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Transgressive segregation for yield traits in Oryza sativa IR58025B X Oryza

meridionalis Ng. Bc2F3population under irrigated and aerobic conditions

Article · September 2012

DOI: 10.1007/s12892-012-0006-1

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J. Crop Sci. Biotech. 2012 (September) 15 (3) : 231 ~ 238
DOI No. 10.1007/s12892-012-0006-1

RESEARCH ARTICLE

Transgressive Segregation for Yield Traits in Oryza sativa

IR58025B X Oryza meridionalis Ng. Bc F Population under 2 3

Irrigated and Aerobic Conditions

Chejerla Mohan Kumar Varma1*, Patil Kalmeshwer Gouda1,2, Surapaneni Saikumar1, Vinay Shenoy1, Halagappa Eshwarappa
Shashidhar1,3, Sarla Neelamraju 4

1
Barwale Foundation, Barwale Chambers, #3-6-666, Street No. 10, Himayathnagar, Hyderabad 500 029, India
2
Monsanto, Bengaluru, India
3
Department of Biotechnology, University of Agricultural Sciences, GKVK, Bangalore 560 065, India
4
Department of Biotechnology, Directorate of Rice Research, Rajendranagar, Hyderabad 500 030, India

Received: February 1, 2012 / Revised: May 6, 2011 / Accepted: May 30, 2012
Ⓒ Korean Society of Crop Science and Springer 2012

Abstract
Wild species of the genus Oryza are a good source of beneficial alleles for enhancing rice yield under normal and adverse condi-
tions. BC2F3 population was derived from a cross between Oryza sativa IR58025B and Oryza meridionalis Ng. (2n = 24, AA) a heat
tolerant wild species to evaluate 12 yield traits under irrigated and aerobic conditions. Analysis of variance and genetic estimates
indicated there is substantial genetic variation among progenies under both conditions. Grain yield had high heritability (61.9%) and
genetic advance (36.4%) under irrigated conditions but moderate heritability (49.6%) and genetic advance (13.3%) under aerobic
conditions indicating that selection for yield will be effective under both conditions. Panicle number, grain number, spikelet fertility,
and test weight showed significant positive correlation with grain yield under both conditions. Families out-performing IR58025B
for yield under both conditions were obtained providing evidence that phenotypically inferior O. meridionalis contributed to yield
increase. This species can be a novel source of natural genetic variation for the improvement of rice under irrigated as well as under
aerobic condition.

Key words : advanced backcross population, aerobic condition, inter-specific cross, Oryza meridionalis, transgressive segregants,
yield related traits

Introduction
Rice (Oryza sativa L.) is a major cereal food crop and con-
sumed by nearly half of the world’s population. It constitutes for
35 - 75% of caloric uptake by Asians and planted over 11% of
global arable land (Khush 2005). Because of its agricultural
importance, rice has been bred intensively resulting in the dou-
bling of the production by adopting high-yielding
varieties/hybrids. However, this achievement has led to the nar-
rowing of the available genetic base in elite germplasm and much
C. Mohan Kumar Varma ( ) variability in cultivated rice is becoming increasingly obvious
Email: mohanvarma@barwalefoundation.org,
mohankumarvarma@gmail.com
concern about the reduced genetic gain in present day rice breed-
ing. In a genetic diversity study, Sun et al. (2001) reported that
cultivated rice has only 60% of the alleles of wild rice. A large
amount of genetic variation in the genus Oryza lies unexploited in
wild progenitors (Wang et al. 1992). Hence, attention is being
shifted to map yield-enhancing QTLs and enrich the cultivated
gene pool by introgression of favorable genes/gene complexes
from wild species (Tanksley and Mccouch 1997; Swamy and
Sarla 2008). The potential of wild relatives to increase the genetic
(Vaughan 1994). The increased variability occurs both by intro-
duction of the pre-existing wild species alleles and also a lot of

