Identification of loci for cold tolerance at reproductive stage and seed shattering in Ethiopian rice cultivar ‘X-Jigna’

Identification of loci for cold tolerance at reproductive stage and seed shattering in Ethiopian rice cultivar ‘X-Jigna’ | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Identification of loci for cold tolerance at reproductive stage and seed shattering in Ethiopian rice cultivar ‘X-Jigna’ Zelalem Zewdu, Mulugeta Atnaf, Assaye Berie, Honoka Takasago, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8244416/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract Genetic improvement is essential for improved rice production in the high-elevation, cool-climate regions of Ethiopia. We conducted QTL-seq analysis using 846 RIL F 5 populations of ‘X-Jigna’, a dominant rice cultivar in Ethiopia, × ‘Hitomebore’, a cold-tolerant elite Japanese cultivar, in field trials in Japan. We detected three QTLs for cold tolerance for inducing spikelet sterility under cold at reproductive stage ( qSFC5 , qSFC8 , and qSFC10 ) and one for seed shattering habit ( qSHT1 ). In haplotype analysis using indel markers in an F 6 population in Japan and Ethiopia, qSFC5 and qSFC8 were confirmed in Japan and qSHT1 was confirmed in both Japan and Ethiopia. Markers for these will be useful to speed up breeding in Ethiopia. Cold tolerance Ethiopia QTL-seq Rice Shattering Figures Figure 1 Figure 2 1. Introduction Rice consumption in Ethiopia has increased 10× since 2010, to > 2.5 Mt annually in 2021 (FAO, 2024 ). It has become a challenge to meet this demand solely through domestic production, which is < 10% of the self-sufficiency rate in spite of 30 million hectares of potential cropland. In Ethiopia, rice is grown mostly in high-elevation regions above 1800 m in altitude, notably on the Fogera Plain, which accounts for 71% of the total rice production in the country (Ministry of Agriculture and Rural Development, 2010 ). Owing to the high elevation, low temperatures place a major constraint on rice production (Dessie, 2020 ; Zewdu et al., 2025 ): the minimum temperature drops to 13°C during the reproductive stage, which is the most sensitive period in terms of induction of sterility and severe yield losses (Satake 1976 ; Shimono et al., 2002 ). The rice cultivar ‘X-Jinga’ is the Ethiopian farmers’ favorite cultivar, and was grown on > 80% of the Fogera area in 2021 (Beyene et al., 2022 ). Introduced into Ethiopia in the 1980s from North Korea, it is favored for its high yield, its ability to make good-quality injera , a staple food traditionally made from teff ( Eragrostis tef ) flour, and its high biomass and long culm for forage. However, ‘X-Jigna’ is highly susceptible to cold stress during the reproductive stage, which induces spikelet sterility, unlike in the Japanese cold-tolerant elite cultivar ‘Hitomebore’ (Alemayehu et al. 2021 ; Zewdu et al., 2025 ). Identification of DNA markers may facilitate the development of suitable rice genotypes through marker-assisted selection. The processes involved in cold damage during fertilization are complex. The molecular and physiological aspects of damage to male gametes are controlled by many parallel and interacting processes (Shimono, 2018 ), including metabolic processes related to carbohydrates (Oliver et al., 2005 ), amino acids (Ito et al., 1970 ), reactive oxygen species (Suzuki et al., 2015 ), heat-shock proteins (Sato et al., 2011 ; Suzuki et al., 2015 ), and plant hormones such as abscisic acid (Oliver et al., 2007 ) and gibberellins (Sakata et al., 2014 ). It is important to identify the genomic regions causing genotypic differences in the weak cold tolerance of ‘X-Jigna’ for future breeding. The cold-tolerant ‘Hitomebore’ may hold novel quantitative trait loci (QTLs). While a number of QTLs for cold tolerance have been identified in various cultivars (Andaya and Mackill, 2003 ; Kuroki et al., 2007 ; Xu et al., 2008 ; Shimono et al., 2016 ), no study has tested Ethiopian cultivars. Another shortcoming of ‘X-Jigna’ is its easy seed shattering habit (Zewdu et al., 2025 ), which decreases the yield before harvest (Wang et al., 2024 ). The habit is controlled by the formation of an abscission layer in the joint between the lemma and pedicel (Wu et al., 2023 ). Many QTLs related to seed shattering have been reported and identified through the use of various genetic populations and linkage maps (Wu et al., 2023 ). qSH1 , on chromosome (chr.) 1 (Konishi et al., 2006 ), and sh4 , on chr. 4 (Li et al., 2006 ), are dominant QTLs that control formation of the abscission layer, as confirmed in many genetic populations (Wu et al., 2023 ). Introduction of the equivalent genomic region of ‘Hitomebore’ should reduce yield loss. However, the farmers’ postharvest management practices are closely linked to the shattering habit: the harvested plants are threshed by hand using sticks or with cattle (Tadesse et al., 2019 ). The farmers thus prefer the easy shattering habit, despite the significant yield losses. Balancing the reduction of yield loss with the efficient management of threshing would improve yields. Again, however, no QTL studies of seed shattering in Ethiopian cultivars have been conducted. The purpose of this study was to identify QTLs related to cold tolerance at the reproductive stage and to seed shattering in ‘X-Jigna’ and ‘Hitomebore’. We conducted QTL-seq analysis of bulk segregants (Takagi et al., 2013 ) using 846 RIL F 5 populations of ‘X-Jigna’ × ‘Hitomebore’ in Japan in 2023 (Experiment 1), and confirmed our results in both Japan and Ethiopia using F 6 populations in 2024 (Experiment 2). Compared with standard QTL analysis using genotyping data of all populations, QTL-seq analysis is rapid and requires less work for identifying QTLs, reducing the time and labor needed without genotyping individual lines, and has proven effective in identifying QTLs rapidly and cost-effectively in different crop species for various target traits (Takagi et al., 2013 ). 2. Materials and methods 2.1. QTL-seq analysis for detecting QTLs in 2023 2.1.1. Phenotyping in Japan Cold tolerance for inducing spikelet sterility by low temperature at reproductive stage was determined at a field-based cool-irrigation facility at the Iwate Prefecture Agricultural Research Center, Kitakami, Iwate, Japan (39°35′N, 141°11′E). The facility is commonly used for screening cold tolerance in breeding. The method maintains water depth at > 20 cm to cover the developing panicles uniformly so as to minimize errors caused by difference in rates of maturity, and controls the temperature of the developing panicles (Matsunaga, 2005 ; Shimono et al., 2005 ). Cool water sourced from aquifers or refrigeration units is combined with warmer irrigation water to achieve a constant 18–19°C, which is based on the cold tolerance of the population under test (Matsunaga, 2005 ). The method can control the temperature of the developing panicle uniformly under water before culm elongation especially including the most sensitive microspore and booting stage (Satake, 1976 ), although it can not control that of the developing panicle after it has emerged above the water (Shimono et al., 2005 ). The method has been used in Japan since 1929 (Matsunaga, 2005 ; Shimono et al., 2007 ) and is now widely used in Korea (Jeong et al., 2020 ), China (Li et al., 2021 ; Yang et al., 2023 ), and Australia (Farrell et al., 2006 ). From 846 F 5 RILs, we randomly selected 200 RILs with similar culm lengths and days to heading as ‘Hitomebore’. ‘X-Jigna’ heads 7 days earlier than ‘Hitomebore’ and its culm is 27 cm taller (Zewdu et al., 2025 ). The 200 RILs and the two parents were transplanted on 19 May 2023. Cool water was applied at 19.5 ± 0.4°C (mean ± SD) for 60 days from 45 to 104 DAT (Fig. S1 a); the water depth was kept at 20 cm until 50 DAT, and thereafter at 30 cm to fully expose the developing panicles to cold at the booting stage. The mean air temperature and solar radiation during this period were 26.1 ± 1.8°C (Fig. S1 a) and 17.9 ± 6.2 MJ m –2 (Fig. S1 c). Water temperature was recorded by data logger (TR52, T&D Corporation, Nagano, Japan), and air temperature and solar radiation at Morioka were obtained from the Japan Meteorological Agency. Basal fertilizer was applied at 50 kg N ha –1 , 22 kg P ha –1 , and 42 kg K ha –1 . After harvesting at 129 DAT, spikelet fertility was measured by backlighting against a fluorescent light (Treviewer A3-400, Trytech Co., Tokyo, Japan): spikelets that cast a shadow were classified as fertile, and those that did not as sterile. The spikelet fertility rate was calculated as the number of fertile spikelets per total spikelets × 100%. Seed shattering of the 846 F 5 RILs and the parents was evaluated in the field at Iwate University, Morioka, Iwate, Japan (39°42′N, 141°8′E). Seedlings were transplanted on 25 May 2023 at a spacing of 15 cm between plants in a single row, with 30 cm between rows, at 5 hills per line (Fig. S2e). Basal fertilizer was applied at 90 kg N ha –1 , 65 kg P ha –1 , and 75 kg K ha –1 before puddling. The irrigation water was maintained at a depth of ~ 5 cm. After harvest, seed shattering was measured by firmly grasping panicles by hand and counting the number of shattered grains (IRRI, 2013 ) from 1 panicle per line or 4 panicles per parent (Fig. S2f). The seed shattering rate was calculated as the number of shattered grains per total grains per panicle × 100%. Seven other agronomic traits were also measured: (1) days to heading (when 50% of the panicles started heading) in each plot; (2) culm length (from the soil surface to the panicle base) and (3) panicle length (from panicle base to tip) of 3 plants per replicate at maturity; and (4) panicle number, (5) biomass, (6) grain yield, and (7) harvest index of 10 plants per replicate, following the method of Takahashi et al. ( 2023 ). Mean air and water temperatures during the reproductive period (30 d before heading of ‘X-Jigna’) were 24.4 ± 1.6°C and 25.5 ± 1.0°C, respectively (Fig. S2a), and mean solar radiation was 16.7 ± 7.5 MJ m –2 d –1 (Fig. S2c). 2.1.2. QTL-seq analysis in Japan Leaf samples of the F 5 RILs in 2023 grown at Iwate University were collected from each line at 67 DAT. For QTL-seq analysis, based on the phenotyping data, each 40 RILs from each lower and upper extremes were identified from the 894 RILs, and the lower extremes denoted as L-bulk, and the upper extremes denoted as U-bulk, and bulk DNA extraction for each trait was conducted. Note that QTL-seq for spikelet fertility was done for top and bottom 20. Total genomic DNA was extracted from leaf blades using NucleoSpin Plant II Kit (Macherey-Nagel). Sequencing libraries were prepared using a Collibri ES DNA Library Prep Kit for Illumina Systems (Thermo Fisher Scientific, Waltham, MA, USA). The Libraries of the L-bulk and U-bulk for each trait were sequenced on a NovaSeq X Plus (Illumina, San Diego, CA, USA) instrument in 151-bp paired-end mode. QTL-seq analysis (Takagi et al., 2013 ) was performed using QTL-seq pipeline v2.2.8 (Sugihara et al, 2022 ) to identify genomic regions associated with the target trait. The differences in SNP indexes between the L-bulk and U-bulk were calculated using a pseudo-reference genome of ‘Hitomebore’, which was constructed by incorporating the SNPs of ‘Hitomebore’ into the IRGSP-1.0 reference genome. SNPs with a read depth of ≥ 10 were considered for SNP index analysis. A SNP index of 1 indicates that all short reads contain the allele from the cultivar ‘X-Jigna’, whereas a SNP index of 0 indicates that all short reads contain the reference allele from ‘Hitomebore’. A SNP index of 0.5 indicates equal contribution from both parental alleles at that position. The SNP index was plotted using a sliding window approach with a 2-Mb window and 1-kb step size for both bulks. The ΔSNP index, defined as the difference in SNP indexes between the two bulks, was calculated to detect genomic regions showing significant allele frequency differences. Statistical confidence intervals (p < 0.05 and p < 0.01) were determined to identify significant peaks, which were regarded as putative QTL regions associated with the traits. The identified QTL regions were checked from the Rice Annotation Project Database ( https://rapdb.dna.affrc.go.jp/ ) to confirm their novelty. 2.2. Confirmation of the detected QTLs in 2024 2.2.1. Phenotyping in Japan and Ethiopia Cold tolerance was again determined in Iwate in 2024 (there is no comparable facility in Ethiopia). We transplanted 98 F 6 RILs randomly selected from the same 846 lines and both parents on 23 May. These RILs included some of the 200 RILs grown in 2023. Cool water was applied at 19.3 ± 0.3°C (mean ± SD) for 54 days from 54 to 107 DAT (Fig. S1 b); the water depth was kept at 30 cm to fully expose the developing panicles to cold at booting. Mean air temperature and solar radiation during this period were 26.3 ± 1.7°C (Fig. S1 b) and 15.8 ± 6.6 MJ m –2 (Fig. S1 d). Seed shattering was confirmed in both Japan and Ethiopia in 2024. In Japan, the same 98 F 6 RILs as tested for cold tolerance plus the parents were transplanted on 23 May 2024 at Iwate University to confirm the QTLs detected by QTL-seq analysis. Each RIL had 3 replications in 4 rows with 7 hills per row (28 plants per replicate). On 4–19 September, 10 plants per replicate were harvested, and seed shattering of 1 panicle per line or parent cultivar was assessed. Air and water temperatures during the reproductive period (30 d before heading of ‘X-Jigna’) were 23.9 ± 1.9°C and 25.5 ± 1.0°C, respectively (Fig. S2b), and solar radiation was 18.6 ± 7.7 MJ m –2 d –1 (Fig. S2d). The other agronomic traits measured in 2023 were also measured. In Ethiopia, we transplanted 52 F 6 RILs randomly selected from the 98 RILs, plus the parents and two standard check cultivars, at the Fogera National Rice Research and Training Center: 15 plants each (3 rows × 5 plants) in 2 replications with a spacing of 25 cm between plants and 20 cm between rows (Fig. S3b). The September mean air temperatures during the reproductive stage (before heading of ‘X-Jigna’ on 2 October) were 23.1°C (mean average), 17.7°C (mean minimum), and 28.4°C (mean maximum) (Fig. S3a). No solar radiation data were available. Fertilizer was applied as per the local recommendations at 184 kg N ha –1 , 46 kg P ha –1 , and 8.5 kg S ha –1 . All P and S fertilizers and 42% of the N were applied basally 1 week after transplanting, followed by 29% of the N (as urea) at tillering stage and 29% at heading stage. Weeding and water management were done as needed. The seed shattering score of 1 panicle from 1 plant in each plot was assessed. The shattering score was based on IRRI ( 2013 ), which scored from 1 to 9, lower shattering to higher shattering. Seven other agronomic traits were also measured: (1) days to heading in each plot; (2) culm length and (3) panicle length of 5 plants per replicate at maturity; and (4) panicle number, (5) biomass, (6) grain yield, and (7) harvest index of 15 plants per replicate. 2.2.3. Marker analysis in Japan and Ethiopia Leaf samples of the 98 F 6 RILs grown at Iwate University, Japan, in 2024 were collected at 40 DAT of the maximum tillering stage. InDel markers were developed from the whole genome sequencing result following the method described by Natsume et al. ( 2023 ). In total, 22 markers were designed to validate 13 QTLs associated with 9 traits, with one to five markers per QTL (Table S1 ). Briefly, InDels polymorphic between the two parental cultivars were identified, and candidates with product sizes greater than 150 bp and GC content above 40% were retained. Subsequently, InDels located within the peak regions of ΔSNP-index values in the target chromosomes were chosen. The polymorphisms of these InDels between the two parents were examined by PCR. For polymorphism testing, a 10 µl PCR reaction containing 2µl template DNA, 0.4µl forward and reverse primers, 2.2µl distilled water, and 5µl Quick Taq was used. PCR conditions were as follows: 94°C for 3 min; 35 cycles of 94°C for 30 sec, 55°C for 30 sec, and 72°C for 30 sec. Amplicons were separated by electrophoresis on a 3% agarose gel (KANTO HC) at 100 V for 30 min, stained with 0.5 µg/ml ethidium bromide for 30 min, and visualized under ultraviolet light. Twenty-two markers showing clear polymorphism between the two parents were selected and used for genotyping the 98 RILs and their parents. Genotyping was conducted using a 5 µl PCR reaction (1µl template DNA, 0.025µl forward and reverse primers, 1.95µl distilled water, and 2µl Go Taq Green Master Mix) under the same thermal conditions as above. Single-marker analysis was performed to confirm the effects of the identified QTLs and to determine the most effective markers representing each QTL region. We did not genotype F 6 plants in Ethiopia, so we applied the Japanese genotyping data to both Japan and Ethiopia. The seeds of F 5 RILs were sent from Japan to Ethiopia in 2022, and the F 6 seeds were raised in Ethiopia in 2023. 