010) ISSN 0931-2250

FLOODING STRESS

The Effects of Planting Pattern and Water Regime on Root

Morphology, Physiology and Grain Yield of Rice
A. Mishra & V. M. Salokhe
Agriculture Systems and Engineering, School of Environment Resources and Development, Asian Institute of Technology, Pathumthani, Thailand

Keywords
plant senescence; planting pattern; root
oxidizing activity; single seedling; system of
rice intensification; water regime
Correspondence
A. Mishra
Agriculture Systems and Engineering, School
of Environment Resources and Development,
Asian Institute of Technology, PO Box 4,
Klong Luang, Pathumthani 12120, Thailand
Tel.: +66-2-524-5477
Fax: +66-2-524-6200
Email: abhamishra@ait.asia
Accepted January 15, 2010
doi:10.1111/j.1439-037X.2010.00421.x

Abstract
Field experiments were conducted over two growing seasons to investigate the
effects of variations in water regime and planting pattern on the growth of rice
plant roots and shoots and on yield. Four water regimes were evaluated with
split plot design: intermittent flooding during the vegetative stage only (IF-V);
intermittent flooding extending into the reproductive stage (IF-R); not flooded
(NF); and continuously flooded (CF), interacting with three different planting
patterns: single seedling per hill with wider 30 30 cm spacing (P1); single
seedling per hill with closer 20 20 cm spacing (P2); and three to four seedlings per hill with 20 20 cm spacing (P3). The treatment combination CF/P3
corresponds most closely with current conventional practice. The other combinations were evaluated to contribute to a better understanding of the effects of
the two parameters studied, respectively and together. IF-V/P1 was considered
as an approximation of System of Rice Intensification (SRI) practice. This
study found that the combination of singly transplanted seedlings, both P1 and
P2, with the IF-V water regime improved root length density, root physiological activity, and chlorophyll content of the upper and lower leaves, leading to
higher grain yield compared with the other treatment combinations. With continuous flooding (CF), P2 gave 23 % more yield compared with the P3 planting pattern. Combining IF-V and P2 produced 32 % more grain yield
compared with the CF/P3 treatment. These results showed a synergistic effect
on grain yield from reduced intra-hill competition and IF-V water management. In these trials, there was no significant yield difference between the IF-V/
P1 and CF/P3 treatments. Wider spacing improved the performance of individual hills when grown under IF-V water regimes, but tiller number per unit area
remained a dominant determinant of yield. The yield reduction observed for
CF/P1 compared with CF/P3 indicated that in more hypoxic CF soils, denser
plant populations can produce more than sparser ones, whereas the latter benefit from more aerobic soil conditions. Intermittent irrigation during the vegetative growth stage and transplanting single seedlings/hill are major elements of
SRI methodology. These findings contribute to an understanding of why SRI
methods can produce the higher yields reported. A consideration of the effects
of interaction between planting pattern and water regime shows the need to
establish empirically the optimum values for these treatments according to
varietal, soil and climatic characteristics for the greatest yield response.

Introduction
Rice production needs to be increased by 50 % or more
above the current production level to meet the rising
368

food demand. Much of the demand will be arising from

Asia, which is the home of two-third of the worlds population and where 90 % of total rice supply is grown in
diverse environments (IRRI, 2008). Current challenges
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Effects of Some SRI Practices on Rice Growth

constraining farmers ability to meet the growing food

demand are: decreasing water resources, increasing costs
of cultivation, an apparent yield ceiling, and sustainable
environmental issues. These challenges have prompted
exploration of alternative crop management options,
including recently the System of Rice Intensification
(SRI).
Operationally, SRI includes careful, rapid and shallow
transplanting of very young seedlings, one per hill with
wider spacing (initially 25 25 cm and up to 50 50 cm
with very fertile soil), practising intermittent irrigation
during the vegetative stage or keeping the soil moist but
not continuously saturated, soil-aerating weeding, and
applying compost in preference to chemical fertilizer
(Stoop et al. 2002).
It has been reported that these practices are raising
yield by 50100 % or sometime more with reduced
requirements for water, seed, fertilizer and crop protection (Stoop et al. 2002). However, because of limited scientific understanding and research, the SRI approach has
attracted considerable controversy (Sheehy et al. 2004,
Sinclair 2004, McDonald et al. 2006; 2008, Uphoff et al.
2008). With these issues in mind, an earlier article (Mishra et al. 2006) reviewed the scientific literature relevant to
the management practices of SRI and found considerable
evidence to suggest that integration of the key practices of
SRI could improve the physiological efficiency of rice
plants.
Recently, experimental evidence has shown that SRI
practices can enhance the physiological efficiency of rice
plants of the same variety grown under SRI conditions
compared with currently recommended management
practices (Thakur et al. 2009a,b). However, the claimed
synergistic effects of SRI practice need to be studied by
examining the respective impacts of specific components
of SRI recommendations on plant growth.
Two components of SRI which have attracted particular interest are reduced water applications and sparser
planting, i.e., radically reduced plant populations. It has
been suggested that these two cultural practices can have
synergistic effects on root growth and subsequently on
grain yield (Mishra et al. 2006, Satyanarayana et al. 2006).
Ceesay et al. (2006) reported higher yield from transplanting single seedlings per hill and practicing intermittent irrigation during the vegetative growth stage. But the
mechanisms for getting higher yield under such conditions were not investigated.
Recently, the contribution of higher root length density
(RLD) to improved shoot growth during the vegetative
growth stage of rice has been reported in non-flooded soil
condition (Mishra and Salokhe 2008a). But what, if any,
effect of these adaptive root traits have on yield remains
to be determined. Lin et al. (2004) found that rice plants
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grown with shallow water depth and alternate wet/dry