The Korean Society of Crop Science

232 Transgressive Segregation for Yield Using O. meridionalis

genetic variability is created de novo (Wang et al. 2005; wang et

al. 2010). Wild relatives were initially used to transfer disease and
insect resistance traits (Brar and Khush 1997; Dalmacio et al.
1995). Following the identification of yield QTLs yld 1.1 and yld
2.1 from Malaysian accession of O. rufipogon, each of which
could help increase yield by about 18% (Xiao et al. 1998), many
researchers exploited the wild species, O. rufipogon (Fu et al.
2010; He et al. 2006; Marri et al. 2005; Moncada et al. 2001;
Septiningsih et al. 2003; Tan et al. 2008; Thomson et al. 2003), O.
glaberrima (Aluko et al. 2004), O. glumaepatula (Brondani et al.
2002), O. grandiglumis (Yoon et al. 2006), and O. nivara
(Kaladhar et al. 2008; Li et al. 2006; Swamy and Sarla 2011) for
yield and grain quality traits.
Oryza meridionalis Ng. is an annual diploid (2n = 24) wild
species, endemic to northern Australia (Ng et al. 1981) and some
parts of Irian Jaya and Indonesia (Lu and Silitonga 1999).
Phylogenetic relationship studies among rice genomes based on
mitochondrial and chloroplast microsatellites (Nishikawa et al. Fig. 1. Flow chart showing the advanced backcross method adopted for the develop-
2005), MITE insertions (Zhu and Ge 2005), and SINE insertions ment of BC2F3 IR58025B/ O. meridionalis mapping population.
(Xu et al. 2005) revealed that it forms a separate lineage within
the genus Oryza. O. meridionalis shows high levels of genetic ent in inter-specific cross with O. meridionalis (IRGC101145), a
diversity among the geographically isolated accessions due to wild relative of northern Australia as the male parent. The popula-
selective pressure on isolated gene pools (Juliano et al. 2005). It tion was derived using the advanced backcross method (Tanksley
has high genomic affinity with O. sativa species as it shows high et al. 1996) which is a successful strategy and widely adopted in
bivalent formation and normal meiosis in inter-specific hybrids transferring favorable alleles from phenotypically poor
since it shares the AA genome (Lu et al. 1997, 1998). This species wild/weedy species into elite cultivars/lines. The breeding proce-
can be a good source for introducing new genes in expansion of dure adopted in the population development is shown in Fig. 1.
cultivated gene pool especially where water saving is required. Six F1 plants produced from the cross IR58025B X O. meridion-
alis were selected based on phenotype and confirmed genotypical-
Wild species of Oryza offers a good source for beneficial alle-
ly using polymorphic simple sequence repeat (SSR) markers
les and can be transferred into cultivars for enhancing yield even
between the parents. The F1s were backcrossed to IR58025B,
under stress conditions as well (McCouch et al. 2007; Price et al.
recurrent parent (RP) to derive 84 BC1F1 plants. In the BC1 gener-
2002). Earlier reports on O. meridionalis indicates that it has
ation plants with seed dormancy, sterility, and weedy characters
adaptation to arid climatic conditions (Second 1988), has drought
were rejected and 50 BC1F1 plants were used to backcross with
avoidance traits (Brar and Kush 1997; Somanthri 2001), and good
the recurrent parent to obtain BC2F1 seeds. Then 4 - 5 BC2F1 seeds
levels of tolerance to heat compared with O. sativa (Andrew et al.
from each BC1F1 plant were planted to generate 239 BC2F1 indi-
2009), yet it has not been exploited very much in rice breeding
viduals. All the plants were allowed to self to get BC2F2 seeds,
programs. Kalmesh et al. 2012 reported that O. meridionalis can
from each BC2F1 individual plant. A total of 239 BC2F2 families
be a good source for root traits contributing to moisture stress.
were advanced to BC2F3 for phenotyping without applying any
Recognizing the potential of wild species and to discover the
selection pressure.
unexplored variability in O. meridionalis, we generated an
advanced backcross population derived from O. sativa IR58025B
Field trial
X O. meridionalis comprising 239 BC2F3 progenies to evaluate 12
yield and yield-related traits under irrigated and aerobic condi- Evaluation of the mapping population along with recurrent par-
tions. ent was done under two moisture conditions, namely, irrigated
and aerobic, following a randomized block design with two repli-
cations during the wet season of 2009. In the irrigated field trial,
Materials and Methods all 239 BC2F3 families along with recurrent parent were sown in
nursery beds and transplanted to the field 24 days after germina-
tion. Each family and RP consisted of 30 plants in three rows of
Location 10 plants each with spacing of 20 cm between the plants and 30
The study was conducted at Barwale Foundation Farm, cm between the rows. After transplanting, approximately 5 - 7 cm
Maharajpet, Hyderabad, located at latitude of 17º 24’ N and longi- of standing water was maintained in the field until draining before
tude of 78º 12’ E, and a altitude of 536 m above mean sea level. harvest. Basal fertilizer was applied at the rate of 50-40-40 kg
NPK ha-1, and were top-dressed twice with 30 kg N ha-1 at 2 and 6
Parent material and population development weeks after planting. Weeds were controlled by application of
IR58025B, a popular maintainer line, was used as female par- post-emergence herbicide Butaclor (3 L ha-1). For the control of
 JCSB 2012 (September) 15 (3) : 231 ~ 238 233

