Developing rice cultivars for high-fertility upland systems in the Asian tropics
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Field Crops Research 97 (2006) 43–52
Developing rice cultivars for high-fertility upland
systems in the Asian tropics
G.N. Atlin a,*, H.R. Lafitte a, D. Tao b, M. Laza a, M. Amante a, B. Courtois c
a International Rice Research Institute, DAPO Box 7777, Metro Manila, Philippinesb Yunnan Academy of Agricultural Sciences, Kunming 650205, People’s Republic of China
c CIRAD-Biotrop TA40/03, Avenue Agropolis, 34398 Montpellier Cedex 5, France
Abstract
Traditionally, upland rice is grown in Asia in low-input, subsistence systems. More productive upland systems, using more fertilizer and
improved varieties, are emerging in China and Philippines, and could contribute to productivity increases in rainfed environments in other
countries. Here, we evaluate, on-station and on-farm, the yield under upland management of improved indica upland cultivars selected for
yield under high-fertility conditions. These cultivars are compared with traditional and improved tropical japonica upland varieties, and with
elite indica high-yielding varieties (HYV) developed for irrigated lowland production, to characterize the features of varieties that produce
high yields in favorable upland environments. Forty-four improved and traditional varieties and experimental lines were evaluated in irrigated
lowland, non-stressed upland, moderately stressed upland, severely water-stressed upland, and low-fertility upland environments in southern
Luzon, Philippines. Correlations between yields in non-stress and mild-stress environments were low but positive. Some cultivars, like
IR55423-01, were among the highest yielding under both conditions, indicating that high yield and moderate water-stress tolerance can be
combined. Upland-selected indica varieties yielded 3.56 t ha�1 in favorable upland environments on-station in southern Luzon, out-
performing improved tropical japonica and irrigated varieties by 23 and 69%, respectively. They were also the highest-yielding class in
infertile, acid soils. The improved upland indica cultivars are about 110 cm tall under favorable upland conditions and maintain a harvest
index of nearly 0.4, or about one-third higher than other cultivar types. The best upland-adapted rice varieties produced average yields on-farm
of 3.3 and 4.1 t ha�1 in southern Luzon and Yunnan, respectively, outyielding traditional checks by 30–50% with moderate N application.
Screening under both high-fertility, non-stress conditions and moderate reproductive-stage stress appears to be needed to develop cultivars
combining high-yield potential with drought tolerance. Upland-adapted indica cultivars offer a new approach to increasing productivity and
reducing risk in Asian rainfed rice systems.
# 2005 Elsevier B.V. All rights reserved.
Keywords: Rice cultivars; Upland environments; Harvest index
1. Introduction
Traditional upland rice crops are grown in unbunded,
unflooded fields, where soil conditions in the root zone
remain aerobic through most of the growing season. In most
traditional Asian upland rice-growing areas, soils are acidic
and infertile. Farmers usually treat upland rice as a
subsistence crop, investing little in inputs beyond family
labor. Because upland rice varieties are grown without
irrigation in unsaturated soils, they are considered to be
* Corresponding author. Tel.: +63 2 845 0563x2586; fax: +63 2 845 0606.
E-mail address: g.atlin@cgiar.org (G.N. Atlin).
0378-4290/$ – see front matter # 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2005.08.014
drought tolerant. However, upland rice yields in traditional
systems are low, averaging 1–2 t ha�1 in most growing
regions. Intensification of management of these systems
with currently available germplasm is difficult because most
traditional upland rice varieties are tall, low-tillering, and
prone to lodging when grown under conditions of favorable
moisture and high-soil fertility.
Recently, a new class of upland-adapted cultivars with
improved lodging resistance, harvest index, and input
responsiveness has been developed by breeding programs
in China, Brazil, and the Philippines. These varieties
combine some of the yield potential-enhancing traits of
lowland high-yield varieties with adaptation to aerobic soils.
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–5244
They are often referred to as aerobic rice varieties to
differentiate them from traditional upland rice varieties
adapted to low-input systems. The distinguishing feature of
aerobic production systems is that crops are direct-seeded in
free-draining, non-puddled soils where no standing water
layer is maintained in the field, and roots grow in a mainly
aerobic environment. A commercial cropping system based
on aerobic rice has spread widely in Brazil as a result of the
development of improved varieties (Pinhiero et al., 2005).
Aerobic rice is also of interest in water-short lowland rice
production systems. Shifting from conventional flooded
systems to direct seeding in non-puddled, non-flooded fields
can reduce water requirements for rice production by over
50% by reducing percolation, seepage, and evaporation
losses (Castaneda et al., 2002). This gain is at the expense of
some reduction in yield relative to that obtainable in fully
irrigated systems, but may not result in significant yield loss
relative to conventional rainfed lowland management in
drought-prone areas. Both water-short irrigated and drought-
prone rainfed lowland systems may, therefore, benefit from
aerobic rice systems. Aerobic production systems based on
high-yield, input-responsive upland cultivars have already
been developed to replace conventional irrigated lowland
rice production on the water-short plains of northeastern
China, producing yields of 4.5–6.5 t ha�1, with substantial
water savings relative to conventional irrigated production
(Wang et al., 2002; Bouman et al., 2005).
