Journal of Agricultural Science, Page 1 of 15.

f 2006 Cambridge University Press 1

doi:10.1017/S0021859605005745 Printed in the United Kingdom

Variation in yield, N uptake and N use efficiency of

medium and late duration rice varieties

D I L L I P K U M A R S W A I N 1*, B U R L A C H A N D R A B H A S K A R 2, P R A M I L A K R I S H N A N 2,
K U R K U R I S R I N I V A S A R A O 2, S A N G R A M K E S H A R I N A Y A K 2
2
AND RABINDRA NATH DASH
1
Environment and Sustainable Development, United Nations University, 53–70 Jingumae, 5-Chome, Shibuya-Ku,
Tokyo-1508925, Japan
2
Central Rice Research Institute, Cuttack 753 006, India
(Revised MS received 18 October 2005)

SUMMARY
Field experiments were conducted at the village Kasiadihi, Dhenkanal district, Orissa, India during
wet seasons 2001, 2002 and 2003 under non water-stressed conditions (0–25 cm standing water) to
assess variability in N uptake and utilization by medium and late duration rice varieties. The N rates
were 0, 40, 80 and 120 kg N/ha applied as urea in four equal splits at transplanting, active tiller
initiation, panicle initiation and flowering stages. The grain yield response was up to 80 kg N/ha. The
optimum grain yield attainable by the efficient medium duration varieties was 4.5 t/ha. The N efficient
late duration varieties produced optimum grain yield of 5.8 t/ha. The relationship for total dry matter
and grain yield production between N fertilized (40, 80 and 120 kg N/ha) and non-fertilized
treatments were all significant, suggesting cultivar selection under optimum N fertilized conditions.
The difference in optimum yield of the medium and late duration varieties was due to the differences
in the amount of N uptake and its use efficiency by the plant for grain production. There was
a curvilinear relationship between grain yield and N use efficiency for grain production. The
relationship between N use efficiency for grain production and N contents of leaf, stem and grain at
maturity was quadratic. The optimum plant N use efficiency of medium duration varieties was 49 kg
grain/kg N uptake, achieved with leaf, stem and grain N contents of 10, 8 and 14 g/kg, respectively, at
maturity. For late duration varieties, the optimum plant N use efficiency was 68 kg grain/kg N uptake
and it was maintained with leaf and stem N content of 4.0 g/kg each and grain N content of 9.0 g/kg
at maturity. The N content in plant organs could be the selection guide used to obtain efficient rice
varieties.

INTRODUCTION 1979). Cereals also release N from plant tissues,

predominantly as ammonia following anthesis
Nitrogen is one of the key nutrients that limit
(Harper et al. 1987 ; Francis et al. 1993).
crop growth of cereals in many production systems.
Overuse of fertilizer and the occurrence of other
Nitrogen deficiency is one of the most serious
macronutrient deficiencies can give rise to lower N
nutritional disorders in lowland rice-producing areas
use efficiency (Von Uexkull 1993). Application of
of the world. The proportion of the applied N
sulphur and N prevents nitrate leaching and enhances
accounted for by the above-ground N accumulation
N use efficiency in grassland soils (Brown et al. 2000).
in rice was 0.30–0.40 (Cassman et al. 1993). Ammonia
By including cowpea in a rice–wheat rotation in the
volatilization, nitrification-denitrification, surface run
productive Indo-Gangetic plains of India, it might
off, and leaching account for major N loss from
be possible to reduce the optimum fertilizer N dose
the soil, which is associated with low N recovery
due to increased N availability and greater N use
efficiency in lowland rice (Craswell & Vlek
efficiency (Yadav et al. 2003). However, genetic
selection to improve N use efficiency has so far
* To whom all correspondence should be addressed. been unsuccessful. Nevertheless, there are significant
E-mail: swain@hq.unu.edu differences in N use efficiency between rice varieties.
2 D. K. S W A I N E T A L.
Sta. Cruz & Wada (1994) evaluated several varieties
and concluded that growth duration, native soil
fertility and cultural practices affect the pattern of N
absorption, which in turn alters the amount of N in
plants leading to profound effects on N use efficiency.
Rice varieties may respond differently to N appli-
cation (Singh et al. 1998). Cultivars selected under
high N fertilizer application may not be suitable for
soils with low N status. Even after the application of
high rates of fertilizer N to rice, expected yield levels
might not be obtained. If plant N status can be
increased without lodging or increasing the incidence
of disease, a significant increase in yield requires
increased sink capacity, maintenance of high leaf N
content and a longer grain filling duration (Aggarwal
et al. 1997).
Rice varieties differ in their ability to extract soil
and fertilizer N and in its distribution to different
plant organs. Understanding N uptake and assimi-
lation is necessary in any attempt to optimize the
efficiency of absorbed N for grain production.
Senanayake et al. (1996) showed that vigorous bio-
mass accumulation could lead to dilution of plant
nitrogen content up to the panicle initiation stage,
which could lead to inefficient use of N for spikelet
formation. It is important to increase the efficiency of
soil and fertilizer N by using nutrient efficient var-
ieties. It is hypothesized that the N use efficiency of
the rice plant can be optimized by critical leaf, stem
and grain N content of rice varieties, which improves
the efficiency of grain production. The first objective
of the present investigation was to assess variability in
grain yield, N uptake and N use efficiency for grain
production of medium and late duration rice
varieties. The second objective was to ascertain the
critical N content in rice plant parts for optimizing
the efficiency of absorbed N for grain production. The
results could contribute to an improvement in N
fertilizer management, and might also prove useful
for the selection and breeding of efficient rice
varieties.
MATERIALS AND METHODS
Field experiment
Field experiments were conducted in a field at
Kasiadihi village, Denkanal district, Orissa, India in
2001, 2002 and 2003, to study variation in N uptake
and utilization by medium and late duration rice
varieties. The site is a rain-fed area spread over 6 km2
in the 20x40kN latitude and 85x38kE longitude at
300 m asl. Medium and late duration rice varieties of
115–150 days maturity are usually grown at this site,
without water stress. The climate is hot and humid.
The weekly mean maximum temperature ranges from
26 xC in December to 41 xC in April. The weekly
mean minimum temperature ranges from 12 xC in
December to 26 xC in April. The soil is acidic, pH 5.8,
and the texture is sandy clay loam with 15 % clay,
72 % sand and 13 % silt by volume. The bulk density
of the soil is 1.45 g/ml. The organic C and total N
contents are 4.7 g/kg and 0.7 g/kg, respectively. There
were no deficiency or toxicity symptoms of any
nutrients other than N. The average annual rainfall of
the district is 1421 mm. The proportion of the total
rainfall received during peak monsoon months (June
to September) is 0.81. The average number of rainy
days is 73 in a year. The relative humidity during the
wet season is always above 90 %.
Two experiments were conducted in the wet season
(June to December) of 2001 under non water-stressed
conditions (0–25 cm standing water). They were side
by side in two separate blocks for medium duration
(115–130 days maturity) and late duration (135–150
days maturity) rice varieties. Before the start of the
main experiments in the wet season, a uniformity trial
was conducted with rice variety IR 36 with N applied
at 20 kg N/ha in the previous dry seasons (January to
May) to reduce soil fertility heterogeneity throughout
the field.
Altogether, 12 popular released rice varieties
from different states of India were included in the
investigation, of which five varieties were of medium
duration and seven of late duration. The character-
istics of these rice varieties are given in Table 1. The
experiments were laid out in a split plot design with
two replications, where varieties were in the main
plot and N rates were in subplots. The subplots were
6r4 m. The N rates were 0, 40, 80 and 120 kg N/ha,
applied as urea in four equal splits at Zadoks
growth stage GS14 (transplanting), GS22 (active
tiller initiation), GS37 (panicle initiation) and GS65
(flowering stage) of the crop (Zadoks et al. 1974).
Phosphorus and potassium were applied uniformly at
the rate of 60 kg P2O5/ha and 60 kg K2O/ha to the
puddled soil at transplanting. The plots were trans-
planted with 30-day-old rice seedlings in the third
week of July each year with two seedlings/hill. The
inter-plant spacing was 15 cm and inter-row spacing
was 20 cm. Plots remained flooded with rainwater
from 0–25 cm throughout the experiment. The sub-
plot bund height was 40 cm whereas the main bund
height was 50 cm so that rainwater did not overflow
to adjacent plots. Polythene lining 30 cm deep was
provided inside the subplots along the bund to
prevent the flow of water and dissolved nutrients
through seepage. The same experimental site was
reused in successive years (2002 and 2003). All the
subplots in each block were broken to make into one
plot before the uniformity trials in the dry seasons of
2002 and 2003. After the uniformity trials, fresh
experimental layouts were made in the wet seasons
of 2002 and 2003 in each block. Even though the
experiments were repeated in the same design in two
wet seasons of 2002 and 2003, the main plot and
 Table 1. Differences in plant characters of the rice varieties used