3. Results 3.1. Cold tolerance at reproductive stage Under cold conditions, spikelet fertility was 90% in ‘Hitomebore’, 44% in ‘X-Jigna’, and 14% to 95% among the 200 RILs, with a mean of 69% (Fig. 1 a, b). QTL-seq analysis of the top 20 lines (U-bulk) and the bottom 20 lines (L-bulk) detected 3 significant QTLs: qSFC5 on chr. 5 (20.0–24.4 Mb) with an average ∆SNP index of − 0.481 (derived from ‘Hitomebore’), qSFC8 on chr. 8 (2.4–7.4 Mb; −0.615; ‘Hitomebore’), and qSFC10 on chr. 10 (20.7–21.9 Mb; 0.488; ‘X-Jigna’) (Fig. 1 c; Table 1 ; Fig. S4). Table 1 Summary of 20 identified QTLs for 9 agronomic traits of rice using 846 F 5 RILs developed from the cross between ‘X-Jigna’ × ‘Hitomebore’ in the QTL-seq approaches in Iwate, Japan, in 2023. Agronomic trait QTL Chr. No. Region (Mb) SNP index 1 SNP index 2 ΔSNP index Contributing parent Start End Spikelet fertility qSFC5 5 20.0 24.4 0.625 0.144 −0.481 Hitomebore qSFC8 8 2.4 7.4 0.683 0.068 −0.615 Hitomebore qSFC10 10 20.7 21.9 0.217 0.706 0.488 X-Jigna Seed shattering qSHT1 1 30.8 41.5 0.220 0.840 0.621 X-Jigna Days to heading qDH8 8 1.0 9.0 0.837 0.175 −0.662 Hitomebore Culm length qCL2 2 21.3 32.6 0.221 0.832 0.611 X-Jigna qCL3.1 3 18.0 21.6 0.249 0.865 0.616 X-Jigna qCL3.2 3 27.8 35.1 0.350 0.872 0.522 X-Jigna qCL8.1 8 1.3 5.1 0.746 0.206 −0.540 Hitomebore qCL8.2 8 23.1 25.7 0.144 0.771 0.627 X-Jigna Panicle length qPL3 3 33.1 34.6 0.306 0.834 0.529 X-Jigna qPL8 8 1.6 7.1 0.817 0.196 −0.621 Hitomebore qPL12 12 17.5 18.0 0.800 0.313 −0.488 Hitomebore Panicles per plant qPN2.1 2 8.1 8.4 0.711 0.284 −0.428 Hitomebore qPN2.2 2 16.1 17.1 0.737 0.309 −0.427 Hitomebore Biomass per plant qBMPL8 8 1.6 6.5 0.788 0.250 −0.538 Hitomebore qBMPL9 9 5.2 6.1 0.748 0.233 −0.516 Hitomebore Biomass per panicle qBMPA3 3 34.8 35.1 0.240 0.760 0.520 X-Jigna qBMPA8 8 2.2 5.2 0.750 0.220 −0.530 Hitomebore Harvest index qHI8 8 1.0 4.7 0.270 0.797 0.527 X-Jigna SNP index 1 and 2 refer to the lower and upper bulks; ΔSNP index gives the difference between the two bulks. In the confirmation study in 2024 (Fig. 1 d; Table 2 ), at qSFC5 , all 3 markers (M14-5-20554705, M24-5-20631657, and M25-5-20673667, 20.5–20.7 Mb) revealed significant phenotypic differences ( P < 0.01), with percentage of phenotypic variation explained (PVE) = 11%–14% between parental types. At qSFC8 , all 4 markers (M15-8-3642818, M16-8-4309379, M7-8-5179432, and M8-8-5378389, 3.6–5.4 Mb) revealed significant phenotypic differences ( P < 0.01), with PVE = 11%–13%. At qSFC10 , none of the 4 markers (21.0–22.6 Mb) revealed a significant difference, although the ‘X-Jigna’ allele indicated positive effects (Fig. 1 d) in agreement with the QTL-seq results (Fig. 1 c; Table 1 ). No spikelet sterility induced by low temperature was observed in Ethiopia, where the mean temperature during the reproductive stage was 23.1°C (Fig. S3a), higher than the threshold of 20°C (Shimono et al., 2005 ). Table 2 Confirmation of QTLs for nine agronomic traits of rice identified using the QTL-seq approach through indel marker genotyping using F 6 RILs at Iwate, Japan ( n = 98), and Fogera, Ethiopia ( n = 52) in 2024. Agronomic trait QTL Marker Iwate, Japan Fogera, Ethiopia Hitomebore type X-Jigna type PVE (%) Hitomebore type X-Jigna type PVE (%) Spikelet fertility qSFC5 M14-5-20554705 83.6 ± 1.0 76.1 ± 2.3 ** 10.6 – – – M24-5-20631657 83.4 ± 1.0 75.1 ± 2.4 ** 14.0 – – – M25-5-20673667 83.4 ± 1.0 75.4 ± 2.4 ** 11.9 – – – qSFC8 M15-8-3642818 84.4 ± 1.2 76.5 ± 1.8 *** 12.2 – – – M16-8-4309379 84.1 ± 1.2 76.3 ± 1.9 ** 12.4 – – – M7-8-5179432 83.8 ± 1.2 75.7 ± 1.9 *** 12.7 – – – M8-8-5378389 83.4 ± 1.1 76.7 ± 2.3 ** 10.5 – – – qSFC10 M13-10-21053134 78.6 ± 1.6 82.3 ± 1.5 ns 2.8 – – – M26-10-2114085 80.0 ± 2.1 80.6 ± 1.4 ns 0.1 – – – M10-10-22599813 78.9 ± 1.7 81.5 ± 1.6 ns 1.3 – – – M11-10-22600962 78.7 ± 1.8 81.6 ± 1.6 ns 1.6 – – – Seed shattering qSHT1 M1-1-29019501 40.2 ± 4.3 48.9 ± 5.9 ns 1.7 3.0 ± 0.5 4.0 ± 0.6 ns 3.5 M2-1-32576298 36.2 ± 4.0 52.7 ± 5.2 ** 6.5 3.0 ± 0.4 4.0 ± 0.6 ns 7.1 M22-1-34731315 27.5 ± 3.0 63.3 ± 5.1 *** 30.0 2.0 ± 0.4 4.0 ± 0.4 ** 19.7 M23-1-34895063 28.1 ± 3.0 64.0 ± 5.2 *** 29.9 2.0 ± 0.4 4.0 ± 0.6 ** 19 M12-1-35259855 28.4 ± 3.0 67.7 ± 5.2 *** 34.3 2.0 ± 0.4 5.0 ± 0.7 ** 18.6 Days to heading qDH8 M15-8-3642818 104.0 ± 0.6 98.0 ± 0.60 *** 36.5 103.0 ± 0.5 102.0 ± 0.7 ns 3.3 M16-8-4309379 105.0 ± 0.5 97.0 ± 0.47 *** 60.6 103.0 ± 0.4 102.0 ± 0.8 ns 2.9 M7-8-5179432 104.0 ± 0.5 97.0 ± 0.64 *** 40.0 103.0 ± 0.4 102.0 ± 1.0 ns 0.8 M8-8-5378389 103.0 ± 0.6 98.0 ± 0.74 *** 26.5 103.0 ± 0.4 102.0 ± 1.1 ns 1.4 Culm length qCL2 M4-2-21821646 96.4 ± 1.2 99.3 ± 1.10 ns 3.1 85.9 ± 1.0 86.3 ± 1.0 ns 0.2 M21-2-23976867 95.2 ± 1.2 100.8 ± 0.95 *** 12.3 85.0 ± 1.0 87.6 ± 1.1 ns 5.1 M20-2-26163645 95.2 ± 1.0 101.9 ± 1.12 *** 17.1 85.0 ± 1.1 87.6 ± 1.2 ns 5.1 M6-2-30732909 96.4 ± 1.1 100.4 ± 1.23 * 6.3 85.9 ± 1.2 87.1 ± 1.2 ns 1 qCL8.1 M15-8-3642818 100.4 ± 1.0 95.4 ± 1.18 ** 10.0 86.8 ± 1.0 85.1 ± 1.2 ns 2.2 M16-8-4309379 101.2 ± 1.1 93.9 ± 1.04 *** 20.1 86.9 ± 1.0 84.5 ± 1.3 ns 4 M7-8-5179432 100.4 ± 1.0 94.8 ± 1.15 *** 12.8 86.7 ± 0.9 85.2 ± 1.5 ns 1.5 qCL8.2 M32-8-23461108 98.2 ± 1.0 97.3 ± 1.11 ns 0.0 86.7 ± 1.2 86.1 ± 1.1 ns 0.003 Panicle length qPL8 M15-8-3642818 23.6 ± 0.2 22.3 ± 0.20 *** 20.5 21.0 ± 0.2 20.5 ± 0.2 ns 5.5 M16-8-4309379 23.8 ± 0.2 22.0 ± 0.16 *** 35.4 20.9 ± 0.2 20.6 ± 0.2 ns 2.4 M7-8-5179432 23.5 ± 0.2 22.2 ± 0.20 *** 19.3 20.5 ± 0.2 20.5 ± 0.2 ns 3.8 M8-8-5378389 23.4 ± 0.2 22.4 ± 0.20 ** 9.5 20.9 ± 0.2 20.6 ± 0.3 ns 1.6 Panicle number qPN2.1 M17-2-8236008 15.2 ± 0.2 15.0 ± 0.30 ns 0.4 9.6 ± 0.3 9.9 ± 0.3 ns 0.9 Biomass per plant qBMPL8 M15-8-3642818 68.9 ± 0.9 63.3 ± 0.97 *** 16.2 64.0 ± 1.6 68.0 ± 3.1 ns 3.9 M16-8-4309379 69.3 ± 0.8 62.1 ± 0.96 *** 26.0 65.0 ± 1.7 66.0 ± 1.6 ns 0.2 M7-8-5179432 68.0 ± 0.8 63.0 ± 1.11 *** 12.8 66.0 ± 1.6 67.0 ± 3.4 ns 0.4 M8-8-5378389 67.9 ± 0.9 63.3 ± 1.23 ** 10.3 66.0 ± 1.6 67.0 ± 4.1 ns 0.2 Biomass per panicle qBMPA8 M15-8-3642818 4.5 ± 0.0 4.3 ± 0.01 ns 1.5 4.5 ± 0.1 4.8 ± 0.2 ns 3.7 M16-8-4309379 4.5 ± 0.0 4.3 ± 0.01 * 5.7 4.6 ± 0.1 4.6 ± 0.2 ns 0.1 M7- 8-5179432 4.5 ± 0.0 4.3 ± 0.01 ns 2.5 4.6 ± 0.1 4.8 ± 0.3 ns 1.9 Harvest index qHI8 M15-8-3642818 0.6 ± 0.0 0.6 ± 0.01 ns 3.4 0.43 ± 0.01 0.42 ± 0.01 ns 0.3 M16-8-4309379 0.6 ± 0.0 0.6 ± 0.01 * 5.3 0.44 ± 0.01 0.42 ± 0.01 ns 2.9 Values are means ± SEM; PVE (%) = percentage of phenotypic variance explained as expressed by R 2 ; a total of 22 InDel marks developed in the peak regions of the QTL-seq result were used for confirmation of the QTL-seq result. 3.2. Seed shattering The rate of seed shattering of the 846 RILs in 2023 ranged from 4% to 100%, with a mean of 49% and a bimodal distribution, reflecting the rates of 19% in ‘Hitomebore’ and 99% in ‘X-Jigna’ (Fig. 1 a, b). QTL-seq analysis of the U-bulk and L-bulk detected a significant peak on chr. 1, at 30.8–41.5 Mb, indicating a significant major QTL, qSHT1 , with an average ∆SNP index of 0.621 (Fig. 2 c; Table 1 ; Fig. S5). This QTL was contributed by ‘X-Jigna’. In the confirmation study in 2024, in Japan, 4 out of 5 markers (M2-1-32576298, M22-1-34731315, M23-1-34895063, and M12-1-35259855, 34.7–35.2 Mb) revealed significant phenotypic differences ( P < 0.01), with PVE = 7%–34% (Fig. 2 d; Table 2 ). The other marker (M1-1-29019501, 29 Mb) indicated a positive effect of the ‘X-Jigna’ allele (Fig. 2 d) in agreement with the QTL-seq result (Fig. 1 c; Table 1 ). In Ethiopia, 3 out of the 5 markers revealed significant effects ( P < 0.01), with PVE = 19%–20% (Fig. 2 e; Table 2 ). The other 2 markers again indicated a positive effect (Fig. 2 e) in agreement with the QTL-seq result (Fig. 1 c; Table 1 ). The three markers at 34–35 Mb on chr. 1 confirmed in both Japan and Ethiopia would be suitable for breeding. 3.3. Other traits QTL-seq analysis for days to heading, culm length, biomass, grain yield, and harvest index (HI) found another 16 QTLs, bringing the total to 20 (Table 1 ). No significant QTL for grain yield was detected (Fig. S6). Among the QTLs for cold tolerance and seed shattering, only qSFC8 , for cold tolerance, coincided with QTLs for other traits (Fig. S7; Table 1 ), namely days to heading ( qDH8 ), culm length ( qCL8.1 ), panicle length ( qPL8 ), and biomass ( qBMPL8 , qBMPA8 ). Whole genome QTL-seq results of the traits are shown in Figures S8–S14. In the confirmation study, 11 of the 12 QTLs found in Japan in 98 RILs (including those for spikelet fertility (Fig. 1 d) and shattering (Fig. 2 d, e) but excluding qPN2.1 (Fig. S15b, d)) were confirmed by more than 1 marker each (Table 2 , Figs. S16b, d, S17b, d, S18b, d, S19b, d). But only 1 of the 9 QTLs found in Ethiopia in 52 RILs for seed shattering ( qSHT1 ) was confirmed (Fig. 2 e), although another 5 QTLs indicated agreement with QTL-seq results for days to heading ( qDH8 , Fig. S16f), culm length ( qCL2 , qCL8.1 , qCL8.2 , Fig. S17f), and panicle length ( qPL2 , Fig. S18f). On the other hand, allele effects of another 4 QTLs indicated opposite effects on biomass per plant ( qBMPL8 , Fig. S19f) and per panicle ( qBMPA8 , Fig. S20f) and HI ( qHI8 , Fig. S21f) compared to QTL-seq and the confirmation in Japan; i.e., the qBMPL8 and qBMPA8 alleles of ‘Hitomebore’ increased the biomass in Japan (Figs. S19b, d, S20b, d), in agreement with the QTL-seq results (Table 1 ), but decreased it in Ethiopia (Figs. S19f, S20f). Note that no significant QTL was detected (Fig. S6) and histogram of grain yield in Japan and Ethiopia was shown in Fig. S22. Our QTL-seq analysis identified QTLs for spikelet fertility under cold temperatures during the reproductive stage ( qSFC5 , qSFC8 , and qSFC10 ) and for seed shattering ( qSHT1 ) in the Ethiopian rice cultivar ‘X-Jigna’ (Table 1 ). To our knowledge, this is the first QTL mapping study of an Ethiopian rice cultivar grown in the field in different environments. For breeding by marker selection in Ethiopia, markers for both traits are critical. 4.1. Cold tolerance at reproductive stage Cold tolerance at the reproductive stage is an important trait in Ethiopia’s high-elevation, cool-climate zones. We found three QTLs for cold tolerance: the ‘Hitomebore’ qSFC5 and qSFC8 alleles and the ‘X-Jigna’ qSFC10 allele increase it (Fig. 1 c; Table 1 ). qSFC5 was detected at 20–24 Mb on chr. 5 and was confirmed by markers at 20.5–20.6 Mb (Fig. 1 d; Table 2 ). This region is similar to regions reported by Xu et al. ( 2008 ) (517 F 2 and 1517 F 3 RILs between tolerant ‘Kunmingxiaobaigu’ and sensitive ‘Towada’), Andaya and Mackill ( 2003 ) (191 F 6 RILs between tolerant ‘M-202’ and sensitive ‘IR50’), and Shimono et al. ( 2016 ) (208 F 7 and F 8 RILs between tolerant ‘Tohoku-PL3’ and sensitive ‘Akihikari’). Combined with the QTL and the transcriptome, Shimono et al. ( 2016 ) associated the region with candidate genes for O -methyltransferase ZRP4 (Os05g0515600, chr. 05:25569090..25570887) and β-1,3-glucanase-like protein (Os05g0535100, chr. 05:26570425..26573958) in a cross between ‘Tohoku-PL3’ and ‘Akihikari’. qSFC8 was detected at 2.4–7.4 Mb on chr. 8 (Fig. 1 c; Table 1 ) and was confirmed by markers at 3.6–5.4 Mb (Fig. 1 d; Table 2 ). This region is similar to regions reported by Kuroki et al. ( 2007 ) (F 3 and F 7 backcross inbred lines between tolerant ‘Hokkai PL’ [donor to ‘Hayahuki’ and ‘Padi Labou Alumbis’] and sensitive ‘Hokkai 287’) and Wainaina et al. ( 2018 ) (108 F 2 between tolerant ‘Hananomai’ and sensitive ‘WAB56-10’). qSFC8 overlapped several QTLs for heading date, culm length, and biomass (Table 2 ; Fig. S7). The qSFC8 region, which is large at ~ 5 Mb, would hold multiple genes, and/or some of which might have pleiotropic effects (Nutan et al., 2020 ). Map-based cloning and near-isogenic lines would be able to narrow down the position of the responsible gene. qSFC10 was detected at 20.7–21.9 Mb on chr. 10 (Fig. 1 c; Tables 1 ) in ‘X-Jigna’, the cold-sensitive parent, with indicative results (Fig. 1 d; Table 2 ). The region is close to a region for drought tolerance reported by Moncada et al. ( 2001 ) (274 BC 2 F 2 backcross inbred lines between ‘Caiapo’ [upland Oryza sativa ] and an accession of Oryza rufipogon ). The ‘X-Jigna’ region might prove useful in boosting cold tolerance. The method used to test for cold tolerance can control panicle temperature after initiation at the shoot base but not after emergence above the water level (Shimono et al., 2005 ), when it is exposed to atmospheric temperature. During 2023, the air temperature during the reproductive period was > 26°C (Fig. S1 a), higher than the lower threshold temperature of 20°C for inducing sterility (Shimono et al., 2005 , 2007 ). The difference in culm length and days to heading among RILs would not indirectly affect cold tolerance through the panicle temperature affected by atmospheric temperature. 4.2. Seed shattering Seed shattering is important to yield loss before harvest (Wang et al., 2024 ). We found a significant QTL on chr. 1, qSHT1 , at 30.8–41.5 Mb (Fig. 2 c; Table 1 ), and confirmed it at 34–35 Mb in both Japan and Ethiopia (Fig. 2 d, e; Table 2 ). qSH1 encodes a BEL1-type homeobox gene (Konishi et al., 2006 ; Wu et al., 2023 ), which causes seed shattering in rice ( Os01g0848400 , chr. 01:36 445 456..36 449 951). Our RILs showed a wide range of seed shattering habits (Fig. 2 b ), attributable to a haplotype mutation in this QTL region (Konishi et al., 2006 ; Wu et al., 2023 ). In Ethiopia, postharvest management practices are manual (Tadesse et al., 2019 ), so farmers rely on the easy-shattering characteristics, which cause significant yield losses. Future breeding in Ethiopia should seek to balance reducing yield losses with efficiency of threshing. 4.3. Future strategy for improving Ethiopian rice cultivars To improve the production of ‘X-Jigna’ in Ethiopia, phenotyping and genotyping studies should be conducted on-site. However, technical and management constraints made it necessary to do this in Japan and then confirm the detected QTLs in Ethiopia. This approach worked well for identifying QTLs for seed shattering ( qSHT1 ; Fig. 2 c, d, e), days to heading ( qDH8 ; Fig. S16f), culm length ( qCL2 , qCL8.1 , qCL8.2 ; Fig. S17f), and panicle length ( qPL8 ; Fig. S18f). DNA markers for these traits would be useful to develop new cultivars by marker-assisted selection in Ethiopia. However, the important quantitative traits of biomass production ( qBMPL8 , qBMPA8 ; Figs. S19b, d, f, S20b, d, f) and HI ( qHI8 ; Fig. S21f) were confirmed in Japan but not in Ethiopia, where the QTLs indicated the opposite effect. This suggests that the strategy is useful for traits controlled by a few dominant genes, as for seed shattering and culm length, but not for complex quantitative traits of biomass and yield, which must be phenotyped on-site in Ethiopia. However, it should be noted that developing countries face limitations on access to phenotyping facilities, such as the cold screening facility used in this study. In current condition, the present strategy of the multi-country studies can compensate for each others’ deficiencies. 4. Discussion Our QTL-seq analysis identified QTLs for spikelet fertility under cold temperatures during the reproductive stage ( qSFC5 , qSFC8 , and qSFC10 ) and for seed shattering ( qSHT1 ) in the Ethiopian rice cultivar ‘X-Jigna’ (Table 1). To our knowledge, this is the first QTL mapping study of an Ethiopian rice cultivar grown in the field in different environments. For breeding by marker selection in Ethiopia, markers for both traits are critical. 