water management (SWD) compared with conventional
water management had better ontogenesis when the
seedlings were transplanted at low densities. However, the
reasons for better ontogenesis also still need to be determined.
Similarly, San-oh et al. (2006) found that rice yield was
higher when single seedlings per hill were transplanted
compared with three seedlings per hill. The explanation
which they advanced was that single plants per hill had
higher cytokinin concentrations in their roots during the
late reproductive stage compared with plants grown three
per hill. Higher cytokinin concentration in the roots was
associated with delayed senescence of the plant and with
higher grain yield. However, the San-oh et al. (2006)
findings were observed under continuously flooded soil
conditions. How different planting densities affect root
and shoot growth under intermittent and non-flooded
regimes still warrants investigation.
Such investigation could contribute to better understanding and optimizing the effects of particular management practices such as those of SRI which have
attracted attention within the research and development
communities. This study was designed to investigate the
interaction effects of planting pattern and water regime
on rice root and shoot growth and on grain yield. It
was not assessing the System of Rice Intensification as a
system, but rather the measurable effects of two of its
key elements.
Materials and Methods
Experiments were conducted during the 2006 and 2007
dry seasons in a rice field at the Asian Institute of Technology in Bangkok, Thailand (1404N latitude and
10037E longitude, with 2.27 m altitude). The experiment was set up with a split-plot design, having four and
three replications, respectively, with water regime and
planting density as the main factor and sub-factor.
The soil composition was 66.2 % clay, 10.2 % sand,
and 23.2 % silt, with pH (1 : 1) of 5.0, and 1.38 %
organic carbon, 0.14 % nitrogen, 11 ppm available P,
212 ppm available K. Pathumthani, a locally-popular,
high-yielding, photo-nonsensitive variety with maturity
period of 120130 days, was the rice cultivar used for this
study.
Experimental plots were first puddled by flooding,
followed by wet ploughing and harrowing. Basal applications of cow manure at the rate of 10 tons ha)1 preceded
the land preparation activities. Inorganic nutrients,
16 : 16 : 16 (NPK) and DAP, were applied to all plots at
the rate of 180 kg ha)1. The total amount of DAP was
applied as a basal application, whereas NPK was top
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Mishra and Salokhe

dressed at 10, 25 and 45 days after transplanting (DAT).

Although the application of alternative kinds and
amounts of fertilization could have important interactive
effects with the two factors assessed here, fertilization was
not a variable in this study.
Seedlings used in this experiments were raised using
SRI seedbed procedures (Mishra and Salokhe 2008a), and
16-day-old seedlings were transplanted at a depth of
22.5 cm with three different transplanting patterns: P1
one seedling per hill with spacing 30 30 cm, termed the
wider spacing/planting pattern; P2 one seedling per hill
with spacing 20 20 cm, termed the narrower spacing/
planting pattern; and P334 seedlings per hill with spacing 20 20 cm, termed the crowded planting pattern.
The respective plant populations per m2 were 10, 25 and
8090. A shallow water ponding depth (23 cm) was
maintained during the transplanting period, and gap-filling operations were carried out during the first week after
transplanting.
After disappearance of transplanting shock (7 DAT),
the following irrigation treatments were carried out:
1. IF-V: intermittent flooding during the vegetative stage
a water level 33.5 cm from the soil surface was maintained for 10 days, then the field was drained and kept
non-flooded for 5 days; water was reapplied on the sixth
day of draining to re-establish a 33.5 cm ponded depth,
and this level was maintained for 10 days. Three similar
draining cycles were performed throughout the vegetative
growth period, and after panicle initiation (PI), a 5 cm
ponded depth of water was maintained until 15 days
before harvesting.
2. IF-R: intermittent irrigation extended into the reproductive stage the irrigation process was similar as IF-V
but with five draining cycles.
3. NF: non-flooding water regime field was kept moist
but not inundated.
4. CF: continuously flooded depth of 5 and 10 cm
ponded water was maintained during vegetative and
reproductive stages, respectively, until 15 days before harvesting.
Monthly mean air temperature ranged from 28 to
32 C during the growing seasons. Average total rainfall
during the experimentation period, i.e., March to June,
was about 103 mm. During the experimentation period,
rainfall was concentrated in late MayJune, i.e., the
normal season of monsoon in Thailand. Irrigation management was controlled successfully during the draining
cycle in the IF-V and IF-R treatments where soil moisture content in the rhizosphere (to 20 cm soil depth)
reached 5060 % of the saturated moisture content on
the sixth day of the draining period. A moisture meter
(wet sensor, type WET-2; Delta-T Device Ltd, Cambridge, England), calibrated for clay soil, was used
370

