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Research Article

QTL Mapping of Rice Root Traits at Different NH4+ Levels in Hydroponic Condition

Pyeong-Sook Kim1, Dong-Min Kim2, Ju-Won Kang1, Hyun-Sook Lee1, Sang-Nag Ahn1,*
Plant Breeding and Biotechnology 2015;3(3):244-252.
Published online: September 30, 2015

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*Corresponding author: Sang-Nag Ahn, ahnsn@cnu.ac.kr, Tel: +82-42-821-5728, Fax: +82-42-822-2631
• Received: September 10, 2015   • Revised: September 15, 2015   • Accepted: September 22, 2015

Copyright © 2015 The Korean Society of Breeding Science

This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/3.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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  • This study was initiated to identify quantitative trait loci (QTL) involved in root system development by measuring root length and weight of rice seedlings grown in hydroponic culture condition. Growth conditions for estimating the root elongation were set up to supply NH4+ as a single nitrogen source. Four rice accessions and 40 introgression lines (ILs) derived from a cross between ‘Ilpumbyeo’, a temperate japonica variety, and ‘Moroberekan’, a tropical japonica accession, were used to detect QTL for two root traits of seedlings grown in 0, 250 and 500 μM NH4+. Seven chromosomal regions were found to be involved in root elongation. The qRW6 QTL detected on the long-arm of chromosome 6 promoted root weight and the ‘Moroberekan’ allele at this QTL was associated with enhanced root weight at 250 μM NH4+ concentration. The Moroberekan alleles were beneficial at all loci except for qRW4 at which the Ilpumbyeo allele increased root weight under 0, 250 and 500 μM NH4+. A field trial was conducted for selected four ILs to know whether the introgressed Moroberekan segments harboring the beneficial QTL were associated with effects on other agronomic traits. No significant difference was detected between four ILs and Ilpumbyeo for days to heading and culm length. Two lines showed significant difference from Ilpumbyeo in spikelets per panicle and 1,000 grain weight, and the difference might be attributed to the Moroberekan segment on chromosome 6. Our QTL results indicate that the Moroberekan introgression on chromosome 6 flanked by RM162-RM3430 has the potential to enhance root traits and yield potential in the temperate japonica background.
  • Key words: Rice, Introgression lines, NH4+ response, Root elongation, QTL
Rice is one of the most important crops for almost 50% of the world’s population. Dramatic increase in the production and use of fertilizer N in rice-growing countries has occurred during the past decades (Craswell and Vlek 1983). The usual cultivation of rice includes major changes in the N supply form as the seedling plants are initially raised in a nursery bed and subsequently transferred to the flooded paddy culture system (Patil et al. 2013). Even though N fertilizer increases plant yield, plants consume much less than half of the fertilizers applied (Frink et al. 1999; Socolow 1999) while a majority of N fertilizers were lost to the atmosphere or leached into groundwater, lakes and rivers, which causes severe adverse effects to the environment.
Nitrogen (N) status strongly affects the photosynthetic rate in crop. Rice evolved with greater capacity to assimilate ammonium ion (NH4+) compared with other plant species (Yamaya and Oaks 2004). Studies have shown NH4+ can improve the capacity to tolerate water stress in rice in comparison with NO3− (Guo et al. 2007). However, NH4+ frequently reaches levels in soils that affect plant growth negatively. These negative effects manifest in stunted root growth, yield depression, and chlorosis of leaves (Britto and Kronzucker 2002; Balkos et al. 2010). High NH4+ mainly inhibited the development of total root length and root area while the formation of lateral roots was not affected (Zhao et al. 2014). Nitrogen deficiency delayed both vegetative and reproductive phenological development, slightly reduced leaf emergence rate, and strongly diminished leaf expands. Therefore, it is important to find ways to enhance nitrogen use efficiency (NUE) in rice.