4.1. Cold tolerance at reproductive stage Cold tolerance at the reproductive stage is an important trait in Ethiopia’s high-elevation, cool-climate zones. We found three QTLs for cold tolerance: the ‘Hitomebore’ qSFC5 and qSFC8 alleles and the ‘X-Jigna’ qSFC10 allele increase it (Fig. 1c; Table 1). qSFC5 was detected at 20–24 Mb on chr. 5 and was confirmed by markers at 20.5–20.6 Mb (Fig. 1d; Table 2). This region is similar to regions reported by Xu et al. (2008) (517 F 2 and 1517 F 3 RILs between tolerant ‘Kunmingxiaobaigu’ and sensitive ‘Towada’), Andaya and Mackill (2003) (191 F 6 RILs between tolerant ‘M-202’ and sensitive ‘IR50’), and Shimono et al. (2016) (208 F 7 and F 8 RILs between tolerant ‘Tohoku-PL3’ and sensitive ‘Akihikari’). Combined with the QTL and the transcriptome, Shimono et al. (2016) associated the region with candidate genes for O -methyltransferase ZRP4 (Os05g0515600, chr. 05:25569090..25570887) and β-1,3-glucanase-like protein (Os05g0535100, chr. 05:26570425..26573958) in a cross between ‘Tohoku-PL3’ and ‘Akihikari’. qSFC8 was detected at 2.4–7.4 Mb on chr. 8 (Fig. 1c; Table 1) and was confirmed by markers at 3.6–5.4 Mb (Fig. 1d; Table 2). This region is similar to regions reported by Kuroki et al. (2007) (F 3 and F 7 backcross inbred lines between tolerant ‘Hokkai PL’ [donor to ‘Hayahuki’ and ‘Padi Labou Alumbis’] and sensitive ‘Hokkai 287’) and Wainaina et al. (2018) (108 F 2 between tolerant ‘Hananomai’ and sensitive ‘WAB56-10’). qSFC8 overlapped several QTLs for heading date, culm length, and biomass (Table 2; Fig. S7). The qSFC8 region, which is large at ~5 Mb, would hold multiple genes, and/or some of which might have pleiotropic effects (Nutan et al., 2020). Map-based cloning and near-isogenic lines would be able to narrow down the position of the responsible gene. qSFC10 was detected at 20.7–21.9 Mb on chr. 10 (Fig. 1c; Tables 1) in ‘X-Jigna’, the cold-sensitive parent, with indicative results (Fig. 1d; Table 2). The region is close to a region for drought tolerance reported by Moncada et al. (2001) (274 BC 2 F 2 backcross inbred lines between ‘Caiapo’ [upland Oryza sativa ] and an accession of Oryza rufipogon ). The ‘X-Jigna’ region might prove useful in boosting cold tolerance. The method used to test for cold tolerance can control panicle temperature after initiation at the shoot base but not after emergence above the water level (Shimono et al., 2005), when it is exposed to atmospheric temperature. During 2023, the air temperature during the reproductive period was >26 °C (Fig. S1a), higher than the lower threshold temperature of 20 °C for inducing sterility (Shimono et al., 2005, 2007). The difference in culm length and days to heading among RILs would not indirectly affect cold tolerance through the panicle temperature affected by atmospheric temperature. 4.2. Seed shattering Seed shattering is important to yield loss before harvest (Wang et al., 2024). We found a significant QTL on chr. 1, qSHT 1 , at 30.8–41.5 Mb (Fig. 2c; Table 1), and confirmed it at 34–35 Mb in both Japan and Ethiopia (Fig. 2d, e; Table 2). qSH1 encodes a BEL1-type homeobox gene (Konishi et al., 2006; Wu et al., 2023), which causes seed shattering in rice ( Os01g0848400 , chr. 01:36 445 456..36 449 951). Our RILs showed a wide range of seed shattering habits (Fig. 2b ), attributable to a haplotype mutation in this QTL region (Konishi et al., 2006; Wu et al., 2023). In Ethiopia, postharvest management practices are manual (Tadesse et al., 2019), so farmers rely on the easy-shattering characteristics, which cause significant yield losses. Future breeding in Ethiopia should seek to balance reducing yield losses with efficiency of threshing. 4.3. Future strategy for improving Ethiopian rice cultivars To improve the production of ‘X-Jigna’ in Ethiopia, phenotyping and genotyping studies should be conducted on-site. However, technical and management constraints made it necessary to do this in Japan and then confirm the detected QTLs in Ethiopia. This approach worked well for identifying QTLs for seed shattering ( qSHT1 ; Fig. 2c, d, e), days to heading ( qDH8 ; Fig. S16f), culm length ( qCL2 , qCL8.1 , qCL8.2 ; Fig. S17f), and panicle length ( qPL8 ; Fig. S18f). DNA markers for these traits would be useful to develop new cultivars by marker-assisted selection in Ethiopia. However, the important quantitative traits of biomass production ( qBMPL8 , qBMPA8 ; Figs. S19b, d, f, S20b, d, f) and HI ( qHI8 ; Fig. S21f) were confirmed in Japan but not in Ethiopia, where the QTLs indicated the opposite effect. This suggests that the strategy is useful for traits controlled by a few dominant genes, as for seed shattering and culm length, but not for complex quantitative traits of biomass and yield, which must be phenotyped on-site in Ethiopia. However, it should be noted that developing countries face limitations on access to phenotyping facilities, such as the cold screening facility used in this study. In current condition, the present strategy of the multi-country studies can compensate for each others’ deficiencies. Declarations Author Contribution CRediT statement: Zelalem Zewdu: Writing – original draft preparation, data curation; Mulugeta Atnaf: Methodology, writing – review & editing; Assaye Berie: data curation; Honoka Takasago: data curation; Reo Sato: data curation; Ryuji Ishikawa: writing – review & editing; Akira Abe: data curation, methodology, writing – review & editing; Maya Matsunam: writing – review & editing; Hiroyuki Shimono: conceptualization, supervision, methodology, writing – review & editing Acknowledgments Zelalem Zewdu is grateful for the financial support of the Japan International Cooperation Agency (JICA) through its EthioRice-2 project. The authors thank the staff of the national rice research program at the Fogera National Rice Research and Training Center for managing the experiment; of the Iwate University Crop Science laboratory for their support in data collection at Iwate; and of the Iwate Biotechnology Research Center laboratory for assisting with the QTL-seq analysis and genotyping. We thank the Iwate Prefectural Research Center for providing the cold screening facilities. CRediT statement: Zelalem Zewdu : Writing – original draft preparation, data curation; Mulugeta Atnaf : Methodology, writing – review & editing; Assaye Berie : data curation; Honoka Takasago : data curation; Reo Sato : data curation; Ryuji Ishikawa : writing – review & editing; Akira Abe : data curation, methodology, writing – review & editing; Maya Matsunam : writing – review & editing; Hiroyuki Shimono : conceptualization, supervision, methodology, writing – review & editing References Alemayehu, H.A., Dumbuya, G., Hasan, M., Tadesse, T., Nakajyo, S., Fujioka, T., Abe, A., Matsunami, M., Shimono, H., 2021. Genotypic variation in cold tolerance of 18 Ethiopian rice cultivars in relation to their reproductive morphology. Field Crops Res. 262, 108042. Andaya, V.C., Mackill, D.J., 2003. QTLs conferring cold tolerance at the booting stage of rice using recombinant inbred lines from a japonica × indica cross. Theor. Appl. Genet. 106, 1084–1090. Beyene, A.M., Gashu, A.T., Tegegne, M.A., Anteneh Mihertie, A., 2022. Is the longstanding local rice cultivar ‘X-Jigna’ being replaced by the improved variety ‘Shaga’ in Fogera Plain, northwest Ethiopia? Cogent Econ. Fin. 10, 2145748. Dessie, A., 2020. Rice breeding achievements, potential and challenges in Ethiopia. International Journal of Research Studies in Agricultural Sciences 6, 35–42. FAO, 2024. FAOSTAT (http://faostat.fao.org/site/291/default.aspx). Food and Agriculture Organization of the United Nations. Farrell, T.C., Fox, K.M., Williams, R.L., Fukai, S., 2006. Genotypic variation for cold tolerance during reproductive development in rice: Screening with cold air and cold water. Field Crops Res. 98, 178–194. IRRI, 2013. Standard evaluation systems for rice, 5th ed. International Rice Research Institute, Manila, Philippines. Ito, N., Hayase, H., Satake, T., Nishiyama, I., 1970. Male sterility caused by cooling treatment at the meiotic stage in rice plants. III. Male abnormalities at anthesis. Proc. Crop Sci. Soc. Jpn. 39, 60–64. Jeong, J.-M., Mo, Y., Hyun, U.-J., Jeung, J.-U., 2020. Identification of quantitative trait loci for spikelet fertility at the booting stage in rice ( Oryza sativa L.) under different low-temperature conditions. Agronomy 10. Konishi, S., Izawa, T., Lin, S.Y., Ebana, K., Fukuta, Y., Sasaki, T., Yano, M., 2006. An SNP caused loss of seed shattering during rice domestication. Science 312, 1392–1396. Kuroki, M., Saito, K., Matsuba, S., Yokogami, N., Shimizu, H., Ando, I., Sato, Y., 2007. A quantitative trait locus for cold tolerance at the booting stage on rice chromosome 8. Theor. Appl. Genet. 115, 593–600. Li, C., Zhou, A., Sang, T., 2006. Rice domestication by reducing shattering. Science 311, 1936–1939. Li, J., Zeng, Y., Pan, Y., Zhou, L., Zhang, Z., Guo, H., Lou, Q., Shui, G., Huang, H., Tian, H., et al., 2021. Stepwise selection of natural variations at CTB2 and CTB4a improves cold adaptation during domestication of japonica rice. New Phytologist 231, 1056–1072. Matsunaga, K., 2005. Establishment of an evaluation method for cold tolerance at the booting stage of rice using deep water irrigation system and development of highly cold tolerant rice varieties by combining cold tolerance genes. Bull. Miyagi Furukawa Agric. Exp. Sta. 4, 1–78 (Japanese with English abstract). Ministry of Agriculture and Rural Development, 2010. National rice research and development strategy of Ethiopia., Ethiopia, p. 61. Moncada, P., Martinez, C.P., Borrero, J., Chatel, M., Gauch Jr, H., Guimaraes, E., Tohme, J., McCouch, S.R., 2001. Quantitative trait loci for yield and yield components in an Oryza sativa × Oryza rufipogon BC 2 F 2 population evaluated in an upland environment. Theor. Appl. Genet. 102, 41–52. Natsume, S., Oikawa, K., Nomura, C., Ito, K., Utsushi, H., Shimizu, M., Terauchi, R., Abe, A., 2023. V-primer: software for the efficient design of genome-wide indel and SNP markers from multi-sample variant call format (VCF) genotyping data. Breed. Sci. 73, 415–420. Nutan KK, Singla-Pareek SL, Pareek A. 2020. The Saltol QTL-localized transcription factor OsGATA8 plays an important role in stress tolerance and seed development in Arabidopsis and rice. J. Exp. Bot. 71: 684–698. Oliver, S.N., Dennis, E.S., Dolferus, R., 2007. ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice. Plant Cell Physiol. 48, 1319–1330. Oliver, S.N., Van Dongen, J.T., Alfred, S.C., Mamun, E.A., Zhao, X., Saini, H.S., Fernandes, S.F., Blanchard, C.L., Sutton, B.G., Geigenberger, P., Dennis, E.S., Dolferus, R., 2005. Cold-induced repression of the rice anther-specific cell wall invertase gene OSINV4 is correlated with sucrose accumulation and pollen sterility. Plant Cell Env. 28, 1534–1551. Sakata, T., Oda, S., Tsunaga, Y., Shomura, H., Kawagishi-Kobayashi, M., Aya, K., Saeki, K., Endo, T., Nagano, K., Kojima, M., Sakakibara, H., Watanabe, M., Matsuoka, M., Higashitani, A., 2014. Reduction of gibberellin by low temperature disrupts pollen development in rice. Plant Physiol. 164, 2011–2019. Satake, T., 1976. Determination of the most sensitive stage to sterile-type cool injury in rice plants. Res. Bull. Hokkaido Nat. Agric. Exp. Stn. 113, 1–44. Sato, Y., Masuta, Y., Saito, K., Murayama, S., Ozawa, K., 2011. Enhanced chilling tolerance at the booting stage in rice by transgenic overexpression of the ascorbate peroxidase gene, OsAPXa. Plant Cell Reports 30, 399–406. Shimono, H., 2018. Male sterility induced by low temperature at the booting stage in rice. Jpn. J. Crop Sci. 87, 113–124 (Japanese with English abstract). Shimono, H., Hasegawa, T., Iwama, K., 2002. Response of growth and grain yield in paddy rice to cool water at different growth stages. Field Crops Res. 73, 67–79. Shimono, H., Hasegawa, T., Moriyama, M., Fujimura, S., Nagata, T., 2005. Modeling spikelet sterility induced by low temperature in rice. Agron. J. 97, 1524–1536. Shimono, H., Okada, M., Kanda, E., Arakawa, I., 2007. Low temperature-induced sterility in rice: Evidence for the effects of temperature before panicle initiation. Field Crops Res. 101, 221–231. Shimono, H., Abe, A., Aoki, N., Koumoto, T., Sato, M., Yokoi, S., Kuroda, E., Endo, T., Saeki, K., Nagano, K., 2016. Combining mapping of physiological quantitative trait loci and transcriptome for cold tolerance for counteracting male sterility induced by low temperatures during reproductive stage in rice. Physiol. Plant. 157, 175–192. Sugihara, Y., Young, L., Yaegashi, H., Natsume, S., Shea, D.J., Takagi, H., Booker, H., Innan, H., Terauchi, R., Abe, A., 2022. High-performance pipeline for MutMap and QTL-seq. . PeerJ 10, e13170 https://doi.org/13110.17717/peerj.13170. Suzuki, K., Aoki, N., Matsumura, H., Okamura, M., Ohsugi, R., Shimono, H., 2015. Cooling water before panicle initiation increases chilling-induced male sterility and disables chilling-induced expression of genes encoding OsFKBP65 and heat shock proteins in rice spikelets. Plant Cell Env. 38, 1255–1274. Tadesse, T., Atnaf, M., Alemu, D., Tadesse, T., Shiratori, K., 2019. Advances in rice research and development in Ethiopia. Ethiopian Institute of Agricultural Research. Takagi, H., Abe, A., Yoshida, K., Kosugi, S., Natsume, S., Mitsuoka, C., Uemura, A., Utsushi, H., Tamiru, M., Takuno, S., Innan, H., Cano, L.M., Kamoun, S., Terauchi, R., 2013. QTL-seq: rapid mapping of quantitative trait loci in rice by whole genome resequencing of DNA from two bulked populations. Plant J. 74, 174–183. Takahashi T, Sato N, Matsunami M, et al., 2023: Yield performance of hybrid rice in a cool climate in Japan. Field Crops Res. 291, 1–6. Wainaina, C.M., Makihara, D., Nakamura, M., Ikeda, A., Suzuki, T., Mizukami, Y., Nonoyama, T., Doi, K., Kikuta, M., Samejima, H., et al., 2018. Identification and validation of QTLs for cold tolerance at the booting stage and other agronomic traits in a rice cross of a Japanese tolerant variety, Hananomai, and a NERICA parent, WAB56–104. Plant Prod. Sci. 21, 132–143. Wang, Y., Wu, W., Xu, J., Wang, Y., Wu, Z., Liu, H., 2024. Expounding the effect of harvest management on rice ( Oryza sativa L.) yield and latent loss based on the accurate measurement of grain data. Agronomy 14, 1346. Wu, H., He, Q., Wang, Q., 2023. Advances in rice seed shattering. Int. J. Mol. Sci. 24, 8889. Xu, L.M., Zhou, L., Zeng, Y.W., Wang, F.M., Zhang, H.L., Shen, S.Q., Li, Z.C., 2008. Identification and mapping of quantitative trait loci for cold tolerance at the booting stage in a japonica rice near-isogenic line. Plant Science 174, 340–347. Yang, L., Lei, L., Wang, J., Zheng, H., Xin, W., Liu, H., Zou, D., 2023. qCTB7 positively regulates cold tolerance at booting stage in rice. Theor. Appl. Genet. 