periodically to measure soil water content. During the

rainfall period, i.e., in the late reproductive phase, water
level was maintained by removing excess water from the
field in IF-V, IF-R and NF treatments, but not in the
CF treatments.
Daily consumption of water in CF was expectedly
higher compared with IF-V and NF. This was resulting
from the fact that the hydraulic head in CF was always
higher than in NF and IF-V. Consumption of water was
least in NF, where only 11.5 cm water was applied,
either on alternate days or on an as-needed basis to
ensure aerobic soil condition. This resulted in 6070 % of
saturated moisture content in the rhizosphere (to 20 cm
soil depth). In IF-R treatments, consumption of water
was increased because of the higher number of draining
cycles (five).
The last two draining cycles coincided with the flowering and grain filling stages when crop water demand was
at maximum. As a result of the high transpiration rate
during this period, soil moisture content dropped sharply
after draining and reached 4555 % at 20 cm soil depth.
This facilitated the development of cracks in the soil,
resulting in higher percolation rate of water in the IF-R
treatments.
Plant parameter measurements
Agronomic parameters
Yield components and grain yield were measured at
maturity. Numbers of productive tillers per m2, spikelets
per panicle, number of ripened grains per panicle, single
grain weight (mg), and 1000 grain weight (g) were all
recorded. Moisture content of grain was measured using
Kett Grain Moisture Tester, Grainer II PM-300 (Kett
Electric laboratory, Santiago, USA), and yield per hectare
was recorded as [(100 ) moisture content of the sample) fresh grain weight]/86 to convert the sample to
14 % moisture content.
Root parameters
For root study, two adaptive root traits were examined:
RLD, and root oxidizing activity. A soil-root core sample
was taken from 1520 cm and 3540 cm soil depth at
flowering (F) and at 20 days after flowering (20 DAF)
using cylindrical tubes (5 cm diameter by 90 cm length).
Roots were washed with water and cut into small pieces.
The root length was calculated using the line intersection
method described by Tennant (1975). RLD was calculated
by using the formula: RLD = RL/V, where RL = root
length, and V = volume of the soil core. Root oxidizing
activity rate was measured at flowering and at 20 days
after flowering by assaying the oxidation of alphanapthylamine (Zhang et al. 1994).
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Effects of Some SRI Practices on Rice Growth

Chlorophyll content
Chlorophyll contents of the flag leaf and third leaf were
recorded from flowering to physiological maturity stages
at intervals of 7 days from flowering stage to physiological maturity, using a chlorophyll meter (SPAD 502;
Minolta Corp; Tokyo). Prior to use, it was calibrated by
using spectrophotometric assays to determine the exponential equation to directly convert its output to leaf
chlorophyll concentration as described by Markwell et al.
(1995).
Statistical analysis
Data were subjected to Levenes test option of sas to
check the homogeneity of variance. The data were subjected to combined (pooled) analysis because measured
parameters showed no significant differences between
2006 and 2007. Data were analysed using the PROC GLM
procedure in sas (SAS Institute, 1999) to determine
single-factor and interaction effects. Computed F values
for some of the important plant parameters are presented
in Tables 1 and 2. Whenever significant interaction was
observed between factors, the level of one factor was
compared with each level of the other factor by DMRT

procedures. All data are presented as means S.E. A significance level of 0.05 was used for evaluating all analyses.
Results
Crop growth and yield
Maximum tiller numbers per m2 were recorded with P2
spacing under IF-V water regimes. Treatments P2 and
P3 both had higher number of tillers compared with P1,
and both were at par under both IF-R and NF water
regimes, whereas under the IF-V and CF treatments, the
P2 pattern had a higher number of tillers compared
with P3 (Table 3). In P1/IF-V, a 7 % tiller reduction
was observed compared with the contrasting treatment
of P3/CF.
While the number of tillers per hill was the highest in the
P1 spacing treatments compared with P2 and P3 under all
water regimes (data not shown), this did not compensate
for the fewer hills per m2 associated with wider spacing
between the hills. P2 treatments grown under IF-V condition had 15 % more tillers compared with P2 under CF
conditions, indicating a significant gain from intermittent
irrigation during the vegetative stage over continuouslyflooded conditions in terms of better plant growth.

Table 1 Computed F value from analysis of variance (anova) of productive tiller per m2, spikelet per panicle , percentage of ripened grain, single
grain weight (mg), thousand-grain weight (g), grain yield (tons h)1) at 14 % moisture content
Computed F

Source

Productive
tiller per m2

Spikelet per
panicle

Percentage of
ripened grain

Single grain
weight

Thousand-grain
weight

Grain yield

Planting pattern (PP)

Water regimes (WR)
PP WR

607.5**
300.35**
41.05**

175.55**
68.97**
18.06**

99.96**
19.73**
0.87ns

279.5**
64.27**
7.38*

752.92**
96.78**
3.63*

142.64**
132.98**
12.05**

Df = 2, 83, for planting pattern; 3, 83, for water regime, and 6, 83, for planting pattern water regime.
ns, not significant.
*P < 0.05 and **P < 0.001.