Morphological characters

Maturity Leaf
duration Height Tillering Lodging* Systems Season length Panicle
Varieties (days) (cm) ability (0–1 scale) Grain type bred for Systems used in of use (cm) Colour type

Medium duration
IR 36 115–120 87¡2.2 High 0.05 Long slender Irrigated Irrigated/Shallow Dry/Wet 46¡2.3 Green Open
low land
Mahamaya 125–130 112¡2.1 Medium 0.05 Long bold Irrigated Irrigated/Shallow Dry/Wet 46¡1.8 Green Open

N use efficiency of rice

low land
Kranti 115–120 101¡2.0 Medium 0.05 Short bold Irrigated Irrigated/Shallow Dry/Wet 33¡2.1 Green Open
low land
Lalat 120–125 111¡2.2 High 0.05 Long slender Irrigated Irrigated/Shallow Dry/Wet 45¡2.4 Green Open
low land
Khitish 115–120 85¡1.7 High 0.05 Long slender Irrigated Irrigated/Shallow Dry/Wet 38¡1.7 Green Open
low land
Late duration
Mahsuri 140–145 140¡2.5 High 0.20 Medium slender Irrigated Shallow low land Wet 43¡1.9 Green Open
Madhuri 135–140 108¡2.1 Medium 0.10 Long slender Irrigated Shallow low land Wet 43¡2.2 Green Open
Savitri 145–150 111¡2.8 High 0.05 Short bold Shallow Shallow low land Wet 44¡2.7 Dark green Open
low land
Rajshree 140–145 142¡2.1 Medium 0.15 Medium slender Shallow Shallow low land Wet 45¡2.3 Green Open
low land
Swarna 135–140 101¡2.0 High 0.05 Medium slender Irrigated Shallow low land Wet 43¡2.0 Dark green Open
Sashi 135–140 131¡2.7 High 0.20 Long slender Shallow Shallow low land Wet 47¡2.4 Green Open
low land
Ranjit 145–150 122¡2.4 High 0.10 Medium bold Shallow Shallow low land Wet 45¡2.1 Green Open
low land

* 1=completely lodged.

3
4 D. K. S W A I N E T A L.
subplot treatments were re-randomized each year.
In total, there were six experiments in 3 years.
Plant sampling and sample analysis
Above-ground plant samples were collected at
physiological maturity. For this purpose, 10 hills of
one side of the plot and another 10 hills of the
opposite side of the same plot were sampled after
discarding the border two rows in each side of the
plot (Thiyagarajan et al. 1995). These plant samples
were uprooted; root portions were rejected and the
remainder separated into leaves, stems and grains. All
these plant parts were oven-dried at 65 xC to constant
weight. The sum of the weights of these plant organs
was taken as the total aerial dry matter production
(DM). The grain yield and harvest index (HI) were
determined at maturity from the harvest of 10 m2 area
(Yoshida et al. 1976). The grain weight was adjusted
to 14 % moisture content. Suitable samples of leaves,
stems and grains were powdered and digested with
salt mixture and concentrated sulphuric acid in the
electronic micro-block digestion system. The digested
materials were analysed for N content by the micro-
Kjeldahl distillation method (Yoshida et al. 1976) in
the electronic automatic distillation system. From
the N contents of the plant parts, the N uptake by
plant organs was computed. The total N uptake by
rice was the sum of the N uptake by stems, leaves and
grains.
Derived parameters
The following variety specific parameters were com-
puted (Singh et al. 1998; Inthapanya et al. 2000 b).
Nitrogen harvest index (NHI)=grain N uptake/total
N uptake ; Physiological N use efficiency (PE)=DM/
total N uptake; Nitrogen use efficiency for grain
production (NUE)=grain yield/total N uptake;
Apparent recovery of applied N (AR)=(N uptake in
treated plot – N uptake in control plot)/amount of N
applied ; Agronomic N use efficiency (ANUE)=
(grain yield in N treated plot – grain yield in control
plot)/amount of N applied
The above parameters are related as follows.
Grain yield=Total N uptakerPErHI
=Total N uptakerNUE
PE=1000={(X) N content in leaf
+(Y) N content in stem
+(HI) N content in grain}
NUE=1000(HI)={(X) N content in leaf
+(Y) N content in stem
+(HI) N content in grain}
where X+Y+HI=1 ; X is the leaf fraction ; Y is the
stem fraction.
Statistical analysis
A combined analysis of variance for 3 years’ data
with two replications was carried out to evaluate the
contribution of year, variety, N rates, and varietyrN
rates interaction for late and medium duration
varieties (Gomez & Gomez 1984). From the 3 years’
pooled data with two replications, analyses of vari-
ance was carried out on grain yield, DM, N content,
N uptake and N use efficiency parameters, HI and
NHI to evaluate the effect of varieties, N rates and
their interactions. The interaction effects were
presented wherever they were significant. Linear
correlations were used to examine the relationship
between non-fertilized and N fertilized treatments
for grain yield and biomass production. Linear
correlations between grain yield and N uptake,
ANUE, AR, NUE, HI and NHI were carried out
separately for medium and late duration varieties.
The relationship between NUE and N content in leaf,
stem and grain was examined to find out optimum N
contents for grain production. Path coefficient
analysis was used to determine the direct and indirect
contributions of different components to NUE.
Multiple regression analysis was used to estimate
the relationship between the NUE and the
components. For this purpose, the values of all 3
years and all treatments with two replications were
taken separately for medium and late duration
varieties.
RESULTS
Genotype variability in yield parameters
In the medium duration group, the effect of variety
was significant (P<0.01) for variation in DM, grain
yield, total N uptake, N content and uptake by stem,
and NUE. The effect of N rates was significant
(P<0.01) for differences in DM, grain yield, HI, N
content, N uptake and NUE. The variety and N
interaction was significant for grain yield and total
N uptake. The rice varieties of the late duration
group showed significant differences (P<0.01) for
grain yield, N content, AR, ANUE, NUE, HI and
NHI. The effect of N rate was significant (P<0.01)
for DM, grain yield, N uptake and PE. The variety
and N interactions were significant for DM, grain
yield, total N uptake and NUE. The variations in
grain yield, N content, NUE, ANUE and NHI were
attributable more to variety than to N rate. The
seasonal variation resulted in significant changes in
DM and HI of medium duration varieties but such
effect did not persist in late duration varieties.
However, variation in the cropping year was not
 N use efficiency of rice 5