Aerobic conditions can comprise a wide range of soil
hydrological environments, from near saturated to well
below field capacity. It is not known to what extent these
conditions should be considered separate breeding targets
(Atlin and Lafitte, 2002). There is also little information in
the literature regarding optimal characteristics of high-yield
upland rice varieties for the Asian tropics, and the relative
yield potential of the major germplasm groups under high-
input upland management. This information is needed to
identify promising parents and develop effective screens for
aerobic rice systems. There is also little information
available on the on-farm performance of improved upland
rice cultivars across locations and years under rainfed
conditions. This information is needed to assess the
productivity and stability of intensified upland rice-based
cropping systems. The objectives of this report, are
therefore, to:
� i
dentify cultivars and germplasm types with high-yieldpotential under a range of upland hydrological conditions;
� d
ocument the yield of currently available upland ricecultivars and breeding lines under purely rainfed
conditions and farmer management in emerging intensi-
fied upland rice-based systems in Philippines and Yunnan,
China;
� c
haracterize some of the common attributes of cultivarswith superior performance in aerobic rice systems;
� p
ropose screening methods that can be used to developsuperior cultivars for aerobic rice systems.
2. Materials and methods
2.1. Performance of upland, lowland, and aerobic
varieties in five management regimes in southern Luzon,
Philippines
To clarify the degree of genotype-specific adaptation to
hydrologic and soil fertility conditions, and to characterize
differences between germplasm groups, 44 upland and
irrigated lowland varieties were screened in 14 three- or
four-replicate randomized complete-block trials classified
into five environment types differing in soil fertility and
hydrology:
(a) i
rrigated lowland fields established via transplanting atIRRI in the dry season, in which the trials experienced
no water stress (NSL);
(b) u
pland, high-fertility fields established via directseeding at IRRI during the wet season, with soil
maintained at or near field capacity via occasional
irrigation, and in which trials experienced little stress
(NSU);
(c) u
pland, low-fertility fields established via direct seedingat Siniloan, Philippines, a P-deficient acid upland site,
during the wet season (LFU);
(d) u
pland fields at IRRI established via direct seeding inthe dry season, with frequent irrigation resulting in mild
to moderate stress (MSU);
(e) u
pland fields at IRRI established via direct seeding in thedry season, with infrequent irrigation after the maximum
tillering stage resulting in severe stress (SSU).
The 44 varieties are listed in Table 1. Thirty-six are
improved upland types, three are traditional Philippine
tropical japonica upland varieties, one is a traditional aus
variety and four are varieties from the IRRI irrigated rice-
breeding program. The improved upland varieties were of
diverse origin, representing both the indica and tropical
japonica groups. They are the output of Philippine, Indian,
Brazilian, Colombian, Indonesian, and West African
breeding programs targeted at a range of management
conditions. All varieties are photoperiod-insensitive or only
weakly sensitive, ranging in duration from approximately
90–130 days in duration. All varieties set seed well in both
the wet and dry seasons in Luzon.
Environments are described in Table 2. In all cases, these
environment types were represented by at least two trials in
different field locations. In all but the SSU environment type,
trials were conducted in at least two seasons. All upland
varieties were included in all trials, but trials conducted
before the dry season of 2002 did not include the four
irrigated varieties. In the dry season, all trials were planted
between January 3 and January 20. In the wet season, all
trials were planted between June 10 and July 10.
Trials were conducted at two sites in southern Luzon: Los
Banos and Siniloan. The Los Banos trials were conducted at
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–52 45
Table 1
Description of 44 varieties evaluated in southern Luzon in five environments differing in soil fertility and hydrology
Cultivar Germplasm group Variety type Adaptation Origin
AUS 196 Aus Traditional Upland India
AZUCENA Tropical japonica Traditional Upland Philippines
B6144F-MR-6-0-0 Indica Improved Upland Indonesia
C 22 Tropical japonica Improved Lowland Philippines
CT13370-12-2-M Indica/tropical japonica Improved Upland Colombia
CT13377-4-2-M Tropical japonica Improved Upland Colombia
CT13382-8-3-M Indica/tropical japonica Improved Upland Colombia
CT6510-24-1-2 Indica Improved Upland Colombia
CT6516-24-3-2 Indica/tropical japonica Improved Upland Colombia
DINORADO Tropical japonica Traditional Upland Philippines
IR64 Indica Improved Lowland Philippines
IR72 Indica Improved Lowland Philippines
IR47686-30-3-2 Tropical japonica Improved Upland Philippines
IR55419-04 Indica Improved Upland Philippines
IR55423-01 Indica Improved Upland Philippines
IR60080-46A Tropical japonica Improved Upland Philippines
IR65261-09-1-B Tropical japonica Improved Upland Philippines
IR65907-116-1-B Indica/tropical japonica Improved Upland Philippines
IR66417-18-1-1-1 Tropical japonica Improved Upland Philippines
IR66421-062-1-1-2 Indica/tropical japonica Improved Upland Philippines
IR66424-1-2-1-5 Indica/tropical japonica Improved Upland Philippines
IR68702-072-1-4-B Tropical japonica Improved Upland Philippines
IR70358-84-1-1 Indica/tropical japonica/aus Improved Upland Philippines
IR70360-38-1-B-1 Tropical japonica Improved Upland Philippines
IR71524-44-1-1 Tropical japonica Improved Upland Philippines
IR71525-19-1-1 Indica/tropical japonica Improved Upland Philippines
IR72768-15-1-1 Tropical japonica Improved Upland Philippines
IRAT 170 Tropical japonica Improved Upland Ivory Coast
IRAT 177 Tropical japonica Improved Upland Ivory Coast
IRAT 212 Tropical japonica Improved Upland Ivory Coast
IRAT 216 Tropical japonica Improved Upland Ivory Coast
MARAVILHA Tropical japonica Improved Upland Brazil
PALAWAN Tropical japonica Traditional Upland Philippines
PRIMAVERA Unknown Improved Upland Brazil
PSBRC 80 Indica Improved Lowland Philippines
PSBRC 82 Indica Improved Lowland Philippines
UPL RI-5 Indica Improved Upland Philippines
UPL RI-7 Indica Improved Upland Philippines
VANDANA Aus/tropical japonica Improved Upland India
WAB 181-18 Tropical japonica Improved Upland Ivory Coast
WAB 56-125 Tropical japonica Improved Upland Ivory Coast
WAB 638-1 Tropical japonica Improved Upland Ivory Coast
WAB 96-1-1 Tropical japonica Improved Upland Ivory Coast
WAY RAREM Indica Improved Upland Indonesia
the IRRI Experiment Station (148110N, 1218150E, 21 masl),
where annual average rainfall was 2100 mm in 1979–2002.