Root morphological and physiological traits are controlled by QTL. Root system architecture (RSA) is generally defined by length, weight, number, thickness and density of primary, lateral and adventitious roots (Obara et al. 2010) but RSA study is difficult to handle in soil condition. To overcome these limitations, many of the root studies grow rice seedlings in hydroponic system using diverse mapping populations (Obara et al. 2010; Ogawa et al. 2014). The results suggest that QTLs for root length uncovered under hydroponics may be predictive of enhanced grain yield under field conditions (Obara et al. 2011). This approach has been widely used to detect QTLs associated with improved root systems in both stressed and non-stressed rice fields (Price et al. 1997; Shimizu et al. 2004). However, QTL studies for root traits using mapping populations from crosses between japonica accessions are limited.
In this study, we established growth condition to reliably evaluate rice seedling traits grown in hydroponic condition to identify chromosomal regions associated with root traits under different NH4+ levels using introgression lines (ILs) derived from a cross between two japonica rice accessions ‘Ilpumbyeo’ and ‘Moroberekan’.
Plant materials
Four rice accessions ‘Ilpumbyeo’, ‘Moroberekan’, ‘Hwaseongbyeo’ and ‘O. rufipogon’ (IRGC 105491) were used for parental screening. In the previous research, 117 BC3F5 lines were developed from a cross between Ilpumbyeo and Moroberekan (Kim et al. 2009). From the 117 BC3F5, 40 ILs (introgression lines) covering the whole Moroberekan chromosome in the genetic background of Ilpumbyeo were selected and used in this study (Fig. 1, Kim et al. 2009).
A field trial was conducted to compare selected four ILs with Ilpumbyeo for agronomic traits. Plots for all genotypes were laid out in a completely randomized block design with two replications, one row per line, 25 plants per each row in 2013. Four phenotypic traits, days to heading (DTH), culm length (CL), 1,000 grain weight in brown rice (GW) and spikelets per panicle (SPP) were measured. The middle 3 plants from each line were chosen for the evaluation and average number of the measurements was used for the phenotype of each line for the four traits: DTH, CN, GW, and SPP. Evaluation of the traits followed the method as described in Kim et al. (2014)
Seeds sterilization
All seeds were pre-selected with NaCl solution (d= 1.113) and processed by three-step of sterilization (Obara et al. 2010). First, the seeds were soaked for 10 min in distilled water at 60°C and then washed for 20 min with running tap water. Second, the seeds were soaked for 30 sec in a 70% (v/v) ethanol solution and they were washed for 2 min with distilled water. Finally, the seeds were soaked for 20 min in a 1% (v/v) sodium hypochlorite solution and then washed four times with distilled water.
Hydroponic condition and trait evaluation
Seeds were kept for germination at 30°C, in dark conditions for 2 days in an incubator. After the tip of the plumule had barely emerged, 50 seeds were placed between moist paper towels and then gently rolled. Rolled papers were then transferred to plastic boxes (456 × 296 × 112 mm). Rolled seedlings were grown in a chamber with a 14-hr light at 30°C/10h dark at 28°C, 70% relative humidity. 7 days after imbibition, uniform seedlings established in moist paper towels were selected and moved in a water culture tube with a sponge. The seedlings were grown in 0, 250, and 500 μM NH4Cl dissolved in 10L half strength of Yoshida solution (Yoshida et al. 1976). The seedlings were grown in a chamber with a 14-h light at 28°C/10-h dark at 25°C, 70% relative humidity. The experiments were conducted in a completely randomized design with two replications for each treatment. When NH4Cl was applied to rice plants as a single source of nitrogen, the pH of the nutrient solutions shifted to the acidic region, around pH 3.0–4.0. To maintain pH, the nutrient solution was replaced every three days to maintain pH 5.5~5.6. 10 days after being treated. Seedling root lengths were measured with a ruler, and root fresh weight (RW) was measured with a scale.