136, 135. Zewdu, Z., Atnaf, M., Sato, R., Ishikawa, R., Abe, A., Matsunami, M., Shimono, H., 2025. Comparison of rice yield potential in Ethiopia and in northern Japan: an experimental and modeling study. Journal of Agricultural Meteorology 81, 36–43. Additional Declarations No competing interests reported. Supplementary Files SupplementrayFig.S1S22.pdf Supplementaltables.docx Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 12 Mar, 2026 Reviews received at journal 11 Jan, 2026 Reviews received at journal 04 Jan, 2026 Reviewers agreed at journal 30 Dec, 2025 Reviewers agreed at journal 22 Dec, 2025 Reviewers invited by journal 22 Dec, 2025 Editor assigned by journal 03 Dec, 2025 Submission checks completed at journal 03 Dec, 2025 First submitted to journal 30 Nov, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8244416","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":564393804,"identity":"b0940076-73a4-4289-88b6-1a630bd1051c","order_by":0,"name":"Zelalem Zewdu","email":"","orcid":"","institution":"Iwate University","correspondingAuthor":false,"prefix":"","firstName":"Zelalem","middleName":"","lastName":"Zewdu","suffix":""},{"id":564393806,"identity":"c6ebc1cf-6cca-407b-8aa3-fe712f1ccec3","order_by":1,"name":"Mulugeta Atnaf","email":"","orcid":"","institution":"Ethiopian Institute of Agricultural Research, Fogera National Rice Research \u0026 Training Center","correspondingAuthor":false,"prefix":"","firstName":"Mulugeta","middleName":"","lastName":"Atnaf","suffix":""},{"id":564393807,"identity":"7b564b54-af13-4928-a344-2c88a0bad026","order_by":2,"name":"Assaye Berie","email":"","orcid":"","institution":"Ethiopian Institute of Agricultural Research, Fogera National Rice Research \u0026 Training Center","correspondingAuthor":false,"prefix":"","firstName":"Assaye","middleName":"","lastName":"Berie","suffix":""},{"id":564393809,"identity":"4e0de5bc-0b8c-443c-8589-d638d324fc88","order_by":3,"name":"Honoka Takasago","email":"","orcid":"","institution":"Iwate Prefectural Agricultural Research Center","correspondingAuthor":false,"prefix":"","firstName":"Honoka","middleName":"","lastName":"Takasago","suffix":""},{"id":564393811,"identity":"51f0d9fc-749d-496b-954f-5fd694a71d19","order_by":4,"name":"Reo Sato","email":"","orcid":"","institution":"Iwate 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16:14:47","extension":"html","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":199782,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/0a972562ed167f2dbafbb628.html"},{"id":98963478,"identity":"4c50a729-35f0-4526-abdb-ca520f77f447","added_by":"auto","created_at":"2025-12-24 18:35:08","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":303770,"visible":true,"origin":"","legend":"\u003cp\u003eQTL identification for rice spikelet fertility loci under cold conditions and their confirmation through InDel marker genotyping. Spikelet fertility of parents under pot conditions (a), histograms of 200 F\u003csub\u003e5\u003c/sub\u003e rice RILs developed from ‘X-Jigna’ and ‘Hitomebore’ for spikelet fertility in 2023 (b), spikelet fertility QTLs (\u003cem\u003eqSFC5\u003c/em\u003e, \u003cem\u003eqSFC8\u003c/em\u003e, and \u003cem\u003eqSFC10\u003c/em\u003e) (c), confirmation of identified QTLs through InDel marker genotyping of 98 F\u003csub\u003e6\u003c/sub\u003e RILs in Iwate, Japan (d) in 2024, L-bulk = lower 20 RILs, U-bulk = upper 20 RILs\u003c/p\u003e","description":"","filename":"Figures1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/07358310d2d110b865c0a881.jpg"},{"id":98963479,"identity":"88ae03f3-6b99-4534-a11b-a253f8c9f62a","added_by":"auto","created_at":"2025-12-24 18:35:08","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":257662,"visible":true,"origin":"","legend":"\u003cp\u003eQTL identification for rice seed shattering and its confirmation through InDel marker genotyping. Low shattering type and high shattering type (a), histograms of 846 F\u003csub\u003e5\u003c/sub\u003e rice RILs developed from ‘X-Jigna’ and ‘Hitomebore’ for spikelet fertility in 2023 (b), seed shattering QTL on chromosome-1 (\u003cem\u003eqSHT1\u003c/em\u003e) (c), confirmation of \u003cem\u003eqSHT1\u003c/em\u003e in Iwate, Japan, 98 F\u003csub\u003e6\u003c/sub\u003e RILs (d), and Fogera, Ethiopia, using 52 F\u003csub\u003e6\u003c/sub\u003e RILs (e) in 2024, L-bulk = lower 40 RILs, U-bulk = upper 40 RILs\u003c/p\u003e","description":"","filename":"Figures2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/2691e8f633f90e1f377d9c37.jpg"},{"id":99322854,"identity":"672878cf-2b5b-4ecb-bb19-4fbc8a814ae8","added_by":"auto","created_at":"2025-12-31 16:44:26","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2043014,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/3df92b87-2105-4310-91ba-2eaa4e2717b5.pdf"},{"id":99311386,"identity":"b8552537-b860-4ad8-9fda-474fb20037bb","added_by":"auto","created_at":"2025-12-31 16:14:44","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":7032145,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementrayFig.S1S22.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/4a9ca7fbc9745de46f5fed5a.pdf"},{"id":98963480,"identity":"3a5ee51f-9cf6-426f-b067-f7d6360315ab","added_by":"auto","created_at":"2025-12-24 18:35:08","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":19213,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementaltables.docx","url":"https://assets-eu.researchsquare.com/files/rs-8244416/v1/e89728b9fa960ffe6a780870.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Identification of loci for cold tolerance at reproductive stage and seed shattering in Ethiopian rice cultivar ‘X-Jigna’","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eRice consumption in Ethiopia has increased 10\u0026times; since 2010, to \u0026gt;\u0026thinsp;2.5 Mt annually in 2021 (FAO, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). It has become a challenge to meet this demand solely through domestic production, which is \u0026lt;\u0026thinsp;10% of the self-sufficiency rate in spite of 30\u0026nbsp;million hectares of potential cropland.\u003c/p\u003e \u003cp\u003eIn Ethiopia, rice is grown mostly in high-elevation regions above 1800 m in altitude, notably on the Fogera Plain, which accounts for 71% of the total rice production in the country (Ministry of Agriculture and Rural Development, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Owing to the high elevation, low temperatures place a major constraint on rice production (Dessie, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zewdu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e): the minimum temperature drops to 13\u0026deg;C during the reproductive stage, which is the most sensitive period in terms of induction of sterility and severe yield losses (Satake \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Shimono et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe rice cultivar \u0026lsquo;X-Jinga\u0026rsquo; is the Ethiopian farmers\u0026rsquo; favorite cultivar, and was grown on \u0026gt;\u0026thinsp;80% of the Fogera area in 2021 (Beyene et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Introduced into Ethiopia in the 1980s from North Korea, it is favored for its high yield, its ability to make good-quality \u003cem\u003einjera\u003c/em\u003e, a staple food traditionally made from teff (\u003cem\u003eEragrostis tef\u003c/em\u003e) flour, and its high biomass and long culm for forage. However, \u0026lsquo;X-Jigna\u0026rsquo; is highly susceptible to cold stress during the reproductive stage, which induces spikelet sterility, unlike in the Japanese cold-tolerant elite cultivar \u0026lsquo;Hitomebore\u0026rsquo; (Alemayehu et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zewdu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIdentification of DNA markers may facilitate the development of suitable rice genotypes through marker-assisted selection. The processes involved in cold damage during fertilization are complex. The molecular and physiological aspects of damage to male gametes are controlled by many parallel and interacting processes (Shimono, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), including metabolic processes related to carbohydrates (Oliver et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), amino acids (Ito et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1970\u003c/span\u003e), reactive oxygen species (Suzuki et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), heat-shock proteins (Sato et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Suzuki et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), and plant hormones such as abscisic acid (Oliver et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and gibberellins (Sakata et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). It is important to identify the genomic regions causing genotypic differences in the weak cold tolerance of \u0026lsquo;X-Jigna\u0026rsquo; for future breeding. The cold-tolerant \u0026lsquo;Hitomebore\u0026rsquo; may hold novel quantitative trait loci (QTLs). While a number of QTLs for cold tolerance have been identified in various cultivars (Andaya and Mackill, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Kuroki et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Xu et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Shimono et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), no study has tested Ethiopian cultivars.\u003c/p\u003e \u003cp\u003eAnother shortcoming of \u0026lsquo;X-Jigna\u0026rsquo; is its easy seed shattering habit (Zewdu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), which decreases the yield before harvest (Wang et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The habit is controlled by the formation of an abscission layer in the joint between the lemma and pedicel (Wu et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Many QTLs related to seed shattering have been reported and identified through the use of various genetic populations and linkage maps (Wu et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u003cem\u003eqSH1\u003c/em\u003e, on chromosome (chr.) 1 (Konishi et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), and \u003cem\u003esh4\u003c/em\u003e, on chr. 4 (Li et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), are dominant QTLs that control formation of the abscission layer, as confirmed in many genetic populations (Wu et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Introduction of the equivalent genomic region of \u0026lsquo;Hitomebore\u0026rsquo; should reduce yield loss. However, the farmers\u0026rsquo; postharvest management practices are closely linked to the shattering habit: the harvested plants are threshed by hand using sticks or with cattle (Tadesse et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The farmers thus prefer the easy shattering habit, despite the significant yield losses. Balancing the reduction of yield loss with the efficient management of threshing would improve yields. Again, however, no QTL studies of seed shattering in Ethiopian cultivars have been conducted.\u003c/p\u003e \u003cp\u003eThe purpose of this study was to identify QTLs related to cold tolerance at the reproductive stage and to seed shattering in \u0026lsquo;X-Jigna\u0026rsquo; and \u0026lsquo;Hitomebore\u0026rsquo;. We conducted QTL-seq analysis of bulk segregants (Takagi et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) using 846 RIL F\u003csub\u003e5\u003c/sub\u003e populations of \u0026lsquo;X-Jigna\u0026rsquo; \u0026times; \u0026lsquo;Hitomebore\u0026rsquo; in Japan in 2023 (Experiment 1), and confirmed our results in both Japan and Ethiopia using F\u003csub\u003e6\u003c/sub\u003e populations in 2024 (Experiment 2). Compared with standard QTL analysis using genotyping data of all populations, QTL-seq analysis is rapid and requires less work for identifying QTLs, reducing the time and labor needed without genotyping individual lines, and has proven effective in identifying QTLs rapidly and cost-effectively in different crop species for various target traits (Takagi et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. QTL-seq analysis for detecting QTLs in 2023\u003c/h2\u003e \u003cdiv id=\"Sec4\" class=\"Section3\"\u003e \u003ch2\u003e2.1.1. Phenotyping in Japan\u003c/h2\u003e \u003cp\u003eCold tolerance for inducing spikelet sterility by low temperature at reproductive stage was determined at a field-based cool-irrigation facility at the Iwate Prefecture Agricultural Research Center, Kitakami, Iwate, Japan (39\u0026deg;35\u0026prime;N, 141\u0026deg;11\u0026prime;E). The facility is commonly used for screening cold tolerance in breeding. The method maintains water depth at \u0026gt;\u0026thinsp;20 cm to cover the developing panicles uniformly so as to minimize errors caused by difference in rates of maturity, and controls the temperature of the developing panicles (Matsunaga, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Shimono et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Cool water sourced from aquifers or refrigeration units is combined with warmer irrigation water to achieve a constant 18\u0026ndash;19\u0026deg;C, which is based on the cold tolerance of the population under test (Matsunaga, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The method can control the temperature of the developing panicle uniformly under water before culm elongation especially including the most sensitive microspore and booting stage (Satake, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1976\u003c/span\u003e), although it can not control that of the developing panicle after it has emerged above the water (Shimono et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The method has been used in Japan since 1929 (Matsunaga, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Shimono et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and is now widely used in Korea (Jeong et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), China (Li et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Yang et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and Australia (Farrell et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFrom 846 F\u003csub\u003e5\u003c/sub\u003e RILs, we randomly selected 200 RILs with similar culm lengths and days to heading as \u0026lsquo;Hitomebore\u0026rsquo;. \u0026lsquo;X-Jigna\u0026rsquo; heads 7 days earlier than \u0026lsquo;Hitomebore\u0026rsquo; and its culm is 27 cm taller (Zewdu et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The 200 RILs and the two parents were transplanted on 19 May 2023. Cool water was applied at 19.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u0026deg;C (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD) for 60 days from 45 to 104 DAT (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003ea); the water depth was kept at 20 cm until 50 DAT, and thereafter at 30 cm to fully expose the developing panicles to cold at the booting stage. The mean air temperature and solar radiation during this period were 26.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8\u0026deg;C (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003ea) and 17.9\u0026thinsp;\u0026plusmn;\u0026thinsp;6.2 MJ m\u003csup\u003e\u0026ndash;2\u003c/sup\u003e (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003ec). Water temperature was recorded by data logger (TR52, T\u0026amp;D Corporation, Nagano, Japan), and air temperature and solar radiation at Morioka were obtained from the Japan Meteorological Agency. Basal fertilizer was applied at 50 kg N ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, 22 kg P ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, and 42 kg K ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e. After harvesting at 129 DAT, spikelet fertility was measured by backlighting against a fluorescent light (Treviewer A3-400, Trytech Co., Tokyo, Japan): spikelets that cast a shadow were classified as fertile, and those that did not as sterile. The spikelet fertility rate was calculated as the number of fertile spikelets per total spikelets \u0026times; 100%.\u003c/p\u003e \u003cp\u003eSeed shattering of the 846 F\u003csub\u003e5\u003c/sub\u003e RILs and the parents was evaluated in the field at Iwate University, Morioka, Iwate, Japan (39\u0026deg;42\u0026prime;N, 141\u0026deg;8\u0026prime;E). Seedlings were transplanted on 25 May 2023 at a spacing of 15 cm between plants in a single row, with 30 cm between rows, at 5 hills per line (Fig. S2e). Basal fertilizer was applied at 90 kg N ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, 65 kg P ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, and 75 kg K ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e before puddling. The irrigation water was maintained at a depth of ~\u0026thinsp;5 cm. After harvest, seed shattering was measured by firmly grasping panicles by hand and counting the number of shattered grains (IRRI, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) from 1 panicle per line or 4 panicles per parent (Fig. S2f). The seed shattering rate was calculated as the number of shattered grains per total grains per panicle \u0026times; 100%. Seven other agronomic traits were also measured: (1) days to heading (when 50% of the panicles started heading) in each plot; (2) culm length (from the soil surface to the panicle base) and (3) panicle length (from panicle base to tip) of 3 plants per replicate at maturity; and (4) panicle number, (5) biomass, (6) grain yield, and (7) harvest index of 10 plants per replicate, following the method of Takahashi et al. (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Mean air and water temperatures during the reproductive period (30 d before heading of \u0026lsquo;X-Jigna\u0026rsquo;) were 24.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u0026deg;C and 25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026deg;C, respectively (Fig. S2a), and mean solar radiation was 16.7\u0026thinsp;\u0026plusmn;\u0026thinsp;7.5 MJ m\u003csup\u003e\u0026ndash;2\u003c/sup\u003e d\u003csup\u003e\u0026ndash;1\u003c/sup\u003e (Fig. S2c).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section3\"\u003e \u003ch2\u003e2.1.2. QTL-seq analysis in Japan\u003c/h2\u003e \u003cp\u003eLeaf samples of the F\u003csub\u003e5\u003c/sub\u003e RILs in 2023 grown at Iwate University were collected from each line at 67 DAT. For QTL-seq analysis, based on the phenotyping data, each 40 RILs from each lower and upper extremes were identified from the 894 RILs, and the lower extremes denoted as L-bulk, and the upper extremes denoted as U-bulk, and bulk DNA extraction for each trait was conducted. Note that QTL-seq for spikelet fertility was done for top and bottom 20. Total genomic DNA was extracted from leaf blades using NucleoSpin Plant II Kit (Macherey-Nagel). Sequencing libraries were prepared using a Collibri ES DNA Library Prep Kit for Illumina Systems (Thermo Fisher Scientific, Waltham, MA, USA). The Libraries of the L-bulk and U-bulk for each trait were sequenced on a NovaSeq X Plus (Illumina, San Diego, CA, USA) instrument in 151-bp paired-end mode.