The soil is an Andaqueptic Haplaquol with pH of 6.5. The
Siniloan site is located approximately 60 km east of Los
Banos (148280N, 1218290E, 310 masl), on a light-textured,
acid (pH 3.7), P-deficient Typic Palehumult soil. Long-term
average annual rainfall data are not available for Siniloan,
but the annual mean for 1997–2000 was 4500 mm. Trials
were conducted only in the wet season at Siniloan, and in
both the wet and dry seasons at Los Banos. From 1 January
to 30 April, total rainfall was 257.2, 56.1, and 34.1 mm in the
dry seasons of 2001, 2002, and 2003, respectively; pan
evaporation was 699, 590, and 734 mm for the same three
periods. The vapor pressure deficit in the period from 60
days after sowing to harvest was 0.75 kPa in the dry season
experiments, compared to 0.50 kPa in the wet season. Free
water was not present within 1 m of the soil surface during
the dry season. In the wet season, free water was present
within 0.3 m of the surface during periods of high rainfall,
but usually fell below 0.5 m for several extended periods.
Wet season upland trials experienced little water stress at
either site, and were completely rainfed. Irrigated lowland
trials were maintained with constant standing water between
5 and 15 cm deep from transplanting to pre-harvest
drainage. Dry season upland trials were either sprinkler-
or basin-irrigated, with approximately 30 mm of water
applied at each sprinkler irrigation, and approximately
50 mm applied at each basin irrigation. Dry season irrigation
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–5246
Table 2
Description of trials conducted in southern Luzon in which 44 varieties were evaluated for adaptation to five environments differing in soil fertility and
hydrology
Environment
typeaLocation Season Year N–P2O5–K2O Hydrology Establishment method Irrigation
method
Trial mean
yield (t ha�1)
NSL IRRI Dry 2001 60–60–60 Lowland Transplanting Continuous flood 2.70
NSL IRRI Dry 2002 120–30–30 Lowland Transplanting Continuous flood 2.82
NSL mean 2.76
NSU IRRI Wet 2001 130–100–100 Upland Direct seeding Rainfed 2.33
NSU IRRI Wet 2002 60–60–60 Upland Direct seeding Rainfed 2.15
NSU mean 2.23
MSU IRRI Dry 2002 110–60–60 Upland Direct seeding Sprinkler twice weekly 0.91
MSU IRRI Dry 2001 160–40–40 Upland Direct seeding Sprinkler twice weekly 1.36
MSU IRRI Dry 2001 0–40–40 Upland Direct seeding Sprinkler twice weekly 0.90
MSU IRRI Dry 2003 90–60–60 Upland Direct seeding Basin once every 10 days 1.07
MSU mean 1.05
LFU Siniloan Wet 2000 60–90–90 Upland Direct seeding Rainfed 1.42
LFU Siniloan Wet 2001 30–30–30 Upland Direct seeding Rainfed 0.59
LFU Siniloan Wet 2002 30–0–0 Upland Direct seeding Rainfed 0.99
LFU mean 1.00
SSU IRRI Dry 2002 90–60–60 Upland Direct seeding Sprinkler once weekly 0.55
SSU IRRI Dry 2002 0–0–0 Upland Direct seeding Sprinkler once weekly 0.36
SSU IRRI Dry 2002 70–30–30 Upland Direct seeding Sprinkler once weekly 0.49
SSU mean 0.46
a NSL, non-stressed lowlands; NSU, non-stressed uplands; MSU, moderately water-stressed uplands; LFU, low-fertility uplands; SSU, severely water-
stressed uplands.
regimes were constant from sowing to harvest. Because
irrigation frequencies and amounts were maintained con-
stant through the growing season, intermittent, repeated
stress between irrigations was experienced by all varieties,
regardless of flowering date and duration.
Weeds were controlled though a combination of early
season herbicide applications and manual weeding. Che-
mical insect control was used as required. A severe
infestation of mole crickets damaged seedlings in the DS
2002 trials at Los Banos. Heavy rainfall events shortly after
sowing resulted in severe localized erosion within the
Siniloan site in 2001 and 2002.
Plot size ranged from 4.5 to 7.5 m2. The entire plot was
harvested for grain yield. Yields given are for air-dried paddy,
at approximately 12% moisture content. Harvest index and
panicle number were determined from a 0.25 m2 sample.