leaf folder attack, insecticide monocrotophos was sprayed at the Statistical analysis
rate of 1.5 L ha-1. Phenotypic data generated from both irrigated and aerobic con-
In the aerobic field trial, the same 239 BC2F3 families were dition was analyzed without any transformation. Data analyses
directly sown into level, unpuddled, unflooded upland fields. Each were performed in SAS (SAS Institute. Cary, NC) considering all
family and recurrent parent was sown in three rows of 10 plants factors random. Analyses of variance (ANOVA) was performed
each with a spacing of 15 cm between the plants and 30 cm using the general linear modeling (Proc GLM). The variance com-
between the rows. The soil was amended with zinc sulphate at the ponents used for the estimation of heritability (h2) of each trait
rate of 2.5 kg ha-1 before sowing. NPK fertilizer was applied at the were obtained accord to (Hanson et al. 1956) variance compo-
rate of 100-40-40 kg ha-1. N was applied in three equal splits at nents procedure (PROC VARCOMP) using restricted maximum
sowing, 40 days after sowing (DAS), and 60 DAS. Weeds were likelihood (REML) method while the genetic advance as % of
controlled by applying post-emergence herbicide Butaclor (3 L ha- mean was estimated as suggested by Falconer and Mackay
1
), 7 DAS followed by manual hand weeding at later stages. (1996). Trait correlations were studied with n-2 degrees of free-
Irrigation was given once in 4 days to maintain soil near field dom at 0.5% and 0.1% level of significance using SPAR 2.0
capacity till the germination of seedling to three-leaf stage. (Sangeeta et al. 2008).
Drought stress was imposed to the trial when crop reached flower-
ing by reducing the frequency of irrigations to once in 10 - 12
days from 7 weeks after sowing until harvest of the crop. Results
Tensiometers were installed in the field, and soil water status was
monitored. Irrigation was withheld until soil water tension Mean values of recurrent parent and population for yield and
reached about -40 kPa at 30 cm soil depth. The trial was then irri- yield-related traits studied under irrigated and aerobic conditions
gated until the soil was saturated in the root zone. The total are presented in Table 1. The mean of the recurrent parent and
amount of rainfall received during the 2009 crop growth was 520 population was slightly higher under irrigated conditions com-
mm, while the estimated evapotranspiration was 700 mm. pared to aerobic conditions for most of the traits except for BM.
The GY of the recurrent parent was 20 g under irrigated condi-
Phenotypic evaluation tions and 5.2 g under aerobic conditions with relative yield
Five plants were selected from the center at random from each decrease of 74.5% under aerobic conditions compared to irrigated.
of the 239 BC2F3 families in each experiment and evaluated for A similar kind of reduction in mean values of all the traits except
the following 12 yield-related traits: plant height (PH) measured for TN and BM under aerobic conditions was noticed in the
in centimeters (cm) from the soil surface to the tip of the tallest BC2F3 population with the highest reduction for SPY (69.5%) fol-
panicle (awns excluded). Days to 50% flowering (DF) evaluated lowed by GN (45.3%) and PH (41.4%). However, several positive
as the number of days taken by each entry, from sowing to open- transgressive segregants out-performing the recurrent parent were
ing of first flower in 50% of the plants. Tiller number (TN) the observed for yield component traits under irrigated and aerobic
total number of tillers per plant. Panicle number (PN) recorded as conditions as shown in Fig. 2. The maximum number of trans-
total number of panicle bearing tillers in each plant at the time of gressive segregants at 0.05% CD was observed for GY (52) fol-
harvest. Panicle length (PL) measured in centimeters (cm) from lowed by PL (49), SF (48), and GN (28) under irrigated condi-
the panicle neck to the panicle tip (excluding the awn). Grain tions, whereas under aerobic conditions maximum transgressive
number (GN) calculated by counting number of filled spikelets Table 1. Mean values of IR58025B (recurrent parent RP) and BC2F3 popula-
per panicle averaged over five randomly chosen panicles in each tion of IR58025B/ O. meridionalis for yield and related traits under irrigated
plant. Spikelet number (SN) per panicle were calculated as total (I) and aerobic (A) conditions
number of spikelets including empty and filled ones averaged RP Mean Population mean F- value
over five randomly chosen panicles in each plant. Spikelet fertility Trait
(I) (A) (I) (A) (I) (A)
(SF) was calculated as ratio of filled spikelets to the total number
PH 105.3 62.0 114.3 67 4.5** 1.3*
of filled and unfilled spikelets per panicle expressed in percent- DF 100.0 - 103 - 2.9** -
age. Test weight (TW) was measured as weight of 1,000 random- TN 18.2 16.0 20 21 2.5** 4.0**
ly selected seeds and expressed in grams averaged over five sam- PN 17.6 9.0 19 12 2.1** 4.8**
ples taken from the bulk harvested grains from each plant. PL 24.5 19.0 25.6 18 2.6** 1.6**
Biomass (BM) was calculated as the average weight of the five GN 120.5 70.0 133.5 75.8 2.3** 4.2**
SN 180.0 126.0 194.4 132 2.0** 2.2**
well-dried plants whose panicles had been removed. Grain yield SF 67.0 55.5 68.7 58 2.1** 4.4**
(GY) recorded as total weight of all filled grains of a single plant TW 19.2 12.0 21.1 13 1.9** 2.2**
averaged over five randomly chosen plants in each family was BM 35.0 42.0 32 47 2.1** 3.9**
recorded in grams (g). Plot yield (PY) was taken as the weight of GY 20.5 5.2 23.3 7.1 4.2** 6.9**
dried and cleaned bulked harvested grains from all the 30 plants in PY 2.7 - 3.1 - 3.3** -
a plot was taken in grams and extrapolated to tons per ha. Traits Significant at p= 0.05* and p = 0.01**
PH: plant height (cm), DF: days to 50% flowering, TN: tiller number, PN: panicle
DFF and PY were not recorded in the aerobic field trial experi- number, PL: panicle length (cm), GN: grain number, SN: spikelet number, SF: spikelet
ment. fertility (%), TW: 1000 seed weight (g), BM: biomass (g), GY: grain yield (g), PY: plot
yield (t ha-1)
234 Transgressive Segregation for Yield Using O. meridionalis