Table 2 Computed F value from analysis of variance (anova) of root length density (RLD) (cm cm)3) at flowering and at 20 days after flowering
(20 DAF) at the soil depth of 1520 cm and 3540 cm, root oxidizing activity rate (lg g)1 FW h)1) at flowering and 20 DAF
Computed F
At flowering

At 20 DAF

Source

RLD (1520 cm)

RLD (3540 cm)

ROA

RLD (1520 cm)

RLD (3540 cm)

ROA

Planting pattern (PP)

Water regime (WR)
PP WR

276.02**
211.15**
19.73**

108.67**
344.76**
3.78*

6.68**
203.44**
2.56*

224.48**
175.84**
12.18**

191.57**
132.45**
5**

224.48**
175.84**
12.18**

Df = 2, 83, for planting pattern; 3, 83, for water regime, and 6, 83, for planting pattern water regime).
ns, not significant.
*P < 0.05 and **P < 0.001.

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371

Mishra and Salokhe

Planting density
Single seedlings per
hill (20 20 cm) (P2)

Three seedlings per

hill (20 20 cm) (P3)

Productive tillers per m2

IF-V
353.75 (2.17) cA
IF-R
316.25 (2.75) bC
CF
327.50 (3.12) cB
NF
327.75 (3.04) bB

470.00
368.11
410.00
352.38

(3.85)
(5.31)
(2.04)
(3.06)

aA
aC
aB
aC

427.75
374.50
380.63
366.75

Spikelets per panicle

IF-V
232.71 (1.57) aA
IF-R
216.29 (1.25) aB
CF
220.00 (1.59) aB
NF
209.14 (0.63) aC

217.14
210.14
213.86
205.29

(1.77)
(1.06)
(1.67)
(1.32)

bA
bB
bABC
aC

210.71 (2.54) cA
169.14(4.56) cD
201.29 (1.90) cC
191.14 (1.96) bC

Percentage of ripened grains

IF-V
92.86 (0.38) aA
IF-R
90.11 (0.61) aA
CF
90.60 (0.77) aA
NF
94.34 (0.82) aA

89.54
84.98
84.33
89.57

(0.46)
(0.30)
(1.03)
(0.48)

bA
bC
bC
bA

83.96
78.88
78.84
85.53

(0.73)
(2.32)
(1.78)
(0.66)

cB
cC
cC
cA

Single grain weight (mg)

IF-V
26.16
IF-R
24.75
CF
26.33
NF
26.13

(0.17)
(0.16)
(0.08)
(0.17)

aA
aB
aA
aA

25.04
23.25
24.32
25.36

(0.12)
(0.08)
(0.35)
(0.13)

bA
bB
bB
bA

23.31
22.38
22.45
24.20

(0.07)
(0.14)
(0.16)
(0.16)

cB
cC
cC
cA

Thousand-grain weight (g)

IF-V
31.16
IF-R
28.10
CF
28.44
NF
29.10

(0.23)
(0.24)
(0.12)
(0.19)

aA
aC
aC
aB

29.34
27.07
27.68
27.87

(0.36)
(0.23)
(0.09)
(0.10)

bA
bC
bB
bB

25.19
22.40
23.51
24.49

(0.23)
(0.23)
(0.15)
(0.16)

cA
cD
cC
cC

at 14 % moisture content
4.61 (0.16) cA
6.28 (0.15) aA
3.7 (0.15) bBC
4.53 (0.19) aC
4.05 (0.13) cB
5.85 (0.13) aB
3.76 (0.11) bC
4.43 (0.13) aC

5.18
4.20
4.75
4.41

(0.19)
(0.07)
(0.15)
(0.28)

bA
aD
bB
aC

Water regimes

Single seedlings per

hill (30 30 cm) (P1)

Grain yield (tons ha)1)

IF-V
IF-R
CF
NF

(5.07)
(5.58)
(2.46)
(3.50)

Table 3 Effect of planting pattern (P1,

P2 and P3) and water regimes (IF-V, IF-R, NF,
and CF: see Materials and Methods for
symbol details) on yield contributing parameters
of rice

bA
aC
bB
aC

Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test SAS Institute, 1999).

A greater number of spikelets per panicle were

observed in P1 treatments, on the other hand, followed
by P2 and P3 in IF-V, IF-R and CF water regimes. Spikelet numbers were generally the highest under IF-V water
regime compared with IF-R, CF and NF treatments
(Table 3).
Plants grown with P2 and P3 spacing under IF-V and
NF conditions had more ripened grains compared with
IF-R and CF water regimes. The percentage of increase in
ripened grains in the P1/IF-V combination was 16 %
compared with P3/CF, whereas it was 13 % and 5 %
more than P3/CF in the P2/IF-V and P3/IF-V combinations. There was no effect of water regime on ripened
grain when grown with P1 planting pattern, an interesting
finding.
Single grain weight was significantly affected by the
interaction of water regime planting pattern (Table 3).
The P1 planting pattern, i.e., plants grown with single
372

seedlings and wider spacing, had significantly heavier

grains compared with P2 and P3, and this was observed
under all water regimes, also an interesting finding.
Thousand-grain weight was significantly improved in
the P1 treatment compared with P2 and P3 under all
water regimes, also in the IF-V water regime compared
with the IF-R, CF and NF water regimes. Overall, these
findings revealed that yield-contributing parameters
within the plant were significantly improved by transplanting single seedlings with wider spacing, which was in
agreement with the earlier findings of Baloch et al.
(2002).
Final grain yield was significantly affected by the interaction effects of planting density and water regime
(Table 3). Higher grain yield was recorded in the IF-V
treatment under all planting patterns compared with CF
management, whereas under NF and IF-R, a significant
reduction was observed. Within CF water regime, the
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P2/CF combination had a 23 % yield advantage over the