Table 2. Grain yield and total dry matter production at maturity of medium and late duration rice varieties at
0 (N0), 40 (N40), 80 (N80) or 120 (N120) kg N/ha

Grain yield (t/ha) Total dry matter (t/ha)

Varieties N0 N40 N80 N120 N0 N40 N80 N120

Medium duration
IR-36 3.3 4.1 4.5 4.0 7.0 8 .0 9.4 9.4
Mahamaya 3.8 4.3 5.0 4.9 7.7 9 .3 10.2 10.6
Kranti 3.5 3.8 4.6 4.3 7.3 8 .4 9.1 8.9
Lalat 3.6 4.1 4.4 4.9 8.5 9 .3 10.4 11.0
Khitish 3.3 3.6 4.2 3.4 6.4 7 .2 8.5 7.8
S.E.M.
Variety (V) (D.F.=12) 0.10 0.22
Nitrogen (N) (D.F.=45) 0.06 0.13
VrN (D.F.=45) 0.13 0.30
Late duration
Mahsuri 2.8 3.5 3.8 3.4 9.4 10.3 11.1 11.9
Madhuri 1.8 2.0 2.6 2.3 7.9 8 .6 9.5 10.1
Savitri 4.6 5.2 6.0 6.9 10.4 12.7 14.4 15.4
Rajshree 3.1 3.7 4.2 4.3 8.5 10.1 11.3 11.7
Swarna 3.5 4.5 5.4 5.9 8.1 9.768 12.1 14.0
Sashi 3.1 3.6 4.1 4.2 9.1 10.9 12.2 12.8
Ranjit 4.8 5.6 6.2 6.3 10.2 11.3 13.2 14.3
S.E.M.
Variety (V) (D.F.=18) 0.12 0.35
Nitrogen (N) (D.F.=63) 0.06 0.12
VrN (D.F.=63) 0.15 0.32

significant for N content, N uptake and NUE of both
medium and late duration varieties.
Dry matter production and grain yield
The effect of variety and nitrogen interaction for
DM and grain yield production is presented in
Table 2. Under unfertilized conditions, the DM and
grain yield production of the medium duration
varieties was 6.4–8.5 t/ha and 3.3–3.8 t/ha, respect-
ively. For late duration varieties, the DM production
was higher in comparison with medium duration
varieties under non-fertilized conditions and was
7.9–10.4 t/ha, but the grain yield was 1.8–4.8 t/ha,
respectively. There was a response to N up to
80 kg N/ha.
A comparison of the response of rice varieties
to fertilizer N for grain yield and DM production
indicated considerable differences between the two
maturity duration groups. At 40 kg N/ha application,
the N response of late duration varieties for grain yield
and DM production was 32 and 35 % greater than
medium duration varieties (12 kg grain and 27 kg
DM/kg fertilizer N applied), respectively. Increasing
N application from 40 to 80 kg/ha increased the N
response of medium duration varieties to 15 kg
grain/kg fertilizer N applied and this was comparable
to that of late duration varieties. In a similar N
application range, the N response of late duration
varieties for DM production was 30 % greater than
that of medium duration varieties. Late duration
varieties responded poorly, and medium duration
varieties showed no positive response, to an increase
in applied N from 80 to 120 kg/ha for both grain
yield and DM production. The resultant effect was
a decline in grain yield of medium duration
varieties and marginal increases in grain yield of
late duration varieties with N applications above
80 kg N/ha.
The grain yields of the medium duration varieties
were 4.2–5.0 t/ha at 80 kg N/ha. Of all the medium
duration varieties, the variety Khitish produced the
lowest DM and grain yield at all N rates. The rice
variety Mahamaya produced the highest grain
yield, whereas Lalat produced the highest DM at all
N rates. The late duration varieties Savitri and
Ranjit produced higher grain yields (4.6 and 4.8 t/ha,
respectively) than other varieties under non-fertilized
conditions. The varieties Swarna, Sashi and Rajshree
produced grain yields of 3.1–3.5 t/ha. The grain yield
production of the variety Mahsuri and Madhuri
was very low (1.8–2.8 t/ha) in the similar situation.
Swarna, Savitri and Ranjit responded better to
increased N application rates than other varieties.
Their grain yields and DM were 5.4–6.2 t/ha and
12.1–14.4 t/ha, respectively at 80 kg N/ha.
6 D. K. S W A I N E T A L.

Table 3. Nitrogen content and N uptake by leaf, stem and grain of medium and late duration rice varieties at
maturity as affected by different N applications

Leaf Stem Grain

N content N uptake N content N uptake N content N uptake

Treatments (g/kg) (kg/ha) (g/kg) (kg/ha) (g/kg) (kg/ha)

Medium duration
Varieties
IR-36 6.4 6 .4 5.2 15.0 11.2 44.7
Mahamaya 7.8 7 .8 6.0 19.5 11.0 49.8
Kranti 7.2 6 .9 5.6 15.4 11.9 48.5
Lalat 7.0 7 .7 5.5 19.6 11.6 49.8
Khitish 6.9 5 .4 6.2 13.9 11.2 40.3
N rates (kg/ha)
0 6.1 4 .7 4.8 11.4 9.7 33.8
40 6.5 5 .8 5.2 14.6 10.6 41.7
80 7.5 8 .0 6.0 18.8 12.1 55.1
120 8.2 9 .0 6.8 21.9 13.0 56.0
S.E.M.
Variety (D.F.=12) 0.70 0.68 0.17 0.65 0.25 1.72
Nitrogen (D.F.=45) 0.20 0.20 0.14 0.50 0.18 1.01
Late duration
Varieties
Mahsuri 7.7 12.3 5.8 28.4 12.6 43.0
Madhuri 8.4 11.8 6.9 26.9 15.3 33.4
Savitri 5.5 7 .8 4.2 23.7 11.2 63.8
Rajshree 6.8 9 .9 5.9 26.3 13.4 51.4
Swarna 6.0 7 .2 5.5 23.5 13.4 65.5
Sashi 7.5 10.4 5.9 28.6 13.2 50.1
Ranjit 5.8 8 .3 5.0 25.9 10.6 61.3
N rates (kg/ha)
0 5.9 6 .7 4.8 18.6 10.8 35.1
40 6.2 8 .0 5.2 23.4 12.7 49.3
80 7.1 10.8 5.8 27.8 13.7 61.7
120 8.1 13.1 6.7 35.1 14.1 64.4
S.E.M.
Variety (D.F.=18) 0.20 0.33 0.20 1.04 0.50 1.88
Nitrogen (D.F.=63) 0.20 0.28 0.10 0.47 0.20 1.14