Data analysis
The differences among mean values for seedling root traits (4 accessions) were statistically analyzed by one-way ANOVA (Tukey’s test with Minitab 6.0) (Ryan et al. 1992). Previously, 134 SSR markers were mapped for the 117 BC3F5 lines and these data were used for detecting QTLs in 40 ILs (Kim et al. 2009). Single point analysis (SPA) was performed to determine the effect of each marker on each trait using Q-gene program (Nelson 1997). In SPA, QTL was declared if the phenotype was associated with a marker locus at P<0.005 or with two adjacent marker loci at P<0.01.
Variation of root traits in response to NH4+ supply among four rice accessions
Root length and weight of ‘Ilpumbyeo’, ‘Hwaseongbyeo’, ‘Moroberekan’ and ‘O. rufipogon’ plants grown in hydroponic culture with three NH4+ concentrations was measured as an indicator of genetic variation (Table 1). Four rice accessions were grown for 11 days. Moroberekan and O. rufipogon showed significantly higher values in root length and weight than Ilpumbyeo and Hwaseongbyeo (P<0.05) at all NH4+ levels. At 250 μM Hwaseong showed the shorter roots than Ilpumbyeo. Moroberekan and O. rufipogon were not significantly different in root length and weight at all NH4+ levels. Ilpumbyeo displayed better growth for roor length and weight at 250 μM NH4+ than the other two NH4+ levels. Hwaseongbyeo performed poor showing shortest roots of 14.3 cm at 250 μM NH4+ whereas no difference was observed for root weight between 250 and 500 μM levels.
Frequency distribution of 40 CSSLs and the parents
Seedlings of 40 ILs and their parental lines were grown in hydroponic culture conditions, supplemented with 0, 250, and 500 μM NH4+. Frequency distributions of phenotypes of RL and RW of the 40 ILs are shown in Fig. 2. RL value showed bimodal distributions at three NH4+ levels and ranged from 14.0 to 23.2 cm at 250 μM NH4+. RW also showed bimodal distributions. Transgressive lines with phenotypic values larger than Moroberekan were observed for RL and RW.
The root length of Ilpumbyeo and Moroberekan were 15.5 ± 1.8 and 20.4 ± 2.7 cm under the 0 μM NH4+ level, 18.8 ± 2.1 and 19.9 ± 2.8 cm under the 250 μM NH4+ level and 15.7 ± 2.0 and 19.5 ± 2.4 cm under the 500 μM NH4+ level, indicating that genetic variation of root length in response to exogenous NH4+ concentration exists between the parental lines in hydroponic condition.
The root weight of Ilpumbyeo and Moroberekan were 0.05 ± 0.02 and 0.09 ± 0.03 g under the 0 μM NH4+ level, 0.06 ± 0.02 and 0.10 ± 0.03 g under the 250 μM NH4+ level and 0.05 ± 0.01 and 0.10 ± 0.03 g under the 500 μM NH4+ level, indicating genetic variation of root weight.
QTL analysis
Seven QTLs distributed on six chromosomes (1, 2, 4, 6, 8, and 10) were detected in this study (Table 2, and Fig. 3). One QTL, qRW4 was detected under 0, 250 and 500 NH4+ levels among seven QTLs. The other QTLs include three QTLs (qRW2, qRW8, and qRL10)under 0 μM NH4+, one QTL (qRW6)under 250 μM NH4+ level, 2 QTLs (qRL1 and qRW1) detected under 500 μM NH4+ level for RL and RW.
Two QTL of root length (RL) were detected on chromosomes 1 and 10. One QTL, qRL1 was detected at the marker interval of RM259-RM140 under 500 μM NH4+ level, and the other QTL, qRL10 was detected near the marker interval of RM474-RM216 under 0 μM NH4+ level. The phenotypic variance and LOD peak of two QTL were 31.4% and 3.3 and 17.7% and 1.7, respectively. Moroberekan alleles increased root length at two loci.
Five QTL of root weight (RW) were detected on chromosomes 1, 2, 4, 6 and 8. qRW1 on chromosome 1 was detected near the marker RM302-RM265 under 500 μM NH4+ level. qRW2 on chromosome 2 was detected near the marker interval RM263-RM5404 under the 0 μM NH4+ level. The QTL on chromosome 4, qRW4 was detected at the marker interval of RM7410-RM3333 at three NH4+ levels and at this locus the Ilpumbyeo allele increased root weight. The qRW4 QTL explained 26.6–27.7% of the phenotype variance for root weight (data not shown). qRW6 on chromosome 6 was detected near the marker RM7269-RM3430 under 250 μM NH4+ level. qRW8 on chromosome 8 was detected under the 0 μM NH4+ level.