\u003c/p\u003e \u003cp\u003eQTL-seq analysis (Takagi et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) was performed using QTL-seq pipeline v2.2.8 (Sugihara et al, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) to identify genomic regions associated with the target trait. The differences in SNP indexes between the L-bulk and U-bulk were calculated using a pseudo-reference genome of \u0026lsquo;Hitomebore\u0026rsquo;, which was constructed by incorporating the SNPs of \u0026lsquo;Hitomebore\u0026rsquo; into the IRGSP-1.0 reference genome. SNPs with a read depth of \u0026ge;\u0026thinsp;10 were considered for SNP index analysis. A SNP index of 1 indicates that all short reads contain the allele from the cultivar \u0026lsquo;X-Jigna\u0026rsquo;, whereas a SNP index of 0 indicates that all short reads contain the reference allele from \u0026lsquo;Hitomebore\u0026rsquo;. A SNP index of 0.5 indicates equal contribution from both parental alleles at that position. The SNP index was plotted using a sliding window approach with a 2-Mb window and 1-kb step size for both bulks. The ΔSNP index, defined as the difference in SNP indexes between the two bulks, was calculated to detect genomic regions showing significant allele frequency differences. Statistical confidence intervals (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05 and p\u0026thinsp;\u0026lt;\u0026thinsp;0.01) were determined to identify significant peaks, which were regarded as putative QTL regions associated with the traits. The identified QTL regions were checked from the Rice Annotation Project Database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://rapdb.dna.affrc.go.jp/\u003c/span\u003e\u003cspan address=\"https://rapdb.dna.affrc.go.jp/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e)\u003c/span\u003e to confirm their novelty.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Confirmation of the detected QTLs in 2024\u003c/h2\u003e \u003cdiv id=\"Sec7\" class=\"Section3\"\u003e \u003ch2\u003e2.2.1. Phenotyping in Japan and Ethiopia\u003c/h2\u003e \u003cp\u003eCold tolerance was again determined in Iwate in 2024 (there is no comparable facility in Ethiopia). We transplanted 98 F\u003csub\u003e6\u003c/sub\u003e RILs randomly selected from the same 846 lines and both parents on 23 May. These RILs included some of the 200 RILs grown in 2023. Cool water was applied at 19.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026deg;C (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD) for 54 days from 54 to 107 DAT (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003eb); the water depth was kept at 30 cm to fully expose the developing panicles to cold at booting. Mean air temperature and solar radiation during this period were 26.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7\u0026deg;C (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003eb) and 15.8\u0026thinsp;\u0026plusmn;\u0026thinsp;6.6 MJ m\u003csup\u003e\u0026ndash;2\u003c/sup\u003e (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003eSeed shattering was confirmed in both Japan and Ethiopia in 2024. In Japan, the same 98 F\u003csub\u003e6\u003c/sub\u003e RILs as tested for cold tolerance plus the parents were transplanted on 23 May 2024 at Iwate University to confirm the QTLs detected by QTL-seq analysis. Each RIL had 3 replications in 4 rows with 7 hills per row (28 plants per replicate). On 4\u0026ndash;19 September, 10 plants per replicate were harvested, and seed shattering of 1 panicle per line or parent cultivar was assessed. Air and water temperatures during the reproductive period (30 d before heading of \u0026lsquo;X-Jigna\u0026rsquo;) were 23.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.9\u0026deg;C and 25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026deg;C, respectively (Fig. S2b), and solar radiation was 18.6\u0026thinsp;\u0026plusmn;\u0026thinsp;7.7 MJ m\u003csup\u003e\u0026ndash;2\u003c/sup\u003e d\u003csup\u003e\u0026ndash;1\u003c/sup\u003e (Fig. S2d). The other agronomic traits measured in 2023 were also measured.\u003c/p\u003e \u003cp\u003eIn Ethiopia, we transplanted 52 F\u003csub\u003e6\u003c/sub\u003e RILs randomly selected from the 98 RILs, plus the parents and two standard check cultivars, at the Fogera National Rice Research and Training Center: 15 plants each (3 rows \u0026times; 5 plants) in 2 replications with a spacing of 25 cm between plants and 20 cm between rows (Fig. S3b). The September mean air temperatures during the reproductive stage (before heading of \u0026lsquo;X-Jigna\u0026rsquo; on 2 October) were 23.1\u0026deg;C (mean average), 17.7\u0026deg;C (mean minimum), and 28.4\u0026deg;C (mean maximum) (Fig. S3a). No solar radiation data were available. Fertilizer was applied as per the local recommendations at 184 kg N ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, 46 kg P ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e, and 8.5 kg S ha\u003csup\u003e\u0026ndash;1\u003c/sup\u003e. All P and S fertilizers and 42% of the N were applied basally 1 week after transplanting, followed by 29% of the N (as urea) at tillering stage and 29% at heading stage. Weeding and water management were done as needed. The seed shattering score of 1 panicle from 1 plant in each plot was assessed. The shattering score was based on IRRI (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), which scored from 1 to 9, lower shattering to higher shattering. Seven other agronomic traits were also measured: (1) days to heading in each plot; (2) culm length and (3) panicle length of 5 plants per replicate at maturity; and (4) panicle number, (5) biomass, (6) grain yield, and (7) harvest index of 15 plants per replicate.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section3\"\u003e \u003ch2\u003e2.2.3. Marker analysis in Japan and Ethiopia\u003c/h2\u003e \u003cp\u003eLeaf samples of the 98 F\u003csub\u003e6\u003c/sub\u003e RILs grown at Iwate University, Japan, in 2024 were collected at 40 DAT of the maximum tillering stage. InDel markers were developed from the whole genome sequencing result following the method described by Natsume et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In total, 22 markers were designed to validate 13 QTLs associated with 9 traits, with one to five markers per QTL (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Briefly, InDels polymorphic between the two parental cultivars were identified, and candidates with product sizes greater than 150 bp and GC content above 40% were retained. Subsequently, InDels located within the peak regions of ΔSNP-index values in the target chromosomes were chosen. The polymorphisms of these InDels between the two parents were examined by PCR.\u003c/p\u003e \u003cp\u003eFor polymorphism testing, a 10 \u0026micro;l PCR reaction containing 2\u0026micro;l template DNA, 0.4\u0026micro;l forward and reverse primers, 2.2\u0026micro;l distilled water, and 5\u0026micro;l Quick Taq was used. PCR conditions were as follows: 94\u0026deg;C for 3 min; 35 cycles of 94\u0026deg;C for 30 sec, 55\u0026deg;C for 30 sec, and 72\u0026deg;C for 30 sec. Amplicons were separated by electrophoresis on a 3% agarose gel (KANTO HC) at 100 V for 30 min, stained with 0.5 \u0026micro;g/ml ethidium bromide for 30 min, and visualized under ultraviolet light.\u003c/p\u003e \u003cp\u003eTwenty-two markers showing clear polymorphism between the two parents were selected and used for genotyping the 98 RILs and their parents. Genotyping was conducted using a 5 \u0026micro;l PCR reaction (1\u0026micro;l template DNA, 0.025\u0026micro;l forward and reverse primers, 1.95\u0026micro;l distilled water, and 2\u0026micro;l Go Taq Green Master Mix) under the same thermal conditions as above. Single-marker analysis was performed to confirm the effects of the identified QTLs and to determine the most effective markers representing each QTL region.\u003c/p\u003e \u003cp\u003eWe did not genotype F\u003csub\u003e6\u003c/sub\u003e plants in Ethiopia, so we applied the Japanese genotyping data to both Japan and Ethiopia. The seeds of F\u003csub\u003e5\u003c/sub\u003e RILs were sent from Japan to Ethiopia in 2022, and the F\u003csub\u003e6\u003c/sub\u003e seeds were raised in Ethiopia in 2023.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\n \u003ch2\u003e3.1. Cold tolerance at reproductive stage\u003c/h2\u003e\n \u003cp\u003eUnder cold conditions, spikelet fertility was 90% in \u0026lsquo;Hitomebore\u0026rsquo;, 44% in \u0026lsquo;X-Jigna\u0026rsquo;, and 14% to 95% among the 200 RILs, with a mean of 69% (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ea, b). QTL-seq analysis of the top 20 lines (U-bulk) and the bottom 20 lines (L-bulk) detected 3 significant QTLs: \u003cem\u003eqSFC5\u003c/em\u003e on chr. 5 (20.0\u0026ndash;24.4 Mb) with an average ∆SNP index of \u0026minus;\u0026thinsp;0.481 (derived from \u0026lsquo;Hitomebore\u0026rsquo;), \u003cem\u003eqSFC8\u003c/em\u003e on chr. 8 (2.4\u0026ndash;7.4 Mb; \u0026minus;0.615; \u0026lsquo;Hitomebore\u0026rsquo;), and \u003cem\u003eqSFC10\u003c/em\u003e on chr. 10 (20.7\u0026ndash;21.9 Mb; 0.488; \u0026lsquo;X-Jigna\u0026rsquo;) (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table\u0026nbsp;\u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig. S4).\u003c/p\u003e\n \u003cdiv class=\"gridtable\"\u003e\u0026nbsp;\u003ctable id=\"Tab1\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eSummary of 20 identified QTLs for 9 agronomic traits of rice using 846 F\u003csub\u003e5\u003c/sub\u003e RILs developed from the cross between \u0026lsquo;X-Jigna\u0026rsquo; \u0026times; \u0026lsquo;Hitomebore\u0026rsquo; in the QTL-seq approaches in Iwate, Japan, in 2023.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003ccolgroup cols=\"10\"\u003e\u003c/colgroup\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eAgronomic trait\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eQTL\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eChr. No.\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\" colspan=\"2\"\u003e\n \u003cp\u003eRegion (Mb)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eSNP\u003c/p\u003e\n \u003cp\u003eindex 1\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eSNP\u003c/p\u003e\n \u003cp\u003eindex 2\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003e\u0026Delta;SNP\u003c/p\u003e\n \u003cp\u003eindex\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eContributing parent\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eStart\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eEnd\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003eSpikelet fertility\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC5\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e20.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e24.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.625\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.144\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.481\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e7.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.683\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.068\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.615\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC10\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e20.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e21.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.217\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.706\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.488\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSeed shattering\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqSHT1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e30.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e41.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.220\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.840\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.621\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eDays to heading\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqDH8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e9.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.837\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.175\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.662\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"5\"\u003e\n \u003cp\u003eCulm length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e21.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e32.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.221\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.832\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.611\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL3.1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e18.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e21.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.249\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.865\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.616\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL3.2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e27.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e35.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.350\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.872\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.522\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL8.1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.746\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.206\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.540\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL8.2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e23.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e25.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.144\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.771\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.627\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003ePanicle length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPL3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e33.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e34.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.306\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.834\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.529\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPL8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e7.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.817\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.196\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.621\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPL12\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e17.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e18.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.800\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.313\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.488\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003ePanicles per plant\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPN2.1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.711\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.284\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.428\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPN2.2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e16.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e17.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.737\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.309\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.427\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eBiomass per plant\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPL8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.788\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.538\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPL9\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.748\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.233\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.516\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eBiomass per panicle\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPA3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e34.