Table 2. Estimates of GCV, PCV, h2, and GA for yield and related traits in
BC2F3 population of IR58025B/ O. meridionalis under irrigated (I) and aero-
bic (A) conditions
GCV PCV h2 GA
Trait
(I) (A) (I) (A) (I) (A) (I) (A)
PH 11.4 5.5 13.8 14.4 68.3 14.4 19.4 4.3
DF 4.2 - 6.6 - 40.5 - 5.5 -
TN 15.3 18.1 23.3 23.3 43.0 60.4 20.7 29.0
PN 14.1 23.7 23.5 29.3 36.0 65.6 17.4 39.6
PL 6.2 5.4 8.6 11.2 52.4 23.5 9.3 5.4
GN 16.4 21.5 25.9 27.4 39.8 61.4 21.3 34.6
SN 11.3 13.3 20.8 21.5 29.7 38.3 12.7 17.0
SF 8.0 17.2 16.2 21.6 24.4 63.2 8.2 28.1
TW 5.8 10.4 11.6 16.7 25.0 38.2 6.0 13.2
BM 15.4 22.9 25.6 29.8 35.8 59.3 18.9 36.4
Fig. 2. Transgressive segregants obtained in the mapping population for yield compo-
GY 22.5 36.8 28.6 52.2 61.9 49.6 36.4 13.3
nent traits under irrigated and aerobic conditions during WS 2009.
PY 25.0 - 34.2 - 53.4 - 37.7 -
PH: plant height (cm), DF: days to 50% flowering, TN: tiller number, PN: panicle
number, PL: panicle length (cm), GN: grain number, SN: spikelet number, SF: spikelet
fertility (%), TW: 1000 seed weight (g), BM: biomass (g), GY: grain yield (g), PY: plot
yield (t ha-1), GCV: genotypic coefficient of variation (%), PCV: phenotypic coefficient
of variation (%), h2: broadsense heritability (%), GA: genetic advance as % of mean