Agronomic traits of selected ILs
To determine whether the introgressed segments harboring the beneficial QTL were associated with effects on other yield component and grain traits, a field trial was conducted using seeds from selected four ILs, and the parent, Ilpumbyeo (Table 3). These four lines performed better than Ilpumbyeo in root weight and length in varying NH4+ levels and were phenotypically similar to Ilpumbyeo. No significant difference was detected between four ILs and Ilpumbyeo for days to heading and culm length. Two lines, IL14 and IL24 showed significant difference from Ilpumbyeo for spikelets per panicle and 1,000 grain weight, and the difference might be attributed to the Moroberekan segment on chromosome 6 (Kim et al. 2014).
Nitrogen (N) is essential for plant growth and development (Su et al. 2005). Even though N fertilizer increases plant yield, nitrogen use efficiency of plants is generally low (Frink et al. 1999). Rice plants mainly depend on NH4+ that is the major form of nitrogen in paddy fields (Yamaya et al. 2004). Excessive NH4+ inhibits the growth of most crop species (Roosta et al. 2008). Several studies have been conducted to identify QTL for root traits associated with different NH4+ concentrations using various mapping populations (Obara et al. 2010; Obara et al. 2011; Chun et al. 2014; Ogawa et al. 2014). However, report on root traits using ILs derived from a cross between japonica accessions is lacking.
We evaluated four rice accessions under different NH4+, and Moroberekan and O. rufipogon consistently displayed enhanced RL and RW compared with the other two temperate japonica genotypes, Ilpumbyeo and Hwaseongbyeo. Two temperate japonica accessions at 250 μM experienced the maximum growth compared to 0 and 500 μM NH4+ levels. Moroberekan and O. rufipogon were not significantly different for two root traits in various NH4+ concentrations. Therefore, Moroberekan and O. rufipogon were classified as NH4+ less-sensitive. Ogawa et al. (2013) also reported that O. rufipogon constitutively elongated seminal roots under 5, 50, and 500 μM NH4+ concentrations being not NH4+ sensitive. These results also confirm the previous results that Moroberekan and O. rufipogon possess beneficial alleles for root architectural traits in breeding programs to enhance O. sativa. Moroberekan has been previously used as a donor of many useful agronomical traits, such as resistance to drought or blast disease (Lilley et al. 1996; Ju et al. 2004; Kang et al. 2011). O. rufipogon has also been utilized as a source of biotic and abiotic stress tolerance (Brar and Khush 1997) and as a source of yield-enhancing alleles for O. sativa (Imai et al. 2013). Because O. rufipogon can be easily crossed with O. sativa, this progenitor species represents a valuable source of useful traits for rice improvement, including root architectural traits.
To identify and map QTL associated with root traits, we evaluated 40 ILs from a cross between Ilpumbyeo and Moroberekan and detected a total of 7 QTLs. When the chromosomal locations of the QTL detected in this study were compared with those of the previously detected QTL (www.gramene.org), several overlapping loci were observed. qRW1 near the marker RM302-RM265 on chromosome 1 was colocated with a QTL for seminal root elongation by Obara et al. (2010 and 2011), which was detected in rice seedlings grown in hydrophonic condition using two different mapping populations derived from Koshihikari × Kasalath and Taichung 65 × O. glaberrima. qRW1 was also colocalized with a major QTL for seminal root elongation under 500 μM NH4+ using a CSSL population from a cross between ‘Curinga’ and O. rufipogon (Ogawa et al. 2014) and a QTL for shoot dry weight, and plant dry weight (Wei et al. 2012). In our study, a qRW1 introgression from Moroberekan in the Ilpumbyeo genetic background was associated with higher root weight at 500 μM NH4+ level. The qRW4 QTL on chromosome 4 shared the similar chromosomal regions related to root elongation (Obara et al. 2010) and to nitrogen-use and nitrogen-deficiency tolerance (Wei et al. 2012). qRW6 detected in 250 μM NH4+ level in our study shared the