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e35.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.240\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.760\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.520\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPA8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u003cem\u003e8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.750\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.220\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026minus;0.530\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHitomebore\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHarvest index\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqHI8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.270\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.797\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.527\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eX-Jigna\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003ctfoot\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"10\"\u003eSNP index 1 and 2 refer to the lower and upper bulks; \u0026Delta;SNP index gives the difference between the two bulks.\u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tfoot\u003e\n \u003c/table\u003e\n \u003c/div\u003e\n \u003cp\u003eIn the confirmation study in 2024 (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e), at \u003cem\u003eqSFC5\u003c/em\u003e, all 3 markers (M14-5-20554705, M24-5-20631657, and M25-5-20673667, 20.5\u0026ndash;20.7 Mb) revealed significant phenotypic differences (\u003cem\u003eP\u0026thinsp;\u0026lt;\u003c/em\u003e\u0026thinsp;0.01), with percentage of phenotypic variation explained (PVE)\u0026thinsp;=\u0026thinsp;11%\u0026ndash;14% between parental types. At \u003cem\u003eqSFC8\u003c/em\u003e, all 4 markers (M15-8-3642818, M16-8-4309379, M7-8-5179432, and M8-8-5378389, 3.6\u0026ndash;5.4 Mb) revealed significant phenotypic differences (\u003cem\u003eP\u0026thinsp;\u0026lt;\u003c/em\u003e\u0026thinsp;0.01), with PVE\u0026thinsp;=\u0026thinsp;11%\u0026ndash;13%. At \u003cem\u003eqSFC10\u003c/em\u003e, none of the 4 markers (21.0\u0026ndash;22.6 Mb) revealed a significant difference, although the \u0026lsquo;X-Jigna\u0026rsquo; allele indicated positive effects (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed) in agreement with the QTL-seq results (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). No spikelet sterility induced by low temperature was observed in Ethiopia, where the mean temperature during the reproductive stage was 23.1\u0026deg;C (Fig. S3a), higher than the threshold of 20\u0026deg;C (Shimono et al., \u003cspan class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e\n \u003cdiv class=\"gridtable\"\u003e\u0026nbsp;\u003ctable id=\"Tab2\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eConfirmation of QTLs for nine agronomic traits of rice identified using the QTL-seq approach through indel marker genotyping using F\u003csub\u003e6\u003c/sub\u003e RILs at Iwate, Japan (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;98), and Fogera, Ethiopia (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;52) in 2024.\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003ccolgroup cols=\"16\"\u003e\u003c/colgroup\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eAgronomic trait\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eQTL\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eMarker\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" colspan=\"6\"\u003e\n \u003cp\u003eIwate, Japan\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\" colspan=\"6\"\u003e\n \u003cp\u003eFogera, Ethiopia\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003cth align=\"left\" colspan=\"2\"\u003e\n \u003cp\u003eHitomebore type\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" colspan=\"2\"\u003e\n \u003cp\u003eX-Jigna type\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePVE (%)\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\" colspan=\"2\"\u003e\n \u003cp\u003eHitomebore type\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\" colspan=\"2\"\u003e\n \u003cp\u003eX-Jigna type\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\u0026nbsp;\u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePVE (%)\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"11\"\u003e\n \u003cp\u003eSpikelet fertility\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC5\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM14-5-20554705\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e83.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e76.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;2.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e10.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM24-5-20631657\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e83.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e75.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e14.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM25-5-20673667\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e83.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e75.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e11.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e84.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e76.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e84.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e76.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7-8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e83.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e75.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM8-8-5378389\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e83.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e76.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;2.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e10.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqSFC10\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM13-10-21053134\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e78.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e82.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM26-10-2114085\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e80.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;2.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e80.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM10-10-22599813\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e78.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e81.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM11-10-22600962\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e78.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e81.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u0026ndash;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"5\"\u003e\n \u003cp\u003eSeed shattering\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"5\"\u003e\n \u003cp\u003e\u003cem\u003eqSHT1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM1-1-29019501\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e40.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;4.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e48.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;5.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM2-1-32576298\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e36.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e52.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM22-1-34731315\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e27.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e63.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e30.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e19.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM23-1-34895063\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e28.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e64.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e29.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM12-1-35259855\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e28.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e67.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e34.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e18.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003eDays to heading\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqDH8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e104.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e98.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.60\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e36.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e103.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e102.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e105.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e97.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e60.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e103.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e102.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7-8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e104.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e97.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e40.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e103.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e102.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM8-8-5378389\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e103.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e98.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.74\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e26.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e103.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e102.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"8\"\u003e\n \u003cp\u003eCulm length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqCL2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM4-2-21821646\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e96.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e99.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM21-2-23976867\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e95.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e100.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.95\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e87.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM20-2-26163645\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e95.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e101.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e17.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e87.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM6-2-30732909\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e96.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e100.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e6.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e87.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003e\u003cem\u003eqCL8.1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e100.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e95.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e10.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e101.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e93.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e20.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e84.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7-8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e100.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e94.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e85.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqCL8.2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM32-8-23461108\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e98.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e97.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e86.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.003\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003ePanicle length\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqPL8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e23.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e22.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e21.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e23.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e22.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e35.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7-8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e23.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e22.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e19.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM8-8-5378389\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e23.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e22.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e9.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePanicle number\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e\u003cem\u003eqPN2.1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM17-2-8236008\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e15.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e15.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.30\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003eBiomass per plant\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"4\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPL8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e68.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e63.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.97\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e16.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e64.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e68.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e69.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e62.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.96\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e26.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e65.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e66.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7-8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e68.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e63.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e12.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e66.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e67.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM8-8-5378389\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e67.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e63.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e**\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e10.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e66.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e67.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;4.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003eBiomass per panicle\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"3\"\u003e\n \u003cp\u003e\u003cem\u003eqBMPA8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM7- 8-5179432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e4.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e2.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003eHarvest index\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\" rowspan=\"2\"\u003e\n \u003cp\u003e\u003cem\u003eqHI8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM15-8-3642818\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.43\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eM16-8-4309379\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e5.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\u0026nbsp;\u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"char\"\u003e\n \u003cp\u003e\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ens\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003ctfoot\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"16\"\u003eValues are means\u0026thinsp;\u0026plusmn;\u0026thinsp;SEM; PVE (%)\u0026thinsp;=\u0026thinsp;percentage of phenotypic variance explained as expressed by \u003cem\u003eR\u003c/em\u003e\u003csup\u003e2\u003c/sup\u003e; a total of 22 InDel marks developed in the peak regions of the QTL-seq result were used for confirmation of the QTL-seq result.