Genetic parameters, viz., GCV, PCV, h2, and GA estimated for

Fig. 3. Comparision of mean grain yield of the population and recurrent parent
(IR58025B) under irrigated (I) and aerobic conditions (A).
Fig. 4. Box plots of yield and related traits in BC2F3 population under irrigated (A) and
aerobic (B) conditions.
segregants were noticed for GY (66) followed by SF (39), GN
(35), and TW (27) (Fig. 2.). No positive phenotypic transgressive
variants were observed for TN and PN as O. meridionalis itself
had a significantly high number of TN and PN under both the
conditions evaluated. Comparison of mean grain yield data of the
population and RP under aerobic and irrigated conditions is pre-
sented in the form of an interaction plot in Fig. 3 and it shows the
GY of the population is significantly higher than the RP under
both conditions. There was substantial variation in the population
for all the traits under both conditions as indicated by the results
of ANOVA (Table 1) as well as range values of all the traits pre-
sented in the form of box plots (Fig. 4). All the traits studied fol-
lowed a normal distribution under both conditions (Fig. 5).
all traits under irrigated and aerobic conditions are presented in
Table 2. PY (25.0 and 34.2%) had the maximum GCV and PCV
values followed by GY (22.5 and 28.6%) and GN (16.4 and
25.9%) under irrigated condition, whereas, under aerobic condi-
tion GY (36.79 and 52.24%) had the maximum values followed
by PN (23.7 and 29.3%) and BM (22.9 and 29.7%). The differ-
ence between the GCV and PCV values for most of the traits stud-
ied under both conditions was significant, but difference was
greater under aerobic conditions. High to moderate h2 and GA val-
ues were obtained for GY (61.9 and 36.4%), PY (53.4 and
37.7%), PHT (68.28 and 19.4%), and GN (39.8 and 21.3%) under
irrigated condition, whereas under aerobic condition PN (65.62
and 39.58%), SF (63.2 and 28.1%), and GN (61.4 and 34.6%)
recorded high to moderate values. As expected, the h2 value of
GY was high (61.9%) under irrigated conditions compared to aer-
obic conditions (49.5%).
Correlation values among yield and yield-attributing traits
under both the conditions are summarized in Table 3. GY showed
positive correlation with all the yield-related traits under both con-
ditions. A significant positive correlation was observed for GY
with GN (0.46), followed by SN (0.36), TW (0.32), and PL (0.25)
under irrigated conditions. Under aerobic conditions, PN (0.50)
had the highest positive correlation followed by GN (0.47) and SF
(0.37). Among yield-related traits under both PHT and SN, PL
and GN, SF and GN showed a significantly high positive correla-
tion. All the morphological traits had a significant positive associ-
ation with BM under both conditions. However, the association
pattern differed for the traits such as PN with PH, PL with TN,
and GN with TN across the conditions.
Discussion
Genetic variation observed in the advanced backcross progenies
of the inter-specific cross for yield and yield components under
 JCSB 2012 (September) 15 (3) : 231 ~ 238 235