P3/CF combination, whereas it was 32 % more in the
P2/IF-V combination compared with the P3/CF treatment
(Table 3). This indicated that minimizing intra-hill competition could be one way to improve grain yield even
under continuously flooded conditions. Moreover, transplanting single seedlings per hill had a stronger synergy
effect on grain yield under intermittent irrigation during
the vegetative stage compared with continuously flooded
conditions throughout the crop cycle, as reported earlier
by Lin et al. (2004).
Changes in chlorophyll content
As expected, chlorophyll content of the flag leaves and
third leaves decreased with ripening (Fig. 1). However,
the rate of decrease varied under different planting densities and water regimes. At flowering and at late maturing
stage, chlorophyll content was higher in the P1 treatment
under all water regimes. The difference in chlorophyll

Root growth pattern and root oxidizing activity

The RLD was higher in the top 1520 cm soil below
the hill under all studied planting patterns compared to
P3 (20 x 20) cm

P2 (20 x20) cm

P1 (30 X 30) cm

600

content of the flag leaves and the third leaves was also less
with P1 spacing compared with P2 and P3. At full heading and in middle of the heading stage, there was not
much difference in chlorophyll content with the P2 and
P3 planting patterns. However, at the late ripening stage,
the P2 treatment had higher chlorophyll content compared with P3. This was resulting from the lower rate of
reduction in chlorophyll content over the crop cycle with
P2 treatment compared with P3.
Under different water regimes, the rate of chlorophyll
reduction was higher in the CF water regimes than with
IF-V. This resulted in earlier senescence of plants under
continuously flooded conditions. Moreover, the P1/IF-V
combination had considerably delayed senescence compared with the conventional practice (P3/CF).

IF-V

Chlorophyll content
(mol m2)

500
400
300
200
100

Flag leaf
Third leaf

Flag leaf
Third leaf

Flag leaf
Third leaf

600

IF-R

Chlorophyll content
(mol m2)

500
400
300
200
100

Flag leaf
Third leaf

Flag leaf
Third leaf

Flag leaf
Third leaf

NF

Chlorophyll content
(mol m2)

600
500
400
300
200
100

Flag leaf
Third leaf

Flag leaf
Third leaf

Flag leaf
Third leaf

600

CF

Chlorophyll content
(mol m2)

500
400
300
200
100

Flowering

Flag leaf
Third leaf

7 DAF

Flag leaf
Third leaf

Flag leaf
Third leaf

14 DAF

21 DAF

Flowering

7 DAF

14 DAF

21 DAF

Flowering

7 DAF

14 DAF

21 DAF

Fig. 1 Changes in chlorophyll content of flag leaf and third leaf of rice plant from flowering to 21 DAF (days after flowering) under different
water regimes: intermittent draining for three times (IF-V), intermittent draining for five times (IF-R), continuously flooded (CF) and non-flooded
(NF) and planting patterns. Vertical bar represents the standard error (n = 7). For each replicate, three leaves were used for measurement
(P < 0.05, DMRT test, SAS Institute, 1999).

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Mishra and Salokhe

Table 4 Effect of planting pattern (P1, P2 and P3) and water regimes (IF-V, IF-R, CF and NF) on root length density per hill (cm cm)3) at flowering and 20 days after flowering (DAF) for soil layer of 1520 cm and 3540 cm
Water
regimes

IF-V soil depth

1520

IF-R soil depth

NF soil depth

CF soil depth

3540

1520

3540

1520

3540

1520

3540

RLD at flowering
P1
36.52bA (0.41)
P2
35.44bA (0.23)
P3
29.54bB (0.28)

17.40bA (0.38)
15.42aB (0.54)
12.28aC (0.79)

36.80bA (0.29)
35.85bA (0.21)
27.25cB (0.34)

17.34bA (0.65)
15.43aB (0.65)
12.29aC (0.37)

29.54cA (0.41)
27.76cB (0.54)
27.10cB (0.58)

18.73bA (0.44)
15.73aB (0.48)
13.23aB (0.79)

39.12aA (0.59)
37.62aB (0.47)
32.32aC (0.36)

6.92aA (0.19)
5.32bB (0.18)
4.55bC (0.18)

RLD at 20 DAF
P1
20.13aA (0.44)
P2
18.13aB (0.31)
P3
10.13aC (0.45)

7.86bA (0.06)
7.75aA (0.27)
5.55aB (0.15)

18.99aA (1.27)
17.15aB (0.42)
9.13abC (0.6)

7.67bA (0.23)
7.45aA (0.17)
5.75aB (0.21)

12.64bA (0.16)
10.45bB (0.24)
8.28bC (0.17)

8.54aA (0.20)
7.27aA (0.38)
5.62aB (0.28)

10.37cA (0.20)
8.17cB (0.43)
5.13Cc (0.28

5.44cA (0.28)
4.87bB (0.22)
3.65bC (0.12)

Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test, SAS
Institute, 1999).