N content and uptake by plant organs
The leaf, stem and grain N content and uptake by the
medium and late duration varieties increased with
increasing N rate (Table 3). For all varieties in the
medium and late groups, the stem N content was the
lowest, followed by leaf and grain. The N uptake by
leaf was less than that of stem and grain.
The mean leaf, stem and grain N content of
medium duration varieties across the N rates
was 6.4–7.8 g/kg, 5.2–6.2 g/kg, and 11.0–11.9 g/kg,
respectively. The N uptake of leaf, stem and grain was
lowest in Khitish and highest in Mahamaya and
Lalat. The variations in the N content and N uptake
by plant organs were very large in late duration
varieties. The mean N contents in leaf, stem and grain
were 5.5–8.4 g/kg, 4.2–6.9 g/kg and 10.6–15.3 g/kg,
respectively. Leaf, stem and grain N contents were
relatively higher in the varieties Madhuri, Mashuri,
Rajshree and Sashi than in Swarna, Savitri and
Ranjit. Similar trends were observed in these varieties
for leaf and stem N uptake, but the opposite trend
was found for grain N uptake.
Total N uptake and plant N use efficiency for
grain production
The total N uptake by medium or late duration
rice varieties increased with the increasing N rates,
whereas the NUE decreased (Table 4). At each N
rate, the N uptake by medium duration varieties was
lower and NUE was higher as compared with late
duration varieties.
Of the medium duration varieties, Khitish had the
lowest N uptake. Under non-fertilized conditions,
the N uptake by medium duration rice varieties was
 N use efficiency of rice 7

Table 4. Total N uptake at maturity and N use efficiency for grain production of medium and late duration rice
varieties at 0 (N0), 40 (N40), 80 (N80) or 120 (N120) kg N/ha

Total N uptake (kg/ha) N use efficiency (kg grain/kg N uptake)

Varieties N0 N40 N80 N120 N0 N40 N80 N120

Medium duration
IR-36 45 61 75 84 73 68 60 47
Mahamaya 53 67 91 98 71 64 56 51
Kranti 51 62 85 85 68 61 54 50
Lalat 54 68 85 101 66 60 52 48
Khitish 45 53 74 66 72 67 57 51
S.E.M.
Variety (V) (D.F.=12) 2.0 1.3
Nitrogen (N) (D.F.=45) 1.3 1.1
VrN (D.F.=45) 2.8 2.4
Late duration
Mahsuri 61 77 96 102 45 44 40 34
Madhuri 53 65 83 95 34 30 31 24
Savitri 62 90 110 113 74 58 55 61
Rajshree 62 81 98 114 49 45 43 37
Swarna 61 82 111 122 58 54 49 49
Sashi 61 84 97 115 51 43 42 36
Ranjit 63 85 106 129 76 66 58 49
S.E.M.
Variety (V) (D.F.=18) 2.4 1.7
Nitrogen (N) (D.F.=63) 1.3 0.7
VrN (D.F.=63) 3.6 1.9

45–54 kg N/ha. The N uptake rate was faster up to
80 kg N/ha. At 80–120 kg N/ha, the N uptake by the
medium duration varieties was 74–101 kg N/ha.
The N uptake by rice varieties Lalat, Mahamaya
and IR 36 increased beyond 80 kg N/ha. At a high
rate of N application (80 kg N/ha) the N uptake by
Mahamaya was higher than Lalat, though there was
no difference in N uptake between these varieties
under non-fertilized conditions.
The increase in N uptake by late duration rice
varieties was faster up to 80 kg N/ha, after which it
was reduced by 40 % in several varieties. Of all the
varieties, Madhuri showed the lowest N uptake at all
N application rates. Under non-fertilized conditions,
the N uptake by all late duration varieties except
Madhuri was 60–62 kg N/ha. The N uptake by late
duration varieties at 80–120 kg N/ha was 83–129 kg
N/ha. At higher rates of N applications (80 or 120 kg
N/ha) the N uptake by Savitri, Swarna and Ranjit
was significantly greater than in other varieties.
There appeared to be less variation in NUE of
the medium duration varieties at all N rates. The
average NUE of medium duration varieties was 70 kg
grain/kg N uptake under non-fertilized conditions,
reducing to 64, 56 and 49 kg grain/kg N uptake at
40, 80 and 120 kg N/ha. Variations in NUE of the
late duration varieties were very wide. The NUE of
late duration varieties was 34–76, 30–66, 31–58 and
24–61 kg grain/kg N uptake at 0, 40, 80 and
120 kg N/ha. Of all the varieties, the NUE of Ranjit,
Savitri and Swarna was greater than for other
varieties.
Correlations among parameters
There were significant correlations of grain yield with
AR (Fig. 1 a) and with total N uptake (Fig. 1 d) of
medium duration varieties described by the equations
1 a and b.
y=3
04+3
60 AR (R2 =0
464, D:F:=88) (Eqn 1a)
y=1
86+0
03 NU (R2 =0
719, D:F:=118) (Eqn 1b)
The correlations between grain yield and HI (Fig. 1 c)
and NHI (Fig. 1 b) were not significant. However,
correlations of grain yield with AR (Fig. 2 a), with
NHI (Fig. 2 b), with HI (Fig. 2 c) and with N uptake
(Fig 2 d ) of late duration varieties were linear and
significant :
y=2
81+3
42 AR (R2 =0
171, D:F:=124) (Eqn 2a)
y=2
12+10
7 NHI (R2 =0
601, D:F:=166)
(Eqn 2b)
y=0
42+11
97 HI (R2 =0
649, D:F:=166) (Eqn 2c)
8 D. K. S W A I N E T A L.

8 (a) 6 (b)

Grain yield (t/ha)

Grain yield (t/ha)

5
6
4
4
3

2 2
0·0 0·2 0·4 0·6 0·8 0·5 0·6 0·7 0·8
Apparent N recovery N harvest index

6 (c) 6 (d )

Grain yield (t/ha)

Grain yield (t/ha)

5 5

4 4

3 3

2 2
0·3 0·4 0·5 0·6 0·7 20 40 60 80 100 120
Harvest index Total N uptake (kg/ha)
Fig. 1. Relationship between grain yield and (a) apparent N recovery, (b) N harvest index, (c) harvest index and (d) total N
uptake of medium duration rice varieties at 0 (#), 40 ($), 80 (%) or 120 (&) kg N/ha.