Fig. 5. Frequency distribution for yield traits in IR58025B/ O. meridionalis mapping population under irrigated and aerobic conditions. Arrow indicates the value of the recurrent par-
ent IR58025B.

irrigated and aerobic conditions suggest evidence for the contribu- spite of its phenotypically inferior performance, wild species (here
tion of favorable alleles from low-yielding, agronomically inferior O. meridionalis) harbor trait-enhancing alleles that can increase the
O. meridionalis for yield under well-watered conditions as well as yield of modern cultivars under different ecological conditions.
under aerobic conditions. This confirms the known fact that, in Kalmesh et al. (2012) reported the direct selection for grain yield
236 Transgressive Segregation for Yield Using O. meridionalis

Table 3. Trait correlation for yield and related traits in BC2F3 population of IR58025B/ O. meridionalis under irrigated (I) and aerobic (A) conditions
DFF TN PN PL GN SN SF TW BM GY
PH (I) 0.02 - 0.24** - 0.22** 0.50** 0.25** 0.20** 0.11 0.22** 0.41** 0.22**
PH (A) - 0.02 0.21** 0.70** 0.23** 0.25** 0.05 0.30** 0.30** 0.31**
DF (I) 0.13 0.15* - 0.04 - 0.11 0.01 - 0.17 - 0.28** 0.28** - 0.15*
DF (A) - - - - - - - - -
TN (I) 0.94** - 0.16* - 0.19 - 0.19* - 0.07 - 0.11 0.13* 0.13**
TN (A) 0.51** 0.12* 0.03 0.03 - 0.01 0.10 0.33** 0.25**
PN (I) - 0.16* - 0.18 - 0.19* - 0.06 - 0.09 0.15* 0.15*
PN (A) 0.17* 0.11 - 0.02 0.13 0.31** 0.36** 0.50**
PL (I) 0.278** 0.27** 0.08 0.1 0.25** 0.25**
PL (A) 0.32** 0.28** 0.12 0.33** 0.24** 0.37**
GN (I) 0.75** 0.55** 0.12 0.17** 0.46**
GN (A) 0.61** 0.63** 0.12* 0.08 0.47**
SN (I) - 0.11 - 0.01 0.24** 0.36**
SN (A) - 0.21** 0.05 0.06 0.20**
SF (I) 0.17** - 0.04 0.22**
SF (A) 0.24** 0.05 0.37**
TW (I) - 0.15* 0.32**
TW (A) 0.01 0.21**
BM (I) 0.24**
BM (A) 0.35**
Significant at p = 0.05* and p = 0.01**.
PH: plant height (cm), DF: days to 50% flowering, TN: tiller number, PN: panicle number, PL: panicle length (cm), GN: grain number, SN: spikelet number, SF: spikelet fertility
(%), TW: 1000 seed weight (g), BM: biomass (g), GY: grain yield (g), PY: plot yield (t ha-1)