3540 cm soil depth. This pattern was similar under

IF-V, IF-R and CF treatments, but not NF water regime.
Under NF, the total RLD was less compared with other
water treatments. At flowering, higher RLD was observed
with continuously flooded water regime (CF) compared
with the IF-V and IF-R water regimes at upper soil depth,
whereas at lower soil depth, it was the least in CF water
regimes (Table 4). At deeper soil depth, the NF treatment
had the largest distribution of roots compared with other
water regimes.
Similarly, planting pattern significantly influenced root
growth pattern. Single seedlings per hill had higher RLD
compared with three seedlings per hill under all water
regimes. This difference was observed at flowering, and
even at 20 DAF.
The data show that in the top layer of soil, P1and P2
treatments had similar root distribution at flowering
under IF-V and IF-R water regimes, whereas with CF
there was higher root distribution in P1 compared with
P2 and P3 treatments. Under the non-flooded water
regime, P2 and P3 treatments had similar root growth
but this was <P1.
At lower soil depth, planting pattern had a significant
effect on root distribution under IF-V, IF-R, and CF
water regimes. P1 had more roots followed by P2 and P3.
Under non-flooded soil conditions, P2 and P3 had similar
root growth.
At 20 DAF, RLD was significantly reduced under the
continuously flooded water regime (CF) compared with
IF-V, IF-R and NF water regimes. This reflects how continuous flooding is a cause of root degeneration by the
late reproductive stage (Kar et al. 1974). In addition, root
degeneration at upper soil depth was more with the P3
treatment compared with P2 and P1, showing that
crowded planting is another reason for root degeneration
by late reproductive stage.
374

At lower soil depth, there was no difference in RLD

with P1 and P2 spacing under IF-V, IF-R and NF water
regimes. But under continuously flooded conditions, RLD
was significantly reduced under P3 compared with P2
and P1. Moreover, the combination of sparse planting
and intermittent irrigation during the vegetative stage
resulted into significantly slower degeneration of roots
under P1/IF-V practice compared with conventional practice, P3/CF.
Physiological activity of roots as represented by the rate
of root oxidizing activity, was significantly affected by the
interaction of water regimes and planting pattern. Root
oxidizing activity rate was similar and higher in the IF-V,
IF-R and CF treatments at the flowering stage under P1,
whereas under NF treatment, less activity was observed
compared with other water regimes without having any
effects of planting pattern. At the late reproductive stage,
a significant reduction in root oxidizing activity rate was
observed in plant grown with CF water regimes compared
with IF-V, and in P3 planting density compared with P1
and P2 (Table 5). At late reproductive stage, the highest
root oxidizing activity rate was observed under P1/IF-V
combination, whereas it was least under conventional
practice, P3/CF.
Discussion
Among the characteristics that contribute to increased dry
weight and higher grain yield at the later growth stage,
delayed leaf senescence is considered to be very important. This delay, associated with increased production of
dry matter and grain, has been observed previously in rice
(Jiang et al. 1988). San-oh et al. (2006) reported that the
improved photosynthetic characteristics of rice leaves was
the main reason for greater dry matter production in
plants established with a single seedling per hill compared
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Effects of Some SRI Practices on Rice Growth

Table 5 Root oxidizing activity (lg g)1 FW; h)1)

under different water regimes (IF-V, IF-R, CF
and CF) and planting pattern (P1, P2 and
P3)

IF-V

IF-R

NF

CF

Root oxidizing activity at flowering

P1
61.47 (0.54) aA
60.87 (0.55) aA
P2
60.22 (0.80) aB
59.53 (0.53) aB
P3
59.62 (0.51) aB
58.03 (0.83) bB

52.35 (0.47) bA
53.35 (0.61) bA
52.31 (0.92) cAA

60.31 (0.45) aA
59.35 (0.42) aA
59.86 (0.45) aA

Root oxidizing activity at 20 days after flowering

P1
43.28 (0.59) aA
42.01 (0.70) aA
P2
33.1 (0.65) aB
32.23 (0.80) aB
P3
22.31 (0.62) bC
23.35 (0.62) bC

40.30 (0.33) bA
32.54 (0.20) aB
27.70 (0.32) aC

38.32 (0.24) cA
28.62 (0.25) bB
19.23 (0.61) cC

Means with similar small letters within the row and capital letters within the column are not significantly different. (P < 0.05, DMRT test, SAS Institute, 1999).