\u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tfoot\u003e\n \u003c/table\u003e\n \u003c/div\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\n \u003ch2\u003e3.2. Seed shattering\u003c/h2\u003e\n \u003cp\u003eThe rate of seed shattering of the 846 RILs in 2023 ranged from 4% to 100%, with a mean of 49% and a bimodal distribution, reflecting the rates of 19% in \u0026lsquo;Hitomebore\u0026rsquo; and 99% in \u0026lsquo;X-Jigna\u0026rsquo; (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ea, b). QTL-seq analysis of the U-bulk and L-bulk detected a significant peak on chr. 1, at 30.8\u0026ndash;41.5 Mb, indicating a significant major QTL, \u003cem\u003eqSHT1\u003c/em\u003e, with an average ∆SNP index of 0.621 (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e; Fig. S5). This QTL was contributed by \u0026lsquo;X-Jigna\u0026rsquo;.\u003c/p\u003e\n \u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n \u003cp\u003eIn the confirmation study in 2024, in Japan, 4 out of 5 markers (M2-1-32576298, M22-1-34731315, M23-1-34895063, and M12-1-35259855, 34.7\u0026ndash;35.2 Mb) revealed significant phenotypic differences (\u003cem\u003eP\u0026thinsp;\u0026lt;\u003c/em\u003e\u0026thinsp;0.01), with PVE\u0026thinsp;=\u0026thinsp;7%\u0026ndash;34% (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ed; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). The other marker (M1-1-29019501, 29 Mb) indicated a positive effect of the \u0026lsquo;X-Jigna\u0026rsquo; allele (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ed) in agreement with the QTL-seq result (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). In Ethiopia, 3 out of the 5 markers revealed significant effects (\u003cem\u003eP\u0026thinsp;\u0026lt;\u003c/em\u003e\u0026thinsp;0.01), with PVE\u0026thinsp;=\u0026thinsp;19%\u0026ndash;20% (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ee; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). The other 2 markers again indicated a positive effect (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ee) in agreement with the QTL-seq result (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). The three markers at 34\u0026ndash;35 Mb on chr. 1 confirmed in both Japan and Ethiopia would be suitable for breeding.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\n \u003ch2\u003e3.3. Other traits\u003c/h2\u003e\n \u003cp\u003eQTL-seq analysis for days to heading, culm length, biomass, grain yield, and harvest index (HI) found another 16 QTLs, bringing the total to 20 (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). No significant QTL for grain yield was detected (Fig. S6). Among the QTLs for cold tolerance and seed shattering, only \u003cem\u003eqSFC8\u003c/em\u003e, for cold tolerance, coincided with QTLs for other traits (Fig. S7; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e), namely days to heading (\u003cem\u003eqDH8\u003c/em\u003e), culm length (\u003cem\u003eqCL8.1\u003c/em\u003e ), panicle length (\u003cem\u003eqPL8\u003c/em\u003e), and biomass (\u003cem\u003eqBMPL8\u003c/em\u003e, \u003cem\u003eqBMPA8\u003c/em\u003e). Whole genome QTL-seq results of the traits are shown in Figures S8\u0026ndash;S14.\u003c/p\u003e\n \u003cp\u003eIn the confirmation study, 11 of the 12 QTLs found in Japan in 98 RILs (including those for spikelet fertility (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed) and shattering (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ed, e) but excluding \u003cem\u003eqPN2.1\u003c/em\u003e(Fig. S15b, d)) were confirmed by more than 1 marker each (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e, Figs. S16b, d, S17b, d, S18b, d, S19b, d). But only 1 of the 9 QTLs found in Ethiopia in 52 RILs for seed shattering (\u003cem\u003eqSHT1\u003c/em\u003e) was confirmed (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ee), although another 5 QTLs indicated agreement with QTL-seq results for days to heading (\u003cem\u003eqDH8\u003c/em\u003e, Fig. S16f), culm length (\u003cem\u003eqCL2\u003c/em\u003e, \u003cem\u003eqCL8.1\u003c/em\u003e, \u003cem\u003eqCL8.2\u003c/em\u003e, Fig. S17f), and panicle length (\u003cem\u003eqPL2\u003c/em\u003e, Fig. S18f). On the other hand, allele effects of another 4 QTLs indicated opposite effects on biomass per plant (\u003cem\u003eqBMPL8\u003c/em\u003e, Fig. S19f) and per panicle (\u003cem\u003eqBMPA8\u003c/em\u003e, Fig. S20f) and HI (\u003cem\u003eqHI8\u003c/em\u003e, Fig. S21f) compared to QTL-seq and the confirmation in Japan; i.e., the \u003cem\u003eqBMPL8\u003c/em\u003e and \u003cem\u003eqBMPA8\u003c/em\u003e alleles of \u0026lsquo;Hitomebore\u0026rsquo; increased the biomass in Japan (Figs. S19b, d, S20b, d), in agreement with the QTL-seq results (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e), but decreased it in Ethiopia (Figs. S19f, S20f). Note that no significant QTL was detected (Fig. S6) and histogram of grain yield in Japan and Ethiopia was shown in Fig. S22.\u003c/p\u003e\n \u003cp\u003eOur QTL-seq analysis identified QTLs for spikelet fertility under cold temperatures during the reproductive stage (\u003cem\u003eqSFC5\u003c/em\u003e, \u003cem\u003eqSFC8\u003c/em\u003e, and \u003cem\u003eqSFC10\u003c/em\u003e) and for seed shattering (\u003cem\u003eqSHT1\u003c/em\u003e) in the Ethiopian rice cultivar \u0026lsquo;X-Jigna\u0026rsquo; (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). To our knowledge, this is the first QTL mapping study of an Ethiopian rice cultivar grown in the field in different environments. For breeding by marker selection in Ethiopia, markers for both traits are critical.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\n \u003ch2\u003e4.1. Cold tolerance at reproductive stage\u003c/h2\u003e\n \u003cp\u003eCold tolerance at the reproductive stage is an important trait in Ethiopia\u0026rsquo;s high-elevation, cool-climate zones. We found three QTLs for cold tolerance: the \u0026lsquo;Hitomebore\u0026rsquo; \u003cem\u003eqSFC5\u003c/em\u003e and \u003cem\u003eqSFC8\u003c/em\u003e alleles and the \u0026lsquo;X-Jigna\u0026rsquo; \u003cem\u003eqSFC10\u003c/em\u003e allele increase it (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). \u003cem\u003eqSFC5\u003c/em\u003e was detected at 20\u0026ndash;24 Mb on chr. 5 and was confirmed by markers at 20.5\u0026ndash;20.6 Mb (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). This region is similar to regions reported by Xu et al. (\u003cspan class=\"CitationRef\"\u003e2008\u003c/span\u003e) (517 F\u003csub\u003e2\u003c/sub\u003e and 1517 F\u003csub\u003e3\u003c/sub\u003e RILs between tolerant \u0026lsquo;Kunmingxiaobaigu\u0026rsquo; and sensitive \u0026lsquo;Towada\u0026rsquo;), Andaya and Mackill (\u003cspan class=\"CitationRef\"\u003e2003\u003c/span\u003e) (191 F\u003csub\u003e6\u003c/sub\u003e RILs between tolerant \u0026lsquo;M-202\u0026rsquo; and sensitive \u0026lsquo;IR50\u0026rsquo;), and Shimono et al. (\u003cspan class=\"CitationRef\"\u003e2016\u003c/span\u003e) (208 F\u003csub\u003e7\u003c/sub\u003e and F\u003csub\u003e8\u003c/sub\u003e RILs between tolerant \u0026lsquo;Tohoku-PL3\u0026rsquo; and sensitive \u0026lsquo;Akihikari\u0026rsquo;). Combined with the QTL and the transcriptome, Shimono et al. (\u003cspan class=\"CitationRef\"\u003e2016\u003c/span\u003e) associated the region with candidate genes for \u003cem\u003eO\u003c/em\u003e-methyltransferase ZRP4 (Os05g0515600, chr. 05:25569090..25570887) and \u0026beta;-1,3-glucanase-like protein (Os05g0535100, chr. 05:26570425..26573958) in a cross between \u0026lsquo;Tohoku-PL3\u0026rsquo; and \u0026lsquo;Akihikari\u0026rsquo;.\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eqSFC8\u003c/em\u003e was detected at 2.4\u0026ndash;7.4 Mb on chr. 8 (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e) and was confirmed by markers at 3.6\u0026ndash;5.4 Mb (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). This region is similar to regions reported by Kuroki et al. (\u003cspan class=\"CitationRef\"\u003e2007\u003c/span\u003e) (F\u003csub\u003e3\u003c/sub\u003e and F\u003csub\u003e7\u003c/sub\u003e backcross inbred lines between tolerant \u0026lsquo;Hokkai PL\u0026rsquo; [donor to \u0026lsquo;Hayahuki\u0026rsquo; and \u0026lsquo;Padi Labou Alumbis\u0026rsquo;] and sensitive \u0026lsquo;Hokkai 287\u0026rsquo;) and Wainaina et al. (\u003cspan class=\"CitationRef\"\u003e2018\u003c/span\u003e) (108 F\u003csub\u003e2\u003c/sub\u003e between tolerant \u0026lsquo;Hananomai\u0026rsquo; and sensitive \u0026lsquo;WAB56-10\u0026rsquo;). \u003cem\u003eqSFC8\u003c/em\u003e overlapped several QTLs for heading date, culm length, and biomass (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e; Fig. S7). The \u003cem\u003eqSFC8\u003c/em\u003e region, which is large at ~\u0026thinsp;5 Mb, would hold multiple genes, and/or some of which might have pleiotropic effects (Nutan et al., \u003cspan class=\"CitationRef\"\u003e2020\u003c/span\u003e). Map-based cloning and near-isogenic lines would be able to narrow down the position of the responsible gene.\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eqSFC10\u003c/em\u003e was detected at 20.7\u0026ndash;21.9 Mb on chr. 10 (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ec; Tables \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e) in \u0026lsquo;X-Jigna\u0026rsquo;, the cold-sensitive parent, with indicative results (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003ed; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). The region is close to a region for drought tolerance reported by Moncada et al. (\u003cspan class=\"CitationRef\"\u003e2001\u003c/span\u003e) (274 BC\u003csub\u003e2\u003c/sub\u003eF\u003csub\u003e2\u003c/sub\u003e backcross inbred lines between \u0026lsquo;Caiapo\u0026rsquo; [upland \u003cem\u003eOryza sativa\u003c/em\u003e] and an accession of \u003cem\u003eOryza rufipogon\u003c/em\u003e). The \u0026lsquo;X-Jigna\u0026rsquo; region might prove useful in boosting cold tolerance.\u003c/p\u003e\n \u003cp\u003eThe method used to test for cold tolerance can control panicle temperature after initiation at the shoot base but not after emergence above the water level (Shimono et al., \u003cspan class=\"CitationRef\"\u003e2005\u003c/span\u003e), when it is exposed to atmospheric temperature. During 2023, the air temperature during the reproductive period was \u0026gt;\u0026thinsp;26\u0026deg;C (Fig. \u003cspan class=\"InternalRef\"\u003eS1\u003c/span\u003ea), higher than the lower threshold temperature of 20\u0026deg;C for inducing sterility (Shimono et al., \u003cspan class=\"CitationRef\"\u003e2005\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e2007\u003c/span\u003e). The difference in culm length and days to heading among RILs would not indirectly affect cold tolerance through the panicle temperature affected by atmospheric temperature.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\n \u003ch2\u003e4.2. Seed shattering\u003c/h2\u003e\n \u003cp\u003eSeed shattering is important to yield loss before harvest (Wang et al., \u003cspan class=\"CitationRef\"\u003e2024\u003c/span\u003e). We found a significant QTL on chr. 1, \u003cem\u003eqSHT1\u003c/em\u003e, at 30.8\u0026ndash;41.5 Mb (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ec; Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e), and confirmed it at 34\u0026ndash;35 Mb in both Japan and Ethiopia (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ed, e; Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). \u003cem\u003eqSH1\u003c/em\u003e encodes a BEL1-type homeobox gene (Konishi et al., \u003cspan class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wu et al., \u003cspan class=\"CitationRef\"\u003e2023\u003c/span\u003e), which causes seed shattering in rice (\u003cem\u003eOs01g0848400\u003c/em\u003e, chr. 01:36 445 456..36 449 951).\u003c/p\u003e\n \u003cp\u003eOur RILs showed a wide range of seed shattering habits (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003eb ), attributable to a haplotype mutation in this QTL region (Konishi et al., \u003cspan class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wu et al., \u003cspan class=\"CitationRef\"\u003e2023\u003c/span\u003e). In Ethiopia, postharvest management practices are manual (Tadesse et al., \u003cspan class=\"CitationRef\"\u003e2019\u003c/span\u003e), so farmers rely on the easy-shattering characteristics, which cause significant yield losses. Future breeding in Ethiopia should seek to balance reducing yield losses with efficiency of threshing.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e\n \u003ch2\u003e4.3. Future strategy for improving Ethiopian rice cultivars\u003c/h2\u003e\n \u003cp\u003eTo improve the production of \u0026lsquo;X-Jigna\u0026rsquo; in Ethiopia, phenotyping and genotyping studies should be conducted on-site. However, technical and management constraints made it necessary to do this in Japan and then confirm the detected QTLs in Ethiopia. This approach worked well for identifying QTLs for seed shattering (\u003cem\u003eqSHT1\u003c/em\u003e; Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003ec, d, e), days to heading (\u003cem\u003eqDH8\u003c/em\u003e; Fig. S16f), culm length (\u003cem\u003eqCL2\u003c/em\u003e, \u003cem\u003eqCL8.1\u003c/em\u003e, \u003cem\u003eqCL8.2\u003c/em\u003e; Fig. S17f), and panicle length (\u003cem\u003eqPL8\u003c/em\u003e; Fig. S18f). DNA markers for these traits would be useful to develop new cultivars by marker-assisted selection in Ethiopia. However, the important quantitative traits of biomass production (\u003cem\u003eqBMPL8\u003c/em\u003e, \u003cem\u003eqBMPA8\u003c/em\u003e; Figs. S19b, d, f, S20b, d, f) and HI (\u003cem\u003eqHI8\u003c/em\u003e; Fig. S21f) were confirmed in Japan but not in Ethiopia, where the QTLs indicated the opposite effect. This suggests that the strategy is useful for traits controlled by a few dominant genes, as for seed shattering and culm length, but not for complex quantitative traits of biomass and yield, which must be phenotyped on-site in Ethiopia. However, it should be noted that developing countries face limitations on access to phenotyping facilities, such as the cold screening facility used in this study. In current condition, the present strategy of the multi-country studies can compensate for each others\u0026rsquo; deficiencies.\u003c/p\u003e\n\u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eOur QTL-seq analysis identified QTLs for spikelet fertility under cold temperatures during the reproductive stage (\u003cem\u003eqSFC5\u003c/em\u003e, \u003cem\u003eqSFC8\u003c/em\u003e, and \u003cem\u003eqSFC10\u003c/em\u003e) and for seed shattering (\u003cem\u003eqSHT1\u003c/em\u003e)\u0026nbsp;in the Ethiopian rice cultivar ‘X-Jigna’ (Table 1). To our knowledge, this is the first QTL mapping study\u0026nbsp;of an\u0026nbsp;Ethiopian rice cultivar grown in the field in different environments. For breeding by marker selection in Ethiopia, markers for both traits are critical.\u003c/p\u003e\n\u003ch2\u003e4.1. Cold tolerance at reproductive stage\u003c/h2\u003e\n\u003cp\u003eCold tolerance at the reproductive stage is an important trait in Ethiopia’s high-elevation, cool-climate zones. We found three QTLs for cold tolerance: the ‘Hitomebore’ \u003cem\u003eqSFC5\u003c/em\u003e and \u003cem\u003eqSFC8\u003c/em\u003e alleles and the ‘X-Jigna’ \u003cem\u003eqSFC10\u003c/em\u003e allele increase it (Fig. 1c;\u0026nbsp;Table 1). \u003cem\u003eqSFC5\u003c/em\u003e was detected at 20–24 Mb on chr. 5 and was confirmed by markers at 20.5–20.6 Mb (Fig. 1d;\u0026nbsp;Table 2). This region is similar to regions reported by Xu et al. (2008) (517 F\u003csub\u003e2\u003c/sub\u003e and 1517 F\u003csub\u003e3\u003c/sub\u003e RILs between tolerant ‘Kunmingxiaobaigu’ and sensitive ‘Towada’), Andaya and Mackill (2003) (191 F\u003csub\u003e6\u003c/sub\u003e RILs between tolerant ‘M-202’ and sensitive ‘IR50’), and Shimono et al. (2016) (208 F\u003csub\u003e7\u003c/sub\u003e and F\u003csub\u003e8\u003c/sub\u003e RILs between tolerant ‘Tohoku-PL3’ and sensitive ‘Akihikari’). Combined with the QTL and the transcriptome, Shimono et al. (2016) associated the region with candidate genes for \u003cem\u003eO\u003c/em\u003e-methyltransferase ZRP4 (Os05g0515600, chr.\u0026nbsp;05:25569090..25570887) and β-1,3-glucanase-like protein (Os05g0535100, chr.\u0026nbsp;05:26570425..26573958) in a cross between ‘Tohoku-PL3’ and ‘Akihikari’.