under reproductive stage moisture stress using O. meridionalis and
it has not been evaluated under aerobic conditions by exploiting O.
meridionalis for increasing yield in rice. The high-yielding trans-
gressive segregants reported in this study are similar to earlier
reports of obtaining positive transgressive segregants derived from
inter-specific crosses with different wild species of Oryza (O. rufi-
pogon, Marri et al. 2005; Moncada et al. 2001; Septiningsih et al.
2003; Thomson et al. 2003; Xiao et al. 1998; O. glaberrima, Aluko
et al. 2004; O. glumaepatula, Brondani et al. 2002; O. nivara,
Kaladhar et al. 2008; Swamy et al.2011).
Mean values of the population for all the yield component
traits under irrigated and aerobic conditions are significantly high-
er than the recurrent parent which suggests the influence of O.
meridionalis alleles in increasing the yield under both conditions.
GY of RP was reduced (74%) drastically under aerobic compared
to irrigated conditions. This shows the amount of moisture stress
imposed under aerobic conditions and also sensitivity of RP to
moisture stress, whereas in the population the reduction of GY
value is lower (69%) when compared to the reduction of GY of
RP from irrigated to aerobic conditions. This gives evidence for
the presence of positive alleles in O. meridionalis that help in
withstanding moisture stress. O. meridionalis is found growing in
periphery of water bodies and seeds survive the dry season
(Waters et al. 2012). Transgressive families whose performance
exceeded either of the parents in a positive direction were
observed for yield and related traits under both conditions. These
progenies are potentially useful breeding material for further stud-
ies as they serve as diverse germplasm for improving the yield of
hybrids under irrigated and aerobic conditions.
The results of ANOVA, estimates of GCV and PCV revealed
the presence of substantial genetic variation in the population for
yield and related traits under both conditions. Traits such as GY,
GN, and BM recorded high GCV and PCV coupled with high to
moderate h2 and GA values under both the situations indicate
additive genetic control of these traits. The h2 of GY under irrigat-
ed and aerobic conditions was high (61.9%) and moderate
(49.6%), respectively. This is in agreement with the recent reports
on comparable h2 values for grain yield under stress and non-
stress conditions (Lanceras et al. 2004, Venuprasad et al. 2007).
This indicates that direct selection for grain yield under both irri-
gated and aerobic conditions is likely to be effective. In trait cor-
relations, TN (0.13 and 0.25), PN (0.15 and 0.50), PL (0.25 and
0.37), GN (0.46 and 0.47), SN (0.36 and 0.20), SF (0.22 and
0.37), TW (0.32 and 0.21), and BM (0.24 and 0.35) followed a
significant positive relationship with yield under irrigated and aer-
obic conditions (Table 3). Hence, indirect selection of progenies
based on these traits will also result in deriving lines with higher
yields in advanced generations (Yuan et al. 2011) under both con-
ditions. Trait correlations are in agreement with earlier reports
involving O. rufipogon, O. glumaepatula, and O. grandiglumis.
Grain number is highly positively correlated with GY as reported
earlier in O. rufipogon by Moncada et al. (2001), in O. glumaepat-
ula by Brondani et al. (2002), and in O. grandiglumis by Yoon et
al. (2006). Single plant yield showed a positive correlation with
TW, PL, BM, and PH as reported in previous studies (TW,
Moncada et al. 2001; PL, Yoon et al. 2006; BM, Peng et al. 2004;
PH, Moncada et al. 2001). PL had a positive correlation with plant
height as reported by Thomson et al. (2003) and Yoon et al.
(2006) using O. rufipogon and O. grandiglumis, respectively.
Correlation patterns of PH and PN, TN and PL, and PN and PL
differed significantly from irrigated to aerobic conditions, indicat-
ing the role of G X E interaction on these traits. Histograms
showed (Fig. 3) near normal distribution for all traits under both
conditions indicating their quantitative nature.
The results from this investigation indicate that O. meridionalis
has not only yield-enhancing alleles under normal, well-watered
conditions, but also alleles which can help withstand water stress

under aerobic conditions. Hence, this species can be a novel
source of natural genetic variation for the improvement of rice
under irrigated as well as under aerobic conditions and simultane-
ously help in expansion of the cultivated gene pool of O. sativa.
We are generating genotyping data to map yield-related QTLs
from O. meridionalis and it will be interesting to see whether the
location of QTLs contributed by this Australian wild species is
common with those reported for other wild species.
Acknowledgements
We gratefully acknowledge Mr. Dinesh C. Joshi, Executive
Director and the Management of Barwale Foundation, for all the
encouragement and support provided during the project. We thank
Dr. J.R. Diwan for maintaining and providing of O. meridionalis
at the initial stage of this research and Dr. M.S. Ramesha for criti-
cal reviews of this manuscript. We thank Project Director, DRR
Dr. B.C Viraktamath for extending his support.
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