To estimate cultivar correlations across environment
types for mean yield, least-squares means across trials
Table 3
Correlations among mean yields of 44 aerobic, upland, and irrigated varieties ac
Non-stressed uplands Moderately water-s
Non-stressed lowlands 0.72** 0.31*
Non-stressed uplands 0.44**
Moderately water-stressed uplands
Severely water-stressed uplands
* Significant at 0.05.** significant at 0.01.
within environment types were first estimated using a mixed
model in which trials and replicates were considered random
factors, and cultivars were considered fixed. The REML
algorithm of the SAS MIXED procedure (SAS Institute,
1992) was used for the analysis.
A subset of 15 cultivars (Table 3) could be unambigu-
ously classified into five types, based on their Glazsmann
(1987) isozyme genotype and selection history: irrigated
lowland high-yield varieties (HYV), improved upland-
adapted indica varieties selected under high-fertility
management (IHF), traditional Philippine tropical japonica
upland varieties (JTV), improved tropical japonica-derived
upland varieties targeted at low-fertility environments (JLF),
and early maturing, drought-tolerant upland varieties
targeted at drought-prone environments (EDT), with three
cultivars within each type group. Means for each cultivar
type were estimated within environment types, using a
model similar to the one described above, but subdividing
ross five environment types in southern Luzon, 2000–2003
tressed uplands Severely water-stressed uplands Low-fertility uplands
�0.10 0.41**
�0.04 0.57**
0.53** 0.50**
0.27
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–52 47
the cultivar effect into two components: cultivar types and
cultivars within types.
2.2. Performance of improved upland rice varieties
under aerobic management in farmers’ fields in southern
Yunnan, China
Variety trials were conducted on-farm by Yunnan
Academy of Agricultural Sciences research staff in southern
Yunnan, China, to compare improved upland tropical
japonica cultivars (IRAT 104, Yunlu 29, and Yunlu 52)
and the improved indica B6144-MR-6-0-0 with a traditional
upland tropical japonica landrace (Mengwanggu) under
high-input management. Trials were conducted under
upland conditions on rainfed terraces, in communities that
formerly relied upon upland rice production for food
security, but which now grow intensively managed aerobic
rice in rotation with other upland crops. Experiments were
seeded at a rate of 150 kg ha�1. Fertility management
consisted of a basal application of 15 kg ha�1 urea,
15 kg ha�1 zinc sulfate, and 300 kg ha�1 calcium magne-
sium phosphate, plus a top-dressing of 135 kg ha�1 urea at
the panicle initiation stage, for a total N application of
approximately 70 kg ha�1. Three-replicate trials with
harvested plot areas of 13.3 m2 were conducted annually
at 11 sites between 1993 and 2002. The 11 sites are at
elevations ranging from 830 to 1680 masl. They include four
low-elevation (�1250 masl) sites (Jinghong, Menglian,
Ximeng, and Mengla) at which both indica and japonica
cultivars can perform well, and seven short-season, higher-
elevation (>1250 masl) sites (Wengshan, Pingbian, Simao,
Menghai, Langcan, Genma, and Cangyuan) at which
japonica cultivars are usually favored. Means were
estimated from a combined residual maximum likelihood
(REML) analysis over years, using the REML algorithm of
the SAS MIXED procedure (SAS Institute, 1992), with
cultivars and elevation (low versus high) classes considered
fixed, and trials within elevation classes considered random.
Within the 1993–2002 period, each new line was subjected
to a testing cycle of at least 2 years. Mengwanggu was
common to all sites in all years. For the analysis, cultivars
were considered fixed.
2.3. Performance of improved upland rice varieties
under aerobic management in farmers’ fields in
Batangas, Philippines
Experiments were conducted in the province of Batangas,
southern Luzon, Philippines to compare the productivity of
improved indica and traditional tropical japonica varieties
under high-input upland management in farmers’ fields.
This region is characterized by light-textured upland soils of
volcanic origin. Farmers in the region grow a mix of
traditional and improved upland varieties in a double-
cropping system, in which rice is planted at the onset of the
rainy season (usually in May) followed in October by a
maize, sweet potato, or vegetable crop that exploits the late
rains. Varieties were evaluated using both researcher-
managed, replicated on-farm ‘‘Mother’’ trials and farmer-
managed ‘‘Baby’’ trials. In the wet season of 2002, two
traditional Philippine upland varieties (Dinorado and
Azucena) and two high-yielding improved upland rice
cultivars (IR55423-01 and IR74371-46-1-1) were grown in
farmers’ fields in two-replicate researcher-managed trials at
three sites near Lemery, Batangas, as part of a larger
experiment evaluating 15 varieties. A seeding rate of
60 kg ha�1 was used, with 90 kg N ha�1 applied in three
splits in the form of urea. The same varieties were also
screened in farmer-managed trials, in which farms were
treated as incomplete blocks. Farmers received 2 kg of seed
of each of three varieties, and were asked to plant and
manage them as they did their own crop. Farmer fertilizer
application rates are not precisely known, but ranged from
60 to 120 kg N ha�1. In those experiments, seed was
broadcast in May, shortly after the first monsoon rains, at a
rate of about 150 kg ha�1 on furrowed fields, then covered
by harrowing. In both researcher-managed and farmer-
managed trials, rows were plowed into the stand after
emergence. Weeds were managed via inter-row cultivation
using animal traction. At harvest, 5 m2 crop cuts were
harvested and yield determined for each of the introduced
varieties and for the farmer’s field adjacent to the Baby trial.
Cultivar means from both researcher- and farmer-managed
trials were estimated from a combined residual maximum
likelihood (REML) analysis over years, using the REML
algorithm of the SAS MIXED procedure (SAS Institute,
1992), with cultivars considered fixed and farms random.