similar chromosomal region with a QTL, qRL6.1 for seminal root elongation in a wide range of NH4+ concentrations (Obara et al. 2010). qRL6.1 was narrowed down to a 336 kb-long region in the genome sequence and the ‘Kasalath’ allele on qRL6.1 was associated with enhanced root elongation in the ‘Nipponbare’ genetic background. This region also harbors a QTL for maximum root length of seedlings grown in soil culture and a QTL for maximum root length of plants grown in low nitrogen conditions (MacMillan et al. 2006). The QTL, qRW8 shared the similar location with a QTL for nitrogen response (Cho et al. 2007; Obara et al. 2010; Zhao et al. 2014). qRL10 on chromosome 10 under 0 μM NH4+ level detected in RM474-RM216 was colocated with QTL for relative biomass and nitrogen response (Lian et al. 2005; Wei et al. 2012). The QTL, qRW8 shared the similar location with a QTL for nitrogen response (Cho et al. 2007; Obara et al. 2010; Zhao et al. 2014).
However, it is very difficult to conclude whether these QTL are allelic or not because these QTL were mapped by QTL analysis, not by fine-mapping as was conducted for qRL6.1 by Obara et al. (2010). Additional fine-mapping will be required to conclude an allelic relationship among these QTL loci. It is noteworthy that the Moroberekan introgression near RM3430-RM20580 harbors a QTL cluster associated with grain weight and spikelets per panicle in the Ilpumbyeo genetic background (Kim et al. 2014). Our QTL results indicate that the Moroberekan introgression on chromosome 6 flanked by RM162-RM3430 has the potential to enhance root traits and yield potential in the temperate japonica background. All the Moroberekan alleles were associated with an increase in root length and weight except for qRW4. Although Moroberekan had higher root weight than Ilpumbyeo, the Ilpumbyeo allele at qRW4 was favorable in terms of root weight. This result is similar with the study of Obara et al. (2010) in which a QTL for root length was detected on chromosome 4 using CSSLs and the Kasalath allele at this QTL locus inhibited root elongation. Frequency distribution indicated the existence of several lines showing transgressive segregation for RL and RW at 3 NH4+ levels and this might be explained by this QTL due to a combination of favorable alleles from both parents. Further analyses for validating effect of qRW4 and qRW6 on root growth are underway.
In conclusion, this work is only a starting point for characterizing the genetic basis of rice growth in different NH4+ levels. A few lines with beneficial QTLs for root length and weight from Moroberekan in this study would be useful in developing lines with root system development in the breeding program.
This work was supported by the Cooperative Research Program for Agriculture Science and Technology Development (Project No. PJ906910 to S.N.A), Rural Development Administration, Republic of Korea.
Fig. 1
Graphical genotypes of 40 introgression lines used in this study. Black boxes indicat Moroberekan introgressions in the Ilpumbyeo background. Markers are shown on each chromosome.
pbb-3-244f1.jpg
Fig. 2
Distribution of root length and weight in 40 ILs with the parental lines grown in hydroponic condition under 0, 250, and 500 μM NH4+.
pbb-3-244f2.jpg
Fig. 3
Chromosomal regions affecting root length and weight of seedlings grown in hydroponic culture. Open, slashed and closed columns on the left side of the chromosomes 1, 2, 4, 6, 8 and 10 indicate the most likely chromosomal regions of the QTL for root length and weight in 0, 250 and 500 μM NH4+, respectively.
pbb-3-244f3.jpg
Table 1
Comparison of two root traits among 4 accessions grown in hydroponic conditions at three different NH4+ levels.
Table 1
Traitz) NH4+ (μM) Ilpumbyeo Moroberekan Hwaseongbyeo O. rufipogon
0 15.5 ± 1.8 by),bx) 20.4 ± 2.7 a,a 15.9 ± 2.6 a,b 21.2 ± 2.2 a,a
RL 250 18.8 ± 2.1 a,a 19.9 ± 2.8 a,a 17.1 ± 2.9 a,b 20.3 ± 2.8 a,a
500 15.7 ± 2.0 b,b 19.5 ± 2.4 a,a 14.3 ± 2.4 b,b 20.7 ± 1.4 a,a
0 0.05 ± 0.01 b,b 0.09 ± 0.01 a,a 0.05 ± 0.01 b,b 0.11 ± 0.01 a,a
RW 250 0.06 ± 0.01 a,b 0.10 ± 0.01a,a 0.06 ± 0.02 a,b 0.11 ± 0.01 a,a
500 0.05 ± 0.01 b,b 0.10 ± 0.00 a,a 0.07 ± 0.02 a,b 0.11 ± 0.01 a,a