8 (a) 8 (b)
Grain yield (t/ha)

Grain yield (t/ha)

6 6

4 4

2 2

0 0
0·0 0·2 0·4 0·6 0·8 1·0 0·2 0·3 0·4 0·5 0·6 0·7 0·8 0·9
Apparent N recovery N harvest index

8 (c) 8 (d )
Grain yield (t/ha)

Grain yield (t/ha)

6 6

4 4

2 2

0 0
0·0 0·1 0·2 0·3 0·4 0·5 0·6 0·7 20 40 60 80 100 120 140 160
Harvest index Total N uptake (kg/ha)
Fig. 2. Relationship between grain yield and (a) apparent N recovery, (b) N harvest index, (c) harvest index and (d) total N
uptake of late duration rice varieties at 0 (#), 40 ($), 80 (%) or 120 (&) kg N/ha.

y=0
64+0
04 NU (R2 =0
423, D:F:=166) (Eqn 2d) nitrogen harvest index (kg/kg), and NU=total N
uptake (kg/ha). Comparison with Eqns 1 a and 2 a
where y=grain yield (t/ha), AR=apparent N shows that there is no large variation in the rate of
recovery (kg/kg), HI=harvest index (kg/kg), NHI= grain yield change relative to AR by the medium and
 N use efficiency of rice 9

6 (a) 7 (b)
6
Grain yield (t/ha)

Grain yield (t/ha)

5
5
4 4
3
3
2
2 1
70 90 110 130 150 170 190 40 50 60 70 80 90
Physiological N use efficiency N use efficiency for grain production
(kg total dry matter/kg total N uptake) (kg grain/kg total N uptake)

8 (c) 8 (d )

Grain yield (t/ha)

Grain yield (t/ha)

6 6

4 4

2 2

0 0
75 100 125 150 175 200 225 10 20 30 40 50 60 70 80 90
Physiological N use efficiency N use efficiency for grain production
(kg total dry matter/kg total N uptake) (kg grain/kg total N uptake)
Fig. 3. Relationship between grain yield and (a) physiological N use efficiency, (b) N use efficiency for grain production of
medium duration rice varieties, (c) physiological N use efficiency and (d) N use efficiency for grain production of late
duration rice varieties at 0 (#), 40 ($), 80 (%) or 120 (&) kg N/ha.

late duration rice varieties, as the slopes and inter- y=x2
97+0
24 NUEx0
0017 NUE2
cepts of the equations were very similar. Nevertheless,
(R2 =0
478, D:F:=166) (Eqn 4b)
there was a difference in the rate of grain yield change
to N uptake between medium and late duration where y=grain yield (t/ha), PE=physiological N use
varieties. The N uptake efficiency, as indicated by the efficiency (kg DM/kg N uptake), and NUE=N use
slope of Eqn 1 b for medium duration varieties is less efficiency for grain production (kg grain/kg N up-
than the slope of Eqn 2 d for late duration varieties. take). The correlations of NUE with leaf, stem and
The rates of increase in grain yield for unit kg ad- grain N content at maturity for medium duration
ditional N uptake were 31 and 40 kg for medium varieties (Fig. 4 a, b and c) and late duration varieties
and late duration varieties, respectively. There was a (Fig. 5 a, b and c) were highly significant. For medium
profound effect of NHI and HI on the grain yield of duration varieties the curvilinear relationships were
late duration varieties (Eqns 2 b and 2 c). The slopes described by the following quadratic equations :
of the equations indicated 1.1 and 1.2 t/ha increase
in grain yield with 0.1 unit increase in NHI and HI, y=73
9x0
76 A1 x0
16 A21 (R2 =0
31, D:F:=118)
respectively. (Eqn 5a)
The grain yield of medium duration varieties
was negatively and linearly correlated to PE y=116
3x13
5 B1 +0
61 B21 2
(R =0
537, D:F:=118)

(Fig. 3 a) as (Eqn 5b)

2
y=6
30+0
017 PE (R =0
249, D:F:=118) (Eqn 3)
y=217
1x22
5 C1 +0
75 C21 (R2 =0
824, D:F:=118)
The grain yield of late duration varieties was not (Eqn 5c)
correlated with PE (Fig. 3 c). There were significant
quadratic relationships of grain yield with NUE in Similarly, the curvilinear relationship for late
medium (Fig. 3 b) and late (Fig. 3 d) duration rice duration varieties is described by the following
varieties : quadratic equations:
y=1
23+0
12 NUEx0
0012 NUE2 y=101
9x10
2 A2 +0
32 A22 (R2 =0
48, D:F:=166)

(R2 =0
159, D:F:=118) (Eqn 4a) (Eqn 6a)

10 D. K. S W A I N E T A L.

N use efficiency for grain production

N use efficiency for grain production

90 (a) 100 (a)

(kg grain/kg total N uptake)

(kg grain/kg total N uptake)

80 80

70 60

60 40

50 20

40 0
2 4 6 8 10 12 0 2 4 6 8 10 12 14 16
Leaf N content (g/kg) Leaf N content (g/kg)

N use efficiency for grain production

N use efficiency for grain production

100 (b)
90 (b)

(kg grain/kg total N uptake)

(kg grain/kg total N uptake)

80
80
60
70

60 40

50 20

40 0
2 4 6 8 10 0 2 4 6 8 10 12
Stem N content (g/kg) Stem N content (g/kg)
N use efficiency for grain production
N use efficiency for grain production

100 (c)
90 (c)
(kg grain/kg total N uptake)
(kg grain/kg total N uptake)

80
80
60
70
40
60

50 20

40 0
6 8 10 12 14 16 0 5 10 15 20 25
Grain N content (g/kg) Grain N content (g/kg)
Fig. 5. Relationship between N use efficiency for grain pro-
Fig. 4. Relationship between N use efficiency for grain pro-
duction and (a) leaf, (b) stem and (c) grain N content of late
duction and (a) leaf, (b) stem and (c) grain N content of
duration rice varieties at 0 (#), 40 ($), 80 (%) or 120 (&)
medium duration rice varieties at 0 (#), 40 ($), 80 (%) or 120
kg N/ha.
(&) kg N/ha.

y=107
1x13
9 B2 +0
56 B22 (R2 =0
538, D:F:=166) leaf, stem, and grain N content (g/kg), respectively,
for late duration varieties.
(Eqn 6b)

y=136
0x10
2C2 +0
24 C22 (R2 =0
521, D:F:=166) DISCUSSION

(Eqn 6c) Dry matter and grain yield production of rice varieties
where y=N use efficiency for grain production It was not surprising that the grain yield of rice
(kg grain/kg N uptake); A1, B1 and C1 are the leaf, varieties at various N rates reflected the yield at
stem, and grain N content (g/kg) respectively for control. The correlations between non-fertilized and
medium duration varieties, and A2, B2 and C2 are the N fertilized treatments for DM and grain yield of
 N use efficiency of rice 11

Table 5. Agronomic N use efficiency, physiological N use efficiency, apparent recovery of applied N, N harvest
index and harvest index of medium and late duration rice varieties as influenced by N applications

Agronomic N Physiological
use efficiency N use efficiency Apparent
(kg grain/kg N (kg dry matter/kg N recovery of N harvest Harvest
Treatments applied) uptake) applied N index index