3. Results
3.1. Performance of upland, lowland, and aerobic
varieties in five management regimes in southern Luzon,
Philippines
3.1.1. Correlations across environment types
Yields were determined for 44 diverse cultivars (Table 1)
in five environment types (Table 2) for at least two seasons.
Mean yields across cultivars for environments NSL, NSU,
MSU, LFU, and SSU were 2.76, 2.23, 1.05, 1.00, and
0.46 t ha�1, respectively. Average yields in all environments
were relatively low due to the large number of upland
varieties with poor yield potential under favorable condi-
tions. Correlations of variety mean yields across these
hydrological regimes are presented in Table 3. Cultivar
means in the NSL environment were highly correlated with
those from the NSU environment, indicating that similar
cultivars with high-yield potential were favored in both
lowland and near-saturated upland environments. Correla-
tions between yields in the well-watered NSL and NSU
environments and the MSU environment were positive but
considerably lower, indicating that selection in well-watered
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–5248
Table 4
Subset of cultivars used in analysis of agronomic performance of cultivar
types in southern Luzon 2000–2003
Cultivar type Cultivar
Irrigated lowland high-yield
varieties (HYV)
IR64
IR72
PSBRC 82
Improved upland-adapted IR55423-01
environments is not adequate to identify genotypes that
perform well under moderate stress. Genotypic effects from
the severely stressed SSU environment were correlated with
those from the moderately stressed dry-season environment,
but were not positively associated with those estimated in
the irrigated lowland and aerobic wet season environments.
These results indicate that, in hydrological terms, there are
three broad target environments for upland rice breeding:
indica varieties selected forhigh-fertility environments (IHF)
IR55419-04
UPL RI-7
(1) eTabl
Mea
japo
in fiv
Varie
HYV
IHF
JLF
EDT
JTV
L.S.D
Improved upland tropical
japonica varieties selected for
low-fertility environments (JLF)
IR60080-46A
UPL RI-5
IR47686-30-3-2
nvironments in which there is little water stress, with
soil remaining near saturation throughout the growing
season, with soil moisture tension in the upper 15 cm of
soil never falling below �0.03 MPa;
Improved early maturingvarieties selected for
Vandana
IR70358-84-1-1
(2) mdrought tolerance (EDT) WAB 56-125
Traditional Philippine
upland tropical japonica
varieties (JTV)
Azucena
Palawan
Dinorado
oderate-stress environments in which soil moisture
falls below field capacity, either regularly or during the
critical reproductive period, with soil moisture tension
in the upper 15 cm layer often falling below�0.03 MPa,
reducing yields by 30–50% relative to saturated soil
environments;
(3) s
evere-stress environment where soil is rarely saturatedbetween irrigations or rainfall events, and in which soil
moisture tension frequently falls to�0.03 MPa at 30 cm
depths, resulting in yield reductions of 70% or more
relative to saturated soil environments.
The positive correlation between yields in non-stressed
and moderately stressed environments indicates that it is
possible to combine high-yield potential with tolerance to
moderate levels of water stress, and that the most effective
breeding approach for aerobic rice production systems that
are likely to be subject to intermittent stress is to screen in
both non-stressed and moderately stressed environments,
selecting on the basis of mean yield across these regimes.
Correlations between line means at the low-pH LFU site
at Siniloan and the more fertile NSU site at Los Banos were
positive and significant. These results indicate that
responsive genotypes performing well in favorable environ-
ments can also produce relatively high yields in unfavorable
environments. Selection for yield potential under favorable
conditions, required for gains in intensively managed
aerobic rice systems, is therefore, also likely to produce
yield gains in less fertile environments.
e 5
n yield (t ha�1) of irrigated high-yield varieties (HYV), improved indica upl
nica varieties (JTV), and improved tropical japonica types selected for low-fert
e environment types in southern Luzon, 2000–2003
ty type Environment type
Non-stressed
lowland
Non-stressed
upland
Modera
upland
4.04 2.12 0.84
3.62 3.56 1.47
3.31 2.89 1.10
1.71 1.44 1.11
2.29 1.63 0.81
.0.05 0.82 0.47 0.30
3.1.2. Performance of cultivar groups
Among the varieties listed in Table 1 were representatives
of distinct cultivar groups. Table 4 lists the subset of three
cultivars comprising the sample for each group (HYV, IHF,
JTV, JLF, and EDT) Mean yields of variety groups within
environments are presented in Table 5. The HYV types
outyielded all others under lowland conditions, although the
difference was not significant relative to the IHF and JLF
groups. In the high-fertility, well-watered IRRI wet season
environment, IHF varieties significantly outyielded all other
types, surpassing the HYV types by nearly 70%. The IHF
types also outyielded all others under conditions of moderate
stress in the dry season at IRRI and under low-fertility, acid-
soil conditions at Siniloan. The responsiveness of IHF
varieties to improved fertility conditions is illustrated by the
contrast in their performance in the NSU and LFU
environments; when grown in high-fertility (NSU) environ-
ments at IRRI, the improved indica upland types yielded
2.30 t ha�1 more than when grown in the low-fertility (LFU)
Siniloan environment. The equivalent responses were only
1.75, 0.60, and 0.87 t ha�1 for the JLF, EDT, and JTV
cultivar groups, respectively. The advantage of the EDT
and types selected in high-fertility environments (IHF), traditional tropical
ility and drought-prone environments (JLF and EDT, respectively) evaluated
tely water-stressed Severely water-stressed
upland
Infertile
upland
0.47 0.91
0.60 1.26
0.31 1.14
1.00 0.84
0.26 0.76
0.30 0.38
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–52 49
types was evident only under conditions of extreme stress,
when they significantly outyielded all other cultivar groups.