with three seedlings per hill. Rates of photosynthesis at

the crucial stages in grain formation and filling depend
on the chlorophyll content of the leaves, as well as on
root activity and functioning discussed already.
In this study, the chlorophyll contents of the flag leaf
were similar at the heading stage under single seedling
per hill transplanting and three seedlings per hill. Therefore, it appears that the rate of photosynthesis would be
similar in both treatments at this stage. But, at the later
growth stage, the chlorophyll content of the flag leaf and
the lower leaf was significantly higher with single seedling
per hill transplanting compared with three seedlings per
hill. This reflected a decreased rate of leaf senescence in
the hills with single seedlings. The slower leaf senescence
rate in single seedlings per hill contributed to a higher
photosynthesis rate at the late reproductive stage compared with plants where three seedlings per hill were
transplanted. A similar finding was reported by San-oh
et al. (2006).
In our experiments, singles seedling per hill transplanted with wider spacing had further delayed senescence
compared with the single seedlings per hill transplanted
with narrower spacing. Moreover, leaf senescence was
affected not only by planting density, but also by water
regime. Plants grown under IF-V had delayed senescence
of their flag leaf and lower leaves at the late reproductive
stage, which resulted in higher dry matter production
compared with plants grown under continuously flooded
conditions.
The results revealed significant interaction effects
between planting density and water regime for most of
the yield-contributing parameters such as number of productive tillers, spikelets per panicle, percentage of ripened
grain, single grain weight, and thousand-grain weight.
Overall, these yield-contributing characteristics were generally better under IF-V treatments compared with CF.
One possible reason for this, according to our findings,
could be the better root distribution and the higher root
activity at later growth stage with IF-V compared with
CF.
2010 Blackwell Verlag GmbH, 196 (2010) 368378

The better root distribution and the higher root activity

at later growth stage might be related to delayed senescence of the lower leaves. This inference is based on a
previous report that the lower leaves are most active in
supplying photosynthates and oxygen to the roots
(Tanaka 1958). Withered lowered leaves, especially at later
growth stages, lose their ability to take up and pass on
oxygen and photosynthates to the roots (Arikado 1975).
Consequently, it was considered essential to keep the
lower leaves alive as long as possible, so as to provide
roots with their requisites for metabolism.
In this study, it appeared that reduced planting density
and reduced intra-hill competition, such as with P1 and
P2 spacing, helped plant roots to receive more photosynthates and oxygen through the lower leaves at the later
growth stage, as root growth and root activity rate were
higher under both planting patterns. This also suggests
that there would be less shading in P1 and P2 compared
with P3 treatments. Recently, Thakur et al. (2009b)
reported similarly that widely-spaced hills had less shading effects compared with close-spaced hills.
Further, the data also showed that intermittent irrigation helped in maintaining higher root activity rate at the
later growth stage compared with continuously flooded
condition. These could be a reason for the finding of
Lin et al. (2004) that there was better ontogenesis when
the seedlings were transplanted at low densities under
intermittent irrigation compared with conventional water
management.
However, in our study, we saw that intermittent irrigation followed into the reproductive stage significantly
reduced panicle numbers, spikelets per panicle, single
grain weight and thousand-grain weight compared with
CF and IF-V. The reason for this could be that repeated
drainage in later growth stages, especially the fourth and
fifth drainage cycles in the IV-R treatment, coincided with
the flowering and grain-filling stages when plant requirements for water and other soil nutrients would peak.
Some reduction in these yield-influencing parameters
might be attributable to water deficit-induced pollen
375

Mishra and Salokhe

abortion (Nguyen and Sutton 2009) or leaching losses of

nutrients that would result in an increased rate of plant
senescence subsequently.
The previous research (Steponkus et al. 1980) and our
review on literature relevant to SRI water management
(Mishra et al. 2006) have suggested that the plant makes
osmotic adjustments at the vegetative stage when exposed
to intermittent irrigation. These plants can better tolerate
moisture stress during their later growth stages. However,
our findings, working with heavy clay acid sulphate soil,
did not support this suggestion. It seems that although
plants can tolerate moisture stress even at their peak
growth stage, this is achieved at some cost of grain yield,
especially when the plant is exposed to water stress in the
early grain-filling stage. Soil characteristics need to be
taken into account when assessing these factors and their
interaction.
Significant yield reduction in the non-flooded water
regime trials revealed that in the heavy clay acid
sulphate soil of our experiment farm, continuously
non-flooded water regime was associated with low pH
of the soil that limited nutrient availability to the plant.
Moreover, nitrogen was mainly available in nitrate form
(as a result of aerobic soil condition) and that would
reduce the uptake rate in the plant, and consequently
affect the rate of shoot and root growth. Water management is more critical at the reproductive stage than at
vegetative stage when grain yield is a more dominant
factor than plant growth. What should be the interval
and frequency of intermittent irrigation is a subject warranting further research, assessing farmers preferences
and location-specific adaptations based on biophysical
and socio-economic realities.
This study also indicated that root length densities
which represent the development of root laterals were
higher in the P1 and P2 treatments compared with P3 at
upper soil depth under all water regimes. Noticeably,
reduced intra-hill competition and wider spacing
favoured the development of more root laterals. More laterals helped the roots to achieve a greater surface absorption area with higher cation exchange capacity, favouring
higher nutrient absorption from the soil.
Under continuously flooded water regime, the development of laterals was restricted to the upper soil depth. It
appeared that the development of laterals might be influenced by oxygen concentration in the rhizosphere along
with nutrient availability. Root oxidizing activity was
similar under CF, IF-V and IF-R treatments at flowering
stage, i.e., the rates of oxygen release per gram of root
were similar under these treatments. However, P1 and P2
planting pattern generally had higher RLD compared with
P3. Therefore, no direct correlation was seen between
root branching and the root oxidizing activity rate, i.e.,
376