Medium duration
Varieties
IR-36 12.8 132 0.35 0.68 0.49
Mahamaya 12.7 126 0.39 0.65 0.51
Kranti 9.5 123 0.34 0.68 0.51
Lalat 11.1 132 0.37 0.65 0.45
Khitish 6.4 128 0.24 0.67 0.46
N rates (kg N/ha)
0 – 148 – 0.68 0.48
40 11.5 136 0.30 0.67 0.50
80 13.2 117 0.40 0.67 0.49
120 6.7 111 0.31 0.64 0.46
S.E.M.
Variety (D.F.=12) – 3.5 – 0.011 0.010
Nitrogen (D.F.=45) – 4.7 – 0.013 0.012
Late duration
Varieties
Mahsuri 11.8 132 0.40 0.51 0.32
Madhuri 5.6 126 0.34 0.45 0.27
Savitri 18.0 146 0.57 0.68 0.45
Rajshree 13.0 120 0.46 0.57 0.34
Swarna 22.4 119 0.56 0.69 0.46
Sashi 10.9 130 0.49 0.56 0.37
Ranjit 17.0 136 0.55 0.65 0.48
N rates (kg N/ha)
0 – 153 – 0.57 0.37
40 15.6 132 0.51 0.60 0.38
80 15.3 121 0.50 0.61 0.40
120 11.4 115 0.44 0.56 0.39
S.E.M.
Variety (D.F.=18) – 4.7 – 0.010 0.011
Nitrogen (D.F.=63) – 4.7 – 0.011 0.011

medium duration varieties at N applications of 40 to
120 kg/ha were significant (P<0.01). The coefficients
of determination (R2) were 0.79–0.61 for DM and
0.49–0.33 for grain yield production. R2 values for the
DM and grain yield of the late duration varieties were
0.59–0.26 and 0.86–0.64, respectively, at N rates of
40–120 kg/ha. The decrease in R2 values at high N
rates indicated larger variation in DM and grain yield
due to variety and N interactions. The differences in
grain yield and N utilization of several rice varieties
suggested that N use efficiency and choice of variety
should often be specific to the season (De Datta &
Broadbent 1988; Singh et al. 1998), but the N use
efficiency is determined retrospectively. It implies that
there could be N efficient varieties of medium and late
duration groups that produce high grain yield at both
low and high rates of N application in the wet season
of the tropics. Inthapanya et al. (2000 a) found that
the coefficient of variation in NUE was small under
fertilized conditions, providing more opportunity
to differentiate varieties, statistically. Variety and N
interaction strongly influenced overall performance
of some varieties.
An attempt was made to classify the medium and
late duration rice varieties into N efficient and non-
efficient groups, using the observations of ANUE,
PE, AR, NHI and HI of each variety were taken into
consideration (Table 5). For medium duration
varieties, the average ANUE and AR was the highest
at 80 kg N/ha. The HI and NHI of these varieties
were not affected at 80 kg N/ha but were lower at
120 kg N/ha. The PE decreased as the N applications
increased. Similarly, the NHI and HI of the late
duration varieties were highest at 80 kg N/ha. The
12 D. K. S W A I N E T A L.
differences in AR or ANUE at 40 and 80 kg N/ha
were not significant. The N response of both medium
and late duration varieties in the wet season remained
high up to 80 kg N/ha, beyond which N application
had no further effect on grain yield production.
De Datta & Malabuyoc (1976) obtained N responses
up to 80–90 kg N/ha during the wet season in the
tropics. This indicated that the full efficiency of the
medium and late duration varieties was expressed at
80 kg N/ha. The increase in N uptake and DM of
many medium and late duration varieties beyond
80 kg N/ha did not result in a significant increase
in yield.
The late duration varieties had well-developed root
systems (able to absorb sub-soil N) and a longer
vegetative growth period, both conducive to higher
uptake of N (Vergara et al. 1966). As a result, the
total N uptake by the late duration rice varieties was
higher than the medium duration varieties. High N
applications increased N uptake, but if the uptake
amount exceeded an optimum value, the yield
decreased. In contrast, the medium duration varieties
were of panicle number type, which absorb more
surface N because of their shallow rooting system.
Large N uptake resulted in a very high leaf
area index, which enhanced carbohydrate loss via
dark respiration (Shi & Akita 1993). The limited
vegetative growth period of medium duration
varieties prevented excessive production of vegetative
biomass.
In general, the mean grain yield of all the medium
duration varieties was 4.5 t/ha at 80 kg N/ha. But
large differences were observed in N uptake by these
varieties at 80 kg N/ha. Khitish was not efficient
because it exhibited the least ANUE, AR, N uptake
and grain yield in both fertilized and non-fertilized
conditions. Kranti exhibited poor yield at all N rates
except 80 kg N/ha, which resulted in low ANUE and
AR compared to IR 36, Mahamaya and Lalat, the
efficient varieties. The late duration rice varieties
were also distinguished into two groups, efficient and
non-efficient. The grain yields of these varieties
were 2.6–6.2 t/ha with a mean yield of 4.6 t/ha at
80 kg N/ha. The varieties Ranjit, Savitri and Swarna
were efficient and produced higher grain yield than
the others, with an average of 5.8 t/ha. The DM, N
uptake, HI and NHI of these three varieties were
also high. Mahsuri, Rajshree, Madhuri and Sashi
were not efficient. Non-efficient rice varieties have
poor translocation from source to sink, resulting in
low yield and poor HI (Ladha et al. 1998). The HI and
NHI of medium duration varieties were 0.46–0.51
and 0.65–0.68, respectively, when averaged over the
N rates. The efficient late duration varieties Ranjit,
Savitri and Swarn had HI and NHI of 0.45–0.48
and 0.65–0.69, respectively. Dingkuhn et al. (1991)
reported NHI of 0.60–0.72 for three IRRI semi dwarf
rice varieties differing in growth duration. Guindo
et al. (1994) used N15 fertilizers and concluded that
NHI of two lowland rice cultivars was 0.58–0.62.
The non-efficient late duration varieties have poor HI
(0.27–0.37) and poor NHI (0.45–0.56). Differences
in NUE could be due in part to variety differences
in apical development (despite external growth
stages being similar) leading to different internal
N concentrations in the developing ear. This has
implications for spikelet abortion (Senanayake et al.
1994).
Many non-efficient late duration varieties had high
DM even under non-fertilized condition and they
were very tall (131–142 cm) and susceptible to lodging
in high rainfall areas of the tropics resulting in poor
HI. Grain yield in cereals is related to biological yield
and HI (Donald & Hamblin 1976). Grain yield or
economic yield could be increased either by increasing
DM or by increasing HI. Grain yield increases in
cereals such as rice could be accomplished with an
increase in total biological yield (Rahman 1984) as
demonstrated in barley by Naylor et al. (1998) and
this might be possible through breeding for improved
semi-dwarf N responsive rice varieties of medium
duration. Roberts et al. (1993) observed that the
highest HI exhibited by California lowland rice
cultivars under direct seeding was 0.59. This showed
that the HI of medium or late duration varieties could
be improved through breeding or crop management
methods for yield increase.
The significant correlation between grain yield, N
uptake and N use efficiency for grain production
indicated that the grain yield of medium duration
varieties was mostly dependent on total N uptake of
the crop. However, the grain yield of late duration
varieties was governed by both total N uptake and the
utilization efficiency of the variety. The correlation
coefficient for the relationship between grain yield
and the HI and the NHI of late duration varieties was
higher than that for the N uptake and AR. This
showed that the plant N use efficiency of late duration
varieties strongly contributed to differences in grain
yield. The average grain yield of efficient medium
duration varieties was lower than that of the efficient
late duration varieties. This yield difference was
closely associated with lower N uptake by the
medium duration varieties (Cassman et al. 1993).
The present results show that the rate of increase in
grain yield for 1 kg additional N uptake of late
duration varieties was 29 % higher than that of
medium duration varieties.
Plant N use efficiency and N content for optimum
grain production
Optimum grain yield could be associated with
optimum PE and NUE of the rice plant because
the higher N application (120 kg N/ha) did not
ensure higher grain yield. It might be inferred that
 N use efficiency of rice 13