They were the lowest-yielding cultivar group in well-
watered environments. The JTV landraces were the lowest-
yielding under conditions of water stress and low fertility. In
general, the aerobic-adapted IHF group combined respon-
siveness to favorable conditions with a high degree of
tolerance to both water stress and low fertility.
The superiority of aerobic-adapted improved indica
cultivars under favorable and moderately stressful upland
conditions appears to be a function of their higher harvest
index (HI) in these environments relative to other cultivar
types (Table 6). Both IHF andHYVvariety types achievedHI
of nearly 50% in irrigated lowland environments, but theHI of
HYV declined precipitously in all upland environments,
whereas IHF varieties retained a relatively high HI. JLF
varieties targeted at low-input systems usually producedmore
biomass than aerobic rice varieties (Table 7), but partitioned
much less drymatter to grain.Only in themost severelywater-
stressed environment was the HI of IHF varieties exceeded by
that of the drought-tolerant EDT group. Drought-tolerant
tropical japonica varieties retained much higher levels of
grain-set under extreme stress than other types, achieving HI
of over 30%. However, their low biomass production in all
environments resulted in much reduced yields relative to the
IHF group in all other environments.
Although the drought-tolerant japonica varieties are
much shorter in duration than other cultivar classes,
Table 7
Mean biomass production (t ha�1) of irrigated high-yield varieties (HYV), imp
traditional tropical japonica varieties (JTV), and improved tropical japonica type
respectively) evaluated in five environment types in southern Luzon, 2000–2003
Variety type Environment type
Irrigated
lowland
Non-stressed
upland
Moderate
upland
HYV 8.59 7.96 7.15
IHF 7.57 9.59 5.92
JLF 8.48 9.28 5.93
EDT 4.41 6.37 4.25
JTV 6.89 7.44 6.46
L.S.D.0.05 1.63 1.37 2.00
Table 6
Mean harvest index of irrigated high-yield varieties (HYV), improved indica upl
japonica varieties (JTV), and improved tropical japonica types selected for low-fert
in five environment types in southern Luzon, 2000–2003
Variety type Environment type
Irrigated
lowland
Non-stressed
upland
Moderate
upland
HYV 0.47 0.27 0.21
IHF 0.48 0.37 0.28
JLF 0.39 0.31 0.21
EDT 0.39 0.25 0.29
JTV 0.34 0.22 0.16
L.S.D.0.05 0.07 0.05 0.03
flowering in 64 days under stress, their improved harvest
index and yield under severe stress appears to result from
drought tolerance rather than avoidance. In the MSU and
SSU environments, stress was imposed at least 15 days
before the earliest varieties flowered, causing a reduction in
biomass production of over 30% relative to the favorable
NSU environment. This reduction is proportionately less for
the EDT types than for other variety types but is not
consistent with the hypothesis that the drought-tolerant class
merely escaped stress at flowering. The mechanism of
tolerance may be related to a flowering program in which
peduncle elongation and floret opening occur in spite of
stress. Drought-tolerant varieties exhibited a slight reduction
in days to flower under severe stress relative to the irrigated
lowland environment, whereas for all other cultivar classes,
flowering was delayed by 2 weeks or more as a result of
severe stress (Table 8).
Other features of the aerobic rice cultivars studied in this
experiment include intermediate plant height and panicle
number under favorable upland conditions relative to
irrigated and improved upland varieties. The IHF varieties
had a mean height of approximately 110 cm (data not
shown) compared to means of 93 cm and 126 cm for HYV
and JLF varieties, respectively. The semi-dwarf stature of
irrigated HYVs appears to be detrimental to aerobic
adaptation, but the tall, weak-strawed improved JLF types
are not sufficiently lodging-resistant to support the relatively
high levels of N application needed to achieve high-aerobic
roved indica upland types selected in high-fertility environments (IHF),
s selected for low-fertility and drought-prone environments (JLF and EDT,
ly water-stressed Severely water-stressed
upland
Infertile
upland
3.24 4.15
3.63 4.55
2.24 4.88
3.31 3.05
3.23 3.45
1.54 0.97
and types selected in high-fertility environments (IHF), traditional tropical
ility and drought-prone environments (JLF and EDT, respectively) evaluated
ly water-stressed Severely water-stressed
upland
Infertile
upland
0.17 0.25
0.15 0.28
0.10 0.25
0.32 0.23
0.09 0.20
0.08 0.09
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–5250
Table 9
Mean number of productive tillers m�2 of irrigated high-yield varieties (HYV), improved indica upland types selected in high-fertility environments (IHF),
traditional tropical japonica varieties (JTV), and improved tropical japonica types selected for low-fertility and drought-prone environments (JLF and EDT,
respectively) evaluated in five environment types in southern Luzon, 2000–2003
Variety type Environment type
Irrigated
lowland
Non-stressed
uplands
Moderately water-stressed
uplands
Severely water-stressed
uplands
Infertile
uplands
HYV 354 465 576 428 N/A
IHF 237 318 388 353 183
JLF 255 319 389 276 222
EDT 265 327 367 389 236
JTV 182 213 251 186 99
L.S.D.0.05 63 51 36 59 204
Table 10
Grain yield (t ha�1) of upland and aerobic rice cultivars under high-input