the root respiration rate which is considered as one of the

energy-consuming processes in rice plants.
However, root branching may affect oxygen distribution around the roots and in the rhizosphere (Kirk 2003).
If root oxidizing activity is considered as a unit, it could
be said that with the same oxygen loss, a root system with
greater coverage of laterals, such as in the P1 and P2
treatments, has a larger oxygenated absorbing surface area
compared with the P3 treatment.
At 20 DAF, RLD was drastically reduced under a continuously flooded water regime, and particularly with P3
spacing. Moreover, the rate of senescence of the flag leaf
and lower leaf was faster under a continuously flooded
water regime. This reflected that root senescence is associated with lower leaf senescence. Similar findings were
reported in Mishra and Salokhe (2008b). This study
reconfirmed the causal relationship between root and
shoot growth in rice as this affects grain yield even at late
maturity stage.
Overall, the results indicated the benefit of the SRI
recommendations for plant spacing and water management over those associated with conventional management practices. There was consistently better performance
of transplanting single seedlings per hill compared with
three to four seedlings, and benefit from intermittent
irrigation during the vegetative growth stage compared
with continuously flooding the soil in terms of improved
plant characteristics.
In terms of yield per hectare, we observed that transplanting single seedlings with wide spacing of 30 30 cm
did not perform significantly better than single seedlings
per hill with narrower spacing (still wider than most conventional practice), or than three to four seedlings per hill
transplanted with narrow spacing and grown under continuously flooded water regime. This was mainly attributable to the significantly lower tiller number per m2 in the
P1 treatments. Plants grown with P1/IF-V treatment combination had a prolonged vegetative stage and delayed
senescence with improved yield-contributing parameters
(within the plant) compared with conventional practice.
This compensated for the lower number of tillers per m2
so that similar yields resulted from P1/IF-V and P3/CF
combinations in these trials.
Spacing is a factor always to be optimized, depending
on soil and varietal characteristics such as maturity period, tillering ability, photoperiod sensitivity, etc. This
means that location-specific and varietal-specific agronomic research further evaluating SRI practices, respectively and in combination, should be undertaken for
better yield (and profitability). Such adaptations and
adjustments are considered part of the principles and
philosophy of SRI management (Stoop et al. 2002). In as
much as treatments combining SRI methods such as
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Effects of Some SRI Practices on Rice Growth

P1/IF-V or P2/IF-V required less seed compared with

conventional practice, this modification in practice could
provide opportunity for farmers to achieve similar or better yield with less seed inputs through adaptive on-farm
research.
The findings showed that transplanting single seedlings
per hill even under continuous flooding could give significantly better results than current usual management
practices, i.e., transplanting three to four seedlings per
hill. The benefit of transplanting single seedlings per hill,
which is relatively easier to adopt than water management
changes under some field conditions, was clear. Most of
the yield-contributing parameters along with root growth
and physiological activity were significantly improved by
transplanting single seedlings per hill. That could be one
reason for wider acceptability of this practice among
Asian rice farmers who have been reporting significant
yield achievements under SRI practice. Already 100 years
ago, farmers in Tamil Nadu, India, were promoting the
use of single seedlings, and the British colonial government recommended it; although for unknown reasons
this practice was apparently lost in the 1930s (Thiyagarajan and Gujja 2009).
The water saving possible with reduced irrigation will
probably have more significance over time. The competing demands for water are compelling farmers to find
ways to produce more rice with less water. Given the
incentives to modify irrigation practices, we suggest that
SRI principles might be helpful to develop location-specific management practices to optimize water and other
input use without compromising grain yield. Therefore,
the SRI debate should be taken as an opportunity to
understand, appreciate and address the concern of millions of rice farmers who need to cope with water scarcity
along with other sources of vulnerability.
The findings of this study suggest that there are large
opportunities to improve rice management practices
using SRI principles for better adaptability and profitability according to soil and varietal characteristics to obtain
higher grain yield with less seed, and also less water. Possibly there could be other reductions in the cost of inputs,
but the effects of changing fertilization and crop protection strategies, also recommended with SRI, were not
evaluated here.
Better utilization of SRI or any other crop management practices in a given bio-physical and socio-economic conditions requires better crop management
knowledge and skill. A collaborative research approach
that focuses on developing farmers scientific knowledge
and their productionprotection management skills as
well as on scientists practical understanding of scientific-social issues and their application of scientific
knowledge in locally-specific field situations should be
2010 Blackwell Verlag GmbH, 196 (2010) 368378

promoted by agricultural research institutions and development organizations.

Conclusion
The combination of transplanting single seedlings per hill
and following intermittent irrigation during the vegetative
growth stage improved RLD and root activity rate as well
as shoot growth and delayed senescence of plants, leading
to higher grain yield.
These two crop management practices are key elements
of SRI methodology, and their synergistic effects on root
and shoot growth can account for some of the higher
yield reported when SRI practices are utilized. This study
was conducted using 16-day-old seedlings. Further studies
could be conducted using much younger seedlings, i.e.,
10 days (two-three leaf stage), which is commonly recommended under SRI management practices. In addition,
potential interaction effects of all the recommended SRI
practices (young seedling age, reduced planting density,
less water application, increased use of compost, and soilaerating weeding) which were beyond the scope of this
study should be explored in detail to learn their effects
on root and shoot growth and on grain yield.
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