Table 6. Path coefficient analysis showing direct and varieties and these N contents were 10 g/kg, 8 g/kg
indirect effect of N content in grain, leaf and stem at and 14 g/kg for leaf, stem and grain, respectively, at
maturity on N use efficiency for grain production maturity. Tirol-Padre et al. (1996) identified geno-
(NUE) of medium and late duration rice varieties types possessing promising traits for improved
N uptake and utilization efficiency among 180 rice
Characters Leaf Stem Grain NUE genotypes of different growth duration from 100 to
135 days without the addition of N fertilizer. They
Medium duration indicated genotypic variation in stem N contents and
Leaf x0.1640 x0.1371 x0.1370 x0.8776 suggested low stem N content for yield improvement.
Stem x0.3418 x0.4089 x0.3837 x0.9638 Significant genotypic variation in nutrient use
Grain x0.3718 x0.4178 x0.4452 x0.9659 efficiency also contributed to genotypic variation for
Residual 0.1788
effect
grain yield (Inthapanya et al. 2000 b). A larger effect
R2 0.9680 of variety and smaller effect of variety by location
D .F . 118 interaction was observed for N use efficiency.
Late duration
NUE was significantly and negatively correlated
Leaf x0.1248 x0.0826 x0.0827 x0.6737 (P<0.001) with leaf, stem and grain N content at
Stem x0 2673 x0.4038 x0.3097
. x0.8123 maturity at the phenotypic level (Table 6) for medium
Grain x0.2817 x0.3259 x0.4248 x0.8173 and late duration varieties. Path coefficient analysis
Residual 0.4906 showed that the residual effect at the phenotypic level
effect was low for medium duration varieties, and the
R2 0.7593 independent variables explained about 82 % of the
D .F . 166 variability in NUE. In late duration varieties, about
51 % of the variability in NUE could be explained by
N content in plant organs at maturity. The direct
influence of grain and stem N content at maturity for
improvement in grain yield by late duration varieties NUE was significantly higher than leaf N content of
through PE could be difficult as the correlation medium and late duration varieties. The multiple
between grain yield and PE was not significant. The regressions of NUE on the grain, leaf, and stem N
yield of medium duration varieties was influenced content at maturity was described by the equations
mainly by total N uptake, followed by PE and NUE.
Using Eqns 4a and b, the optimum levels of NUE for y=126
1x1
0 A1 x2
5 B1 x3
9 C1
medium and late duration varieties were computed to (Medium duration varieties)
be 49 and 68 kg grain/kg N uptake, respectively. The
optimum grain yields of 4.5 t/ha by medium duration (R2 =0
89, P<0
001, S:D:=0
949)
varieties and 5.8 t/ha by late duration varieties are (S:E: values: A1 =0
02, B1 =0
032, C1 =0
022)
attained with these optimum values of NUE, which (Eqn 7a)
could serve as a guideline for developing new rice
varieties.
Yoshida (1981) reported a quadratic relationship y=112
6x1
6 A2 x3
6 B2 x2
6 C2
between photosynthetic rate and N content per leaf (Late duration varieties)
area. The indica rice varieties showed less promotion
in their photosynthetic rate by the high N supply than (R2 =0
748, P<0
001, S:D:=1
362)
did japonica varieties. At high N applications, some (S:E: values: A2 =0
04, B2 =0
05, C2 =0
03)
rice varieties have a lower proportion of spikelets (Eqn 7b)
filled. The probability of lodging or culm bending is
higher at high levels of N and disrupts the movement where y=N use efficiency for grain production
of assimilates and absorbed nutrients. High N con- (kg grain/kg N uptake); A1, B1 and C1 are the leaf,
tent and uptake in plant organs due to supra-optimal stem and grain N content (g/kg) of medium duration
N application or soil extraction can reduce the utiliz- varieties, respectively ; and A2, B2 and C2 are the leaf,
ation efficiency for maximum grain production. The stem and grain N content (g/kg) of late duration
leaf, stem and grain N contents at maturity of 4.0, varieties, respectively. The intercept and all the
4.0 and 9.0 g/kg (from Eqns 6 a, b and c respectively), coefficients of grain, leaf and stem N contents at
coincide with the optimum NUE (68 kg grain/kg N maturity were significant and confirmed a higher
uptake) of late duration varieties. The N contents contribution of grain N content for NUE of medium
in plant organs for optimum N use efficiency (49 kg duration varieties and stem N content for NUE of
grain/kg N uptake) of medium duration varieties late duration varieties. According to Sahu et al.
(derived from Eqns 5 a, b and c) appeared to be much (1997) the grain yield and N use efficiency showed a
higher than the corresponding values in late duration moderate level of heritability and selection would not
14 D. K. S W A I N E T A L.
be effective based on N use efficiency alone. Hence, a
selection of rice varieties for grain yield and NUE
should be based on the N content of grain, stem
and leaf.
REFERENCES
AGGARWAL, P. K., KROPFF, M. J., CASSMAN, K. G. & TEN
BERGE, H. F. M. (1997). Simulating genotypic strategies
for increasing rice yield potential in irrigated, tropical
environments. Field Crops Research 51, 5–17.
BROWN, L., SCHOLEFIELD, D., JEWKES, E. C., PREEDY, N.,
WADGE, K. & BUTLER, M. (2000). The effect of sulphur
application on the efficiency of nitrogen use in two
contrasting grassland soils. Journal of Agricultural
Science, Cambridge 135, 131–138.
CASSMAN, K. G., KROPFF, M. J., GAUNT, J. & PENG, S.
(1993). Nitrogen use efficiency of rice reconsidered: what
are the key constraints? Plant and Soil 155/156, 359–362.
CRASWELL, E. T. & VLEK, P. L. G. (1979). Fate of fertilizer
nitrogen applied to wetland rice. In Nitrogen and Rice,
Symposium Proceedings, pp. 17–192. Manila, Philippines:
International Rice Research Institute.
DE DATTA, S. K. & BROADBENT, F. E. (1988). Methodology
for evaluating nitrogen utilization efficiency by rice