management at short or long-season locations in Yunnan, 1993–2002
Cultivar Type Grain yield (t ha�1)
Short-season Long-season
Mengwanggu Traditional japonica 2.57 � 0.10 2.92 � 0.12
IRAT 104 Improved japonica 2.37 � 0.26 3.37 � 0.39
B6144F-MR-6-0-0 Improved indica 1.70 � 0.17 4.13 � 0.20
Yunlu 29 Improved japonica 3.34 � 0.20 3.29 � 0.24
Yunlu 52 Improved japonica 2.86 � 0.23 4.00 � 0.28
Table 8
Mean days to flower of irrigated high-yield varieties (HYV), improved indica upland types selected in high-fertility environments (IHF), traditional tropical
japonica varieties (JTV), and improved tropical japonica types selected for low-fertility and drought-prone environments (JLF and EDT, respectively) evaluated
in irrigated lowland and severely stressed upland environments at IRRI, 2000–2003
Variety type Days to flower in
irrigated lowland
Days to flower in severely
water-stressed upland
Flowering delay
due to stress
HYV 82.6 98.1 15.5
IHF 80.7 94.3 13.6
JLF 90.1 108.0 17.9
EDT 68.8 64.4 �4.4
JTV 90.6 106.0 15.4
rice yields. The panicle number of all improved upland
variety types was significantly higher than those of JTV
types, but lower than that of irrigated lowland types
(Table 9). IHF and JLF varieties produced 100–200 fewer
panicles m�2 than HYVs in all environments. The high-
yielding Brazilian aerobic cultivars are also characterized by
moderate tillering (250 tillers m�2; Pinheiro and de Castro,
2000). However, increased panicle number may be an
important component of higher harvest index and yield
potential under aerobic management. High panicle number
is a feature that distinguishes the high-yielding semi-dwarf
hybrid Magat, which can produce over 700 panicles m�2 in
aerobic experiments (George et al., 2002). Selection for
increased panicle number appears to be a promising avenue
for increasing aerobic rice grain yield.
In general, these results indicate that the main feature
distinguishing improved aerobic-adapted rice varieties of the
IHF type from other cultivar types is their ability to retain
both high-biomass production and relatively high-harvest
index under favorable and moderately stressful upland
conditions. Irrigated rice varieties produced less biomass
and partitioned less of it to grain under upland management.
Traditional tropical japonicas and improved JLF varieties
produced as much or more biomass as aerobic varieties
under upland conditions, but partitioned much less to grain.
The high harvest index of IHF varieties may result from a
lower sensitivity to reproductive-stage water stress than all
but the most drought-tolerant upland materials, without
sacrifice of the biomass production needed for high-yield
potential.
3.2. Performance of aerobic rice varieties over sites and
years in southern Yunnan, China, and in Batangas,
Philippines
Cultivar means for the experiments conducted in Yunnan
are presented in Table 10. Means over all cultivars, seasons,
and locations were 3.36 t ha�1 for the longer season sites,
and 2.72 t ha�1 for the shorter-season sites. F-tests of both
cultivar and cultivar � elevation group interaction were
highly significant (data not shown), indicating that different
cultivars were favored in the two types of environment.
Yunlu 29 was the highest-yielding cultivar in the short-
season, high-elevation environments. With a mean yield of
3.34 t ha�1 in these environments, it outyielded the
traditional check by about 30%. B6144F-MR-6-0-0, an
improved indica introduction from Indonesia, and Yunlu 52,
a locally bred improved tropical japonica upland rice, were
the highest-yielding cultivars in longer season environments.
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–52 51
Table 11
Mean grain yield for aerobic and traditional upland rice varieties grown under farmer and researcher management in a traditional rice–maize double-cropping
system in Batangas Province, Philippines, 2002
Cultivar Cultivar
type
No. of farmer-
managed
trials
Grain yield
under farmer
management (t ha�1)
Yield under
researcher
management (t ha�1)
Duration in
researcher-managed
trials (days)
IR55423-01 Aerobic 16 3.41 � 0.26 3.90 123
IR74371-46-1-1 Aerobic 8 3.42 � 0.33 3.81 112
Azucena Traditional 18 2.04 � 0.25 2.15 125
Dinorado Traditional 21 2.28 � 0.24 2.04 131
Farmers’ variety Mixed 58 3.14 � 0.19 N/A N/A
L.S.D.0.05 0.62 1
With mean yields of over 4.0 t ha�1, they outyielded the
traditional check by about 40%.
Means for yield and crop duration in on-farm upland rice
trials in Batangas are presented in Table 11. Mean yields of
traditional tropical japonica varieties averaged over 3 t ha�1
for 66 reporting farms. The improved indica varieties, with
mean yields of approximately 3.4 t ha�1, outyielded the
traditional tropical japonica varieties by over 50% under
farmer management, and by nearly 100% in researcher-
managed trials. They also outyielded the mixture of
traditional and improved varieties used by farmers, and
were preferred because of their reduced duration and
improved lodging resistance.
4. Discussion
4.1. Key attributes of aerobic cultivars, and
implications for aerobic rice breeding
Rice cultivars for aerobic systems need to combine high-
biomass production, harvest index, and lodging resistance
with moderate drought tolerance, particularly at the highly
sensitive reproductive stage (Garrity and O’Toole, 1994).