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eqSFC8\u0026nbsp;\u003c/em\u003ewas detected at 2.4–7.4 Mb on chr. 8 (Fig. 1c; Table 1)\u0026nbsp;and\u0026nbsp;was confirmed by markers at 3.6–5.4 Mb (Fig. 1d;\u0026nbsp;Table 2). This region is similar to regions reported by Kuroki et al. (2007) (F\u003csub\u003e3\u003c/sub\u003e and F\u003csub\u003e7\u003c/sub\u003e backcross inbred lines between tolerant ‘Hokkai PL’ [donor to ‘Hayahuki’ and ‘Padi Labou Alumbis’] and sensitive ‘Hokkai 287’) and Wainaina et al. (2018) (108 F\u003csub\u003e2\u003c/sub\u003e between tolerant ‘Hananomai’ and sensitive ‘WAB56-10’). \u003cem\u003eqSFC8\u003c/em\u003e overlapped several QTLs for heading date, culm length, and biomass (Table 2; Fig. S7). The \u003cem\u003eqSFC8\u0026nbsp;\u003c/em\u003eregion, which is large at ~5 Mb, would hold multiple\u0026nbsp;genes,\u0026nbsp;and/or some of which might have pleiotropic effects (Nutan et al., 2020). Map-based cloning and near-isogenic lines would be able to narrow down the position of the responsible gene.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eqSFC10\u0026nbsp;\u003c/em\u003ewas detected at 20.7–21.9 Mb on chr. 10 (Fig. 1c;\u0026nbsp;Tables 1) in ‘X-Jigna’, the cold-sensitive parent, with indicative results (Fig. 1d; Table 2). The region is close to a region for drought tolerance reported by Moncada et al. (2001) (274 BC\u003csub\u003e2\u003c/sub\u003eF\u003csub\u003e2\u003c/sub\u003e backcross inbred lines between ‘Caiapo’ [upland \u003cem\u003eOryza sativa\u003c/em\u003e] and an accession of \u003cem\u003eOryza rufipogon\u003c/em\u003e). The ‘X-Jigna’ region might prove useful in boosting cold tolerance.\u003c/p\u003e\n\u003cp\u003eThe method used to test for cold tolerance can control panicle temperature after initiation at the shoot base but not after emergence above the water level (Shimono et al., 2005), when it is exposed to atmospheric temperature. During 2023, the air temperature during the reproductive period was \u0026gt;26 °C (Fig. S1a), higher than the lower threshold temperature of 20 °C for inducing sterility (Shimono et al., 2005, 2007). The difference in culm length and days to heading among RILs\u0026nbsp;would not indirectly affect cold tolerance through the panicle temperature\u0026nbsp;affected by atmospheric temperature.\u003c/p\u003e\n\u003ch2\u003e4.2. Seed shattering\u003c/h2\u003e\n\u003cp\u003eSeed shattering is important to yield loss before harvest (Wang et al., 2024). We found a significant QTL on chr. 1, \u003cem\u003eqSHT\u003c/em\u003e\u003cem\u003e1\u003c/em\u003e,\u0026nbsp;at 30.8–41.5 Mb\u0026nbsp;(Fig. 2c; Table 1), and confirmed it at 34–35 Mb in both Japan and Ethiopia (Fig. 2d, e; Table 2).\u0026nbsp;\u003cem\u003eqSH1\u003c/em\u003e encodes a BEL1-type homeobox gene (Konishi et al., 2006; Wu et al., 2023), which causes seed shattering in rice (\u003cem\u003eOs01g0848400\u003c/em\u003e, chr.\u0026nbsp;01:36 445 456..36 449 951).\u003c/p\u003e\n\u003cp\u003eOur RILs showed a wide range of seed shattering habits (Fig. 2b\u0026nbsp;),\u0026nbsp;attributable to a haplotype mutation\u0026nbsp;in this QTL region (Konishi et al., 2006; Wu et al., 2023). In Ethiopia, postharvest management practices are manual (Tadesse et al., 2019), so farmers rely on the easy-shattering characteristics, which\u0026nbsp;cause significant yield losses. Future breeding in Ethiopia should seek to balance reducing yield losses with efficiency of threshing.\u003c/p\u003e\n\u003ch2\u003e4.3. Future strategy for improving Ethiopian rice cultivars\u003c/h2\u003e\n\u003cp\u003eTo improve the production of ‘X-Jigna’ in Ethiopia, phenotyping and genotyping studies should be conducted on-site. However, technical and management constraints made it necessary to do this in Japan and then confirm the detected QTLs in Ethiopia. This approach worked well for identifying QTLs for seed shattering (\u003cem\u003eqSHT1\u003c/em\u003e; Fig. 2c, d, e), days to heading\u0026nbsp;(\u003cem\u003eqDH8\u003c/em\u003e; Fig. S16f), culm length\u0026nbsp;(\u003cem\u003eqCL2\u003c/em\u003e, \u003cem\u003eqCL8.1\u003c/em\u003e, \u003cem\u003eqCL8.2\u003c/em\u003e; Fig. S17f), and panicle length\u0026nbsp;(\u003cem\u003eqPL8\u003c/em\u003e; Fig. S18f). DNA markers for these traits would be useful to develop new cultivars by marker-assisted selection in Ethiopia.\u0026nbsp;However, the important quantitative traits of biomass production (\u003cem\u003eqBMPL8\u003c/em\u003e, \u003cem\u003eqBMPA8\u003c/em\u003e;\u0026nbsp;Figs. S19b, d,\u0026nbsp;f, S20b, d,\u0026nbsp;f) and HI\u0026nbsp;(\u003cem\u003eqHI8\u003c/em\u003e; Fig. S21f) were confirmed in Japan but not in Ethiopia, where the QTLs indicated the opposite effect. This suggests that the strategy is useful for traits controlled by a few dominant genes, as for seed shattering and culm length, but not for complex quantitative traits of biomass and yield, which must be phenotyped on-site in Ethiopia. However, it should be noted that developing countries face limitations on access to phenotyping facilities, such as the cold screening facility used in this study. \u0026nbsp;In current condition, the present strategy of the multi-country studies can compensate for each others’ deficiencies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eCRediT statement: Zelalem Zewdu: Writing \u0026ndash; original draft preparation, data curation; Mulugeta Atnaf: Methodology, writing \u0026ndash; review \u0026amp; editing; Assaye Berie: data curation; Honoka Takasago: data curation; Reo Sato: data curation; Ryuji Ishikawa: writing \u0026ndash; review \u0026amp; editing; Akira Abe: data curation, methodology, writing \u0026ndash; review \u0026amp; editing; Maya Matsunam: writing \u0026ndash; review \u0026amp; editing; Hiroyuki Shimono: conceptualization, supervision, methodology, writing \u0026ndash; review \u0026amp; editing\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eZelalem Zewdu is grateful for the financial support of the Japan International Cooperation Agency (JICA) through its EthioRice-2 project. The authors thank the staff of the national rice research program at the Fogera National Rice Research and Training Center for managing the experiment; of the Iwate University Crop Science laboratory for their support in data collection at Iwate; and of the Iwate Biotechnology Research Center laboratory for assisting with the QTL-seq analysis and genotyping. We thank the Iwate Prefectural Research Center for providing the cold screening facilities.\u003c/p\u003e \u003cp\u003e \u003cb\u003eCRediT statement: Zelalem Zewdu\u003c/b\u003e: Writing \u0026ndash; original draft preparation, data curation; \u003cb\u003eMulugeta Atnaf\u003c/b\u003e: Methodology, writing \u0026ndash; review \u0026amp; editing; \u003cb\u003eAssaye Berie\u003c/b\u003e: data curation; \u003cb\u003eHonoka Takasago\u003c/b\u003e: data curation; \u003cb\u003eReo Sato\u003c/b\u003e: data curation; \u003cb\u003eRyuji Ishikawa\u003c/b\u003e: writing \u0026ndash; review \u0026amp; editing; \u003cb\u003eAkira Abe\u003c/b\u003e: data curation, methodology, writing \u0026ndash; review \u0026amp; editing; \u003cb\u003eMaya Matsunam\u003c/b\u003e: writing \u0026ndash; review \u0026amp; editing; \u003cb\u003eHiroyuki Shimono\u003c/b\u003e: conceptualization, supervision, methodology, writing \u0026ndash; review \u0026amp; editing\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlemayehu, H.A., Dumbuya, G., Hasan, M., Tadesse, T., Nakajyo, S., Fujioka, T., Abe, A., Matsunami, M., Shimono, H., 2021. Genotypic variation in cold tolerance of 18 Ethiopian rice cultivars in relation to their reproductive morphology. Field Crops Res. 262, 108042.\u003c/li\u003e\n\u003cli\u003eAndaya, V.C., Mackill, D.J., 2003. QTLs conferring cold tolerance at the booting stage of rice using recombinant inbred lines from a \u003cem\u003ejaponica\u003c/em\u003e \u0026times; \u003cem\u003eindica\u003c/em\u003e cross. Theor. Appl. Genet. 106, 1084\u0026ndash;1090.\u003c/li\u003e\n\u003cli\u003eBeyene, A.M., Gashu, A.T., Tegegne, M.A., Anteneh Mihertie, A., 2022. Is the longstanding local rice cultivar \u0026lsquo;X-Jigna\u0026rsquo; being replaced by the improved variety \u0026lsquo;Shaga\u0026rsquo; in Fogera Plain, northwest Ethiopia? Cogent Econ. Fin. 10, 2145748.\u003c/li\u003e\n\u003cli\u003eDessie, A., 2020. Rice breeding achievements, potential and challenges in Ethiopia. International Journal of Research Studies in Agricultural Sciences 6, 35\u0026ndash;42.\u003c/li\u003e\n\u003cli\u003eFAO, 2024. FAOSTAT (http://faostat.fao.org/site/291/default.aspx). Food and Agriculture Organization of the United Nations.\u003c/li\u003e\n\u003cli\u003eFarrell, T.C., Fox, K.M., Williams, R.L., Fukai, S., 2006. Genotypic variation for cold tolerance during reproductive development in rice: Screening with cold air and cold water. Field Crops Res. 98, 178\u0026ndash;194.\u003c/li\u003e\n\u003cli\u003eIRRI, 2013. Standard evaluation systems for rice, 5th ed. International Rice Research Institute, Manila, Philippines.\u003c/li\u003e\n\u003cli\u003eIto, N., Hayase, H., Satake, T., Nishiyama, I., 1970. Male sterility caused by cooling treatment at the meiotic stage in rice plants. III. Male abnormalities at anthesis. Proc. Crop Sci. Soc. Jpn. 39, 60\u0026ndash;64.\u003c/li\u003e\n\u003cli\u003eJeong, J.-M., Mo, Y., Hyun, U.-J., Jeung, J.-U., 2020. Identification of quantitative trait loci for spikelet fertility at the booting stage in rice (\u003cem\u003eOryza sativa\u003c/em\u003e L.) under different low-temperature conditions. Agronomy 10.\u003c/li\u003e\n\u003cli\u003eKonishi, S., Izawa, T., Lin, S.Y., Ebana, K., Fukuta, Y., Sasaki, T., Yano, M., 2006. An SNP caused loss of seed shattering during rice domestication. Science 312, 1392\u0026ndash;1396.\u003c/li\u003e\n\u003cli\u003eKuroki, M., Saito, K., Matsuba, S., Yokogami, N., Shimizu, H., Ando, I., Sato, Y., 2007. A quantitative trait locus for cold tolerance at the booting stage on rice chromosome 8. Theor. Appl. Genet. 115, 593\u0026ndash;600.\u003c/li\u003e\n\u003cli\u003eLi, C., Zhou, A., Sang, T., 2006. Rice domestication by reducing shattering. Science 311, 1936\u0026ndash;1939.\u003c/li\u003e\n\u003cli\u003eLi, J., Zeng, Y., Pan, Y., Zhou, L., Zhang, Z., Guo, H., Lou, Q., Shui, G., Huang, H., Tian, H., et al., 2021. Stepwise selection of natural variations at CTB2 and CTB4a improves cold adaptation during domestication of \u003cem\u003ejaponica\u003c/em\u003e rice. New Phytologist 231, 1056\u0026ndash;1072.\u003c/li\u003e\n\u003cli\u003eMatsunaga, K., 2005. Establishment of an evaluation method for cold tolerance at the booting stage of rice using deep water irrigation system and development of highly cold tolerant rice varieties by combining cold tolerance genes. Bull. Miyagi Furukawa Agric. Exp. Sta. 4, 1\u0026ndash;78 (Japanese with English abstract).\u003c/li\u003e\n\u003cli\u003eMinistry of Agriculture and Rural Development, 2010. National rice research and development strategy of Ethiopia., Ethiopia, p. 61.\u003c/li\u003e\n\u003cli\u003eMoncada, P., Martinez, C.P., Borrero, J., Chatel, M., Gauch Jr, H., Guimaraes, E., Tohme, J., McCouch, S.R., 2001. Quantitative trait loci for yield and yield components in an \u003cem\u003eOryza sativa\u003c/em\u003e \u0026times; \u003cem\u003eOryza rufipogon\u003c/em\u003e BC\u003csub\u003e2\u003c/sub\u003eF\u003csub\u003e2\u003c/sub\u003e population evaluated in an upland environment. Theor. Appl. Genet. 102, 41\u0026ndash;52.\u003c/li\u003e\n\u003cli\u003eNatsume, S., Oikawa, K., Nomura, C., Ito, K., Utsushi, H., Shimizu, M., Terauchi, R., Abe, A., 2023. V-primer: software for the efficient design of genome-wide indel and SNP markers from multi-sample variant call format (VCF) genotyping data. Breed. Sci. 73, 415\u0026ndash;420.\u003c/li\u003e\n\u003cli\u003eNutan KK, Singla-Pareek SL, Pareek A. 2020. The Saltol QTL-localized transcription factor OsGATA8 plays an important role in stress tolerance and seed development in Arabidopsis and rice. J. Exp. Bot. 71: 684\u0026ndash;698.\u003c/li\u003e\n\u003cli\u003eOliver, S.N., Dennis, E.S., Dolferus, R., 2007. ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice. Plant Cell Physiol. 48, 1319\u0026ndash;1330.\u003c/li\u003e\n\u003cli\u003eOliver, S.N., Van Dongen, J.T., Alfred, S.C., Mamun, E.A., Zhao, X., Saini, H.S., Fernandes, S.F., Blanchard, C.L., Sutton, B.G., Geigenberger, P., Dennis, E.S., Dolferus, R., 2005. Cold-induced repression of the rice anther-specific cell wall invertase gene OSINV4 is correlated with sucrose accumulation and pollen sterility. Plant Cell Env. 28, 1534\u0026ndash;1551.\u003c/li\u003e\n\u003cli\u003eSakata, T., Oda, S., Tsunaga, Y., Shomura, H., Kawagishi-Kobayashi, M., Aya, K., Saeki, K., Endo, T., Nagano, K., Kojima, M., Sakakibara, H., Watanabe, M., Matsuoka, M., Higashitani, A., 2014. Reduction of gibberellin by low temperature disrupts pollen development in rice. Plant Physiol. 164, 2011\u0026ndash;2019.\u003c/li\u003e\n\u003cli\u003eSatake, T., 1976. Determination of the most sensitive stage to sterile-type cool injury in rice plants. Res. Bull. Hokkaido Nat. Agric. Exp. Stn. 113, 1\u0026ndash;44.\u003c/li\u003e\n\u003cli\u003eSato, Y., Masuta, Y., Saito, K., Murayama, S., Ozawa, K., 2011. Enhanced chilling tolerance at the booting stage in rice by transgenic overexpression of the ascorbate peroxidase gene, OsAPXa. Plant Cell Reports 30, 399\u0026ndash;406.\u003c/li\u003e\n\u003cli\u003eShimono, H., 2018. Male sterility induced by low temperature at the booting stage in rice. Jpn. J. Crop Sci. 87, 113\u0026ndash;124 (Japanese with English abstract).\u003c/li\u003e\n\u003cli\u003eShimono, H., Hasegawa, T., Iwama, K., 2002. Response of growth and grain yield in paddy rice to cool water at different growth stages. Field Crops Res. 73, 67\u0026ndash;79.\u003c/li\u003e\n\u003cli\u003eShimono, H., Hasegawa, T., Moriyama, M., Fujimura, S., Nagata, T., 2005. Modeling spikelet sterility induced by low temperature in rice. Agron. J. 97, 1524\u0026ndash;1536.\u003c/li\u003e\n\u003cli\u003eShimono, H., Okada, M., Kanda, E., Arakawa, I., 2007. Low temperature-induced sterility in rice: Evidence for the effects of temperature before panicle initiation. Field Crops Res. 101, 221\u0026ndash;231.\u003c/li\u003e\n\u003cli\u003eShimono, H., Abe, A., Aoki, N., Koumoto, T., Sato, M., Yokoi, S., Kuroda, E., Endo, T., Saeki, K., Nagano, K., 2016. Combining mapping of physiological quantitative trait loci and transcriptome for cold tolerance for counteracting male sterility induced by low temperatures during reproductive stage in rice. Physiol. Plant. 157, 175\u0026ndash;192.\u003c/li\u003e\n\u003cli\u003eSugihara, Y., Young, L., Yaegashi, H., Natsume, S., Shea, D.J., Takagi, H., Booker, H., Innan, H., Terauchi, R., Abe, A., 2022. High-performance pipeline for MutMap and QTL-seq. . PeerJ 10, e13170 https://doi.org/13110.17717/peerj.13170.\u003c/li\u003e\n\u003cli\u003eSuzuki, K., Aoki, N., Matsumura, H., Okamura, M., Ohsugi, R., Shimono, H., 2015. Cooling water before panicle initiation increases chilling-induced male sterility and disables chilling-induced expression of genes encoding OsFKBP65 and heat shock proteins in rice spikelets. Plant Cell Env. 38, 1255\u0026ndash;1274.\u003c/li\u003e\n\u003cli\u003eTadesse, T., Atnaf, M., Alemu, D., Tadesse, T., Shiratori, K., 2019. Advances in rice research and development in Ethiopia. Ethiopian Institute of Agricultural Research.\u003c/li\u003e\n\u003cli\u003eTakagi, H., Abe, A., Yoshida, K., Kosugi, S., Natsume, S., Mitsuoka, C., Uemura, A., Utsushi, H., Tamiru, M., Takuno, S., Innan, H., Cano, L.M., Kamoun, S., Terauchi, R., 2013. QTL-seq: rapid mapping of quantitative trait loci in rice by whole genome resequencing of DNA from two bulked populations. Plant J. 74, 174\u0026ndash;183.\u003c/li\u003e\n\u003cli\u003eTakahashi T, Sato N, Matsunami M, et al., 2023: Yield performance of hybrid rice in a cool climate in Japan. Field Crops Res. 291, 1\u0026ndash;6.\u003c/li\u003e\n\u003cli\u003eWainaina, C.M., Makihara, D., Nakamura, M., Ikeda, A., Suzuki, T., Mizukami, Y., Nonoyama, T., Doi, K., Kikuta, M., Samejima, H., et al., 2018. Identification and validation of QTLs for cold tolerance at the booting stage and other agronomic traits in a rice cross of a Japanese tolerant variety, Hananomai, and a NERICA parent, WAB56\u0026ndash;104. Plant Prod. Sci. 21, 132\u0026ndash;143.\u003c/li\u003e\n\u003cli\u003eWang, Y., Wu, W., Xu, J., Wang, Y., Wu, Z., Liu, H., 2024. Expounding the effect of harvest management on rice (\u003cem\u003eOryza sativa\u003c/em\u003e L.) yield and latent loss based on the accurate measurement of grain data. Agronomy 14, 1346.\u003