z)RL: root length (cm), RW: root weight (g)

y)Numbers followed by different letters are significantly different among three NH4+ levels in a row for each trait (Tukey’s test, P<0.05).

x)Numbers followed by different letters are significantly different among four rice accessions in a column for each trait (Tukey’s test, P<0.05).

Table 2
Characteristics of the QTLs detected in the 40 ILs.
Table 2
Chr. Level QTL Marker Trait P Trait meansz)
II MM
1 500 qRL1 RM259 - RM140 RL 0.01 17.6(33)y) 18.8(4)
1 500 qRW1 RM302 - RM265 RW 0.01 0.10(38) 0.13(2)
2 0 qRW2 RM263 - RM5404 RW 0.01 0.11(35) 0.14(5)
4 0 qRW4 RM7410 - RM3333 RW 0.01 0.12(30) 0.08(7)
4 250 qRW4 RM7410 - RM3333 RW 0.01 0.11(30) 0.09(7)
4 500 qRW4 RM7410 - RM3333 RW 0.01 0.11(30) 0.09(7)
6 250 qRW6 RM7269 - RM3430 RW 0.005 0.09(37) 0.14(3)
8 0 qRW8 RM339 - RM42 RW 0.01 0.10(35) 0.13(5)
10 0 qRL10 RM474 - RM216 RL 0.01 18.8(36) 20.8(4)

z)II, MM: Phenotypic means of the Ilpumbyeo and Moroberekan homozygotes, respectively.

y)No. of lines in each genotype class.

Table 3
Comparison of agronomic traits of four lines with Ilpum.
Table 3
Line DTH (days) CL (cm) SPP (no.) GW (g) RL (cm) RW (g) Moroberekan introgression segments
No. Chr.
IL7 115az) 84a 122b 21.2b 18.2a 0.087b 3 2, 10 (qRW10)
IL12 116a 81a 145a 23.4a 15.4b 0.142a 4 4, 6, 10 (qRW10), 11
IL14 117a 85a 125b 22.2b 16.7a 0.110a 5 1(qRL1), 2, 6(qRW6), 7, 8(qRW8)
IL24 114a 85a 142a 23.1a 16.6a 0.107a 2 4, 6 (qRW6)
Ilpum 116a 84a 128b 21.8b 14.6b 0.090b

z)Numbers followed by different letters are significantly different among lines in a row for each trait (Tukey’s test, P<0.05).

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QTL Mapping of Rice Root Traits at Different NH4+ Levels in Hydroponic Condition
Plant Breed. Biotech.. 2015;3(3):244-252.   Published online September 30, 2015
DOI: https://doi.org/10.9787/PBB.2015.3.3.244
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QTL Mapping of Rice Root Traits at Different NH4+ Levels in Hydroponic Condition
Plant Breed. Biotech.. 2015;3(3):244-252.   Published online September 30, 2015
DOI: https://doi.org/10.9787/PBB.2015.3.3.244
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QTL Mapping of Rice Root Traits at Different NH4+ Levels in Hydroponic Condition
Image Image Image
Fig. 1 Graphical genotypes of 40 introgression lines used in this study. Black boxes indicat Moroberekan introgressions in the Ilpumbyeo background. Markers are shown on each chromosome.
Fig. 2 Distribution of root length and weight in 40 ILs with the parental lines grown in hydroponic condition under 0, 250, and 500 μM NH4+.
Fig. 3 Chromosomal regions affecting root length and weight of seedlings grown in hydroponic culture. Open, slashed and closed columns on the left side of the chromosomes 1, 2, 4, 6, 8 and 10 indicate the most likely chromosomal regions of the QTL for root length and weight in 0, 250 and 500 μM NH4+, respectively.

Fig. 1

Fig. 2

Fig. 3

QTL Mapping of Rice Root Traits at Different NH4+ Levels in Hydroponic Condition

Comparison of two root traits among 4 accessions grown in hydroponic conditions at three different NH4+ levels.