Because aerobic systems depend on direct seeding in dry
soil, without accumulation of standingwater, vigorous early
growth is also needed to compete with weeds and to root
deeply to avoid early season drought. The results of this
study indicate that, for the Asian tropics, these traits are
most likely to be found in varieties that are primarily of
indica germplasm. Traditional tropical japonica varieties
and improved varieties derived predominantly from them
are low-tillering and have low HI; they are, therefore,
unlikely to have the yield potential or early vegetative vigor
needed for intensively managed upland systems. High-
yielding cultivars that have emerged from Asian breeding
programs for the favorable uplands are often rather closely
related to elite tropical irrigated varieties, usually through
IR8 and its relatives. For example, examination of the
pedigree of the upland rice variety IR55423-01, one of the
highest-yielding pure-line varieties under our aerobic
management trials, shows that its coefficient of coparentage
of with IR8 is only slightly less than that of the irrigated
varieties IR64 and IR72 (Atlin and Lafitte, 2002). IR55423-
01 does not, however, appears to carry the sd-1 allele from
its irrigated parents, and is of intermediate height. Varieties
like IR55423-01, which is capable of maintaining HI under
moderate water deficit, should be evaluated more thor-
oughly in order to discover mechanisms that underlie this
ability.
Recent research has shown that drought tolerance and
high spikelet fertility under reproductive-stage stress in
aerobic-adapted germplasm are strongly associated with
short duration and minimal flowering delay under stress.
These are likely to be key attributes of upland-adapted
cultivars targeted at drought-prone or severely water-
restricted production environments (Lafitte and Courtois,
2002), and may be useful as screening criteria for drought
tolerance. However, the most efficient approach to the
identification of cultivars adapted to any target environ-
ment is usually to screen directly for performance in that
environment, unless the repeatability (H) of grain yield
estimates in the target system is low (Atlin, 2001). In rice,
H of grain yield estimates in variety trials does not appear
to be much affected by soil water status per se (Atlin and
Lafitte, 2002), although uneven application of irrigation
water to the experimental field can greatly increase error
variances and reduce precision. Direct selection under
high-input aerobic management, is therefore, likely to be
required to optimize performance under aerobic manage-
ment. Because the results of this study indicate that yield in
well-watered environments is not highly correlated with
yield under moderate stress, early screening of breeding
lines under both high-fertility aerobic management in the
wet season and, where possible, moderate water stress in
the dry season appears to be the best approach to
developing cultivars combining high-yield potential and
moderate stress tolerance.
4.2. Implications for aerobic rice target environments
The Yunnan and Batangas results confirm that cultivars
suited to aerobic management systems using moderate N
fertilization rates are currently available, both for hilly areas
G.N. Atlin et al. / Field Crops Research 97 (2006) 43–5252
of southeast Asia that have traditionally relied on shifting
cultivation, and for permanent arable upland cropping
systems of the type found in many parts of south and
southeast Asia, where rice is rotated annually or double-
cropped with other upland crops. About 100,000 ha of
upland rice are grown annually in southern Yunnan, and the
system in Batangas covers several thousand hectares.
Currently available improved upland/aerobic rice varieties
can consistently deliver wet-season yields of between 3
and 4 t ha�1 without supplemental irrigation in these
systems. Such cultivars are the basis for more productive
and sustainable upland cropping systems that are evolving
in several parts of southeast Asia. The Batangas results,
which were generated in farmers’ fields in an indigenous
cropping system in which upland rice is grown once a year,
with one intervening non-rice crop, show that high-aerobic
rice yields can be sustainable in annual double-crop
rotations.
The yields obtained with tropical aerobic varieties are
comparable to those obtained in favorable rainfed lowland
environments in south and southeast Asia, indicating that
aerobic rice-based production systems are likely to be
attractive to farmers in drought-prone rainfed lowlands as
well as in uplands. In drought-prone lowland rice-producing
areas like the Bastar Plateau in Chhattisghar, India, or the
light-soil areas of northeastern Thailand and southern Laos,
yields average less than 3 t ha�1 and drought risk is high. In
these areas, failure to accumulate standing water in bunded
fields can delay transplanting (Rickman et al., 2001) or may
severely disrupt water-dependent weed control operations,
such as beusani (Tomar, 2002). Dry direct-seeded aerobic
rice systems, which do not require standing water to
accumulate either for establishment or weed management,
result in early establishment and more reliable weed control,
improving the synchrony between crop development and
seasonal rainfall. Weed pressure is a major constraint to the
adoption of direct seeding with conventional lowland
germplasm. However, upland-adapted indica rice varieties
tend to be moreweed-competitive than either lowland HYVs
or tropical japonica types (unpublished data), and may
facilitate the adoption of risk-avoiding direct-seeding
technology.
5. Conclusion
Medium-height, aerobic-adapted cultivars consistently
yielding approximately 4 t ha�1 under farmer management
in rainfed conditions are now available to rice growers in the
Asian tropics. These cultivars, which are usually of the
indica type, combine a moderate level of drought tolerance
with input responsiveness. They could permit the replace-
ment of shifting cultivation systems with more productive,
more sustainable, and less extensive systems in southeast
Asia, and may permit rainfed lowland rice growers in
drought-prone areas to adopt risk-avoiding direct-seeding
systems. Mechanisms underlying the high-harvest index of
these cultivars under moderate water stress require further
study, but breeding programs focusing on screening
improved, indica-derived germplasm under both favorable
and water-short conditions are making rapid progress in
developing a new class of rice cultivar for water-short
environments.
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