genotypes. Agronomy Journal 80, 793–798.
DE DATTA, S. K. & MALABUYOC, J. (1976). Nitrogen
response of lowland and upland rice in relation to
tropical environmental conditions. In Climate and Rice,
pp. 509–539. Los Baños, Philippines: International Rice
Research Institute.
DINGKUHN, M., SCHNIER, H. F., DE DATTA, S. K.,
DORFFLING, K. & JAVELLANA, C. (1991). Relationships
between ripening phase productivity and crop duration,
canopy photosynthesis and senescence in transplanted
and direct seeded lowland rice. Field Crops Research 26,
327–345.
DONALD, C. M. & HAMBLIN, J. (1976). The biological yield
and harvest index of cereals as agronomic and plant
breeding criteria. Advances in Agronomy 28, 361–405.
FRANCIS, D. D., SCHEPERS, J. S. & VIGIL, M. F. (1993).
Post-anthesis nitrogen loss from corn. Agronomy Journal
85, 659–663.
GOMEZ, K. A. & GOMEZ, A. A. (1984). Statistical Procedures
for Agricultural Research, 2nd edn. New York: John
Wiley and Sons.
GUINDO, D., NORMAN, R. J. & WELLS, B. R. (1994).
Accumulation of fertilizer nitrogen-15 by rice at different
stages of development. Soil Science Society of America
Journal 58, 410–415.
HARPER, L. A., SHARPE, R. R., LANGDALE, G. W. &
GIDDENS, J. E. (1987). Nitrogen cycling in a wheat crop:
soil, plant, and aerial nitrogen transport. Agronomy
Journal 79, 965–973.
INTHAPANYA, P., SIPASEUTH, SIHAVONG, P., SIHATHEP, V.,
CHANPHENGSAY, M., FUKAI, S. & BASNAYAKE, J. (2000 a).
Genotypic performance under fertilized and non-fertilized
conditions in rain fed lowland rice. Field Crops Research
65, 1–14.
INTHAPANYA, P., SIPASEUTH, SIHAVONG, P., SIHATHEP, V.,
CHANPHENGSAY, M., FUKAI, S. & BASNAYAKE, J. (2000 b).
Genotype differences in nutrient uptake and utilization
for grain yield production of rain fed lowland rice
Acknowledgement is due to the research project
which was financed by the World Bank funded
National Agricultural Technology Project (NATP
RRPS 25) from May 2000 to December 2003.
under fertilized and non-fertilized conditions. Field Crops
Research 65, 57–68.
LADHA, J. K., KIRK, G. J. D., BENNETT, J., PENG, S.,
REDDY, C. K., REDDY, P. M. & SINGH, U. (1998).
Opportunity for increasing nitrogen use efficiency from
improved lowland rice germplasm. Field Crops Research
56, 41–72.
NAYLOR, R. E. L., STOKES, D. T. & MATTHEWS, S. (1998).
Biomass, shoot uniformity and yield of winter
barley. Journal of Agricultural Science, Cambridge 131,
13–21.
RAHAMAN, M. S. (1984). Breaking the yield barriers
in cereals with particular reference to rice. Journal of
the Australian Institute of Agricultural Science 50,
228–232.
ROBERTS, S. R., HILL, J. E., BRANDON, D. M., MILLER, B. C.,
SCARDACI, S. C., WICK, C. M. & WILLIAMS, J. F. (1993).
Biological yield and harvest index in rice: nitrogen
response of tall and semidwarf cultivars. Journal of
Production Agriculture 6, 585–588.
SAHU, R. K., TIROL-PADRE, A., LADHA, J. K., SINGH, U.,
BHAGEL, S. S. & SRIVASTAVA, M. N. (1997). Screening
genotypes for nitrogen use efficiency on a nitrogen
deficient soil. Oryza 34, 350–357.
SENANAYAKE, N., NAYLOR, R. E. L. & DE DATTA, S. K.
(1996). Effect of nitrogen fertilization on rice spikelet
differentiation and survival. Journal of Agricultural
Science, Cambridge 127, 303–309.
SENANAYAKE, N., NAYLOR, R. E. L., DE DATTA, S. K. &
THOMSON, W. J. (1994). Variation in development of
contrasting rice cultivars. Journal of Agricultural Science,
Cambridge 123, 35–39.
SHI, Q. & AKITA, S. (1993). Potential dry matter production
and grain yield of present rice cultivars in the tropics.
Acta Agriculturae Universitatis Jiangxiensis 15, 129–137.
SINGH, U., LADHA, J. K., CASTILLO, E. G., PUNZALAN, G.,
TIROL-PADRE, A. & DUQUEZA, M. (1998). Genotypic
variation in nitrogen use efficiency in medium and long
duration rice. Field Crops Research 58, 35–53.
STA. CRUZ, P. C. & WADA, G. (1994). Genetic variation in
nitrogen uptake by rice and the effects of management
and soil fertility. In Rice Roots, Nutrient and Water Use
(Ed. G. J. D. Kirk), pp. 29–40. Manila, Philippines:
International Rice Research Institute.
THIYAGARAJAN, T. M., SIVASAMY, R. & BUDHAR, M. N.
(1995). Procedure for collecting plant samples at different
growth stages of transplanted rice crop. In SARP
Research Proceedings: Nitrogen Management Studies in
Irrigated Rice (Eds T. M. Thyagarajan, H. F. M. ten
Berge & M. C. S. Wopereis), pp. 99–102. Manila,
Philippines: International Rice Research Institute.
TIROL-PADRE, A., LADHA, J. K., SINGH, U., LAURELES, E.,
PUNZALAN, A. & AKITA, S. (1996). Grain yield perform-
ance of rice genotypes at suboptimal levels of soil N as
affected by N uptake and utilization efficiency. Field
Crops Research 46, 127–143.
 N use efficiency of rice
VERGARA, B. S., TANAKA, A., LILIS, R. & PURANABHAVUNG,
S. (1966). Relationship between growth duration and
grain yield of rice plants. Soil Science and Plant Nutrition
12, 31–39.
VON UEXKULL, H. R. (1993). New trend in rice crop
production: enhancement of nutrient use efficiency.
In New Frontiers in Rice Research (Eds K. Muralidharan
& E. A. Siddiq), pp. 290–299. Hyderabad, India:
Directorate of Rice Research.
YADAV, R. L., SINGH, V. K., DWIVEDI, B. S. & SHUKLA, A. K.
(2003). Wheat productivity and N use-efficiency as
influenced by inclusion of cowpea as a grain legume in a
15
rice-wheat system. Journal of Agricultural Science,
Cambridge 141, 213–220.
YOSHIDA, S. (1981). Fundamentals of Rice Crop Science. Los
Baños, Philippines: The International Rice Research
Institute.
YOSHIDA, S., FORNO, D. A., COCK, J. H. & GOMEZ, K. A.
(1976). Laboratory Manual for Physiological Studies of
Rice (Third Edition). Manila, Philippines: The Inter-
national Rice Research Institute.
ZADOKS, J. C., CHANG, T. T. & KONZAK, C. F. (1974). A
decimal code for the growth stages of cereals. Weed
Research 14, 415–421.