Traitz) NH4+ (μM) Ilpumbyeo Moroberekan Hwaseongbyeo O. rufipogon
0 15.5 ± 1.8 by),bx) 20.4 ± 2.7 a,a 15.9 ± 2.6 a,b 21.2 ± 2.2 a,a
RL 250 18.8 ± 2.1 a,a 19.9 ± 2.8 a,a 17.1 ± 2.9 a,b 20.3 ± 2.8 a,a
500 15.7 ± 2.0 b,b 19.5 ± 2.4 a,a 14.3 ± 2.4 b,b 20.7 ± 1.4 a,a
0 0.05 ± 0.01 b,b 0.09 ± 0.01 a,a 0.05 ± 0.01 b,b 0.11 ± 0.01 a,a
RW 250 0.06 ± 0.01 a,b 0.10 ± 0.01a,a 0.06 ± 0.02 a,b 0.11 ± 0.01 a,a
500 0.05 ± 0.01 b,b 0.10 ± 0.00 a,a 0.07 ± 0.02 a,b 0.11 ± 0.01 a,a

z)RL: root length (cm), RW: root weight (g)

y)Numbers followed by different letters are significantly different among three NH4+ levels in a row for each trait (Tukey’s test, P<0.05).

x)Numbers followed by different letters are significantly different among four rice accessions in a column for each trait (Tukey’s test, P<0.05).

Characteristics of the QTLs detected in the 40 ILs.

Chr. Level QTL Marker Trait P Trait meansz)
II MM
1 500 qRL1 RM259 - RM140 RL 0.01 17.6(33)y) 18.8(4)
1 500 qRW1 RM302 - RM265 RW 0.01 0.10(38) 0.13(2)
2 0 qRW2 RM263 - RM5404 RW 0.01 0.11(35) 0.14(5)
4 0 qRW4 RM7410 - RM3333 RW 0.01 0.12(30) 0.08(7)
4 250 qRW4 RM7410 - RM3333 RW 0.01 0.11(30) 0.09(7)
4 500 qRW4 RM7410 - RM3333 RW 0.01 0.11(30) 0.09(7)
6 250 qRW6 RM7269 - RM3430 RW 0.005 0.09(37) 0.14(3)
8 0 qRW8 RM339 - RM42 RW 0.01 0.10(35) 0.13(5)
10 0 qRL10 RM474 - RM216 RL 0.01 18.8(36) 20.8(4)

z)II, MM: Phenotypic means of the Ilpumbyeo and Moroberekan homozygotes, respectively.

y)No. of lines in each genotype class.

Comparison of agronomic traits of four lines with Ilpum.

Line DTH (days) CL (cm) SPP (no.) GW (g) RL (cm) RW (g) Moroberekan introgression segments
No. Chr.
IL7 115az) 84a 122b 21.2b 18.2a 0.087b 3 2, 10 (qRW10)
IL12 116a 81a 145a 23.4a 15.4b 0.142a 4 4, 6, 10 (qRW10), 11
IL14 117a 85a 125b 22.2b 16.7a 0.110a 5 1(qRL1), 2, 6(qRW6), 7, 8(qRW8)
IL24 114a 85a 142a 23.1a 16.6a 0.107a 2 4, 6 (qRW6)
Ilpum 116a 84a 128b 21.8b 14.6b 0.090b

z)Numbers followed by different letters are significantly different among lines in a row for each trait (Tukey’s test, P<0.05).

Table 1 Comparison of two root traits among 4 accessions grown in hydroponic conditions at three different NH4+ levels.

RL: root length (cm), RW: root weight (g)

Numbers followed by different letters are significantly different among three NH4+ levels in a row for each trait (Tukey’s test, P<0.05).

Numbers followed by different letters are significantly different among four rice accessions in a column for each trait (Tukey’s test, P<0.05).

Table 2 Characteristics of the QTLs detected in the 40 ILs.

II, MM: Phenotypic means of the Ilpumbyeo and Moroberekan homozygotes, respectively.

No. of lines in each genotype class.

Table 3 Comparison of agronomic traits of four lines with Ilpum.

Numbers followed by different letters are significantly different among lines in a row for each trait (Tukey’s test, P<0.05).

Table 1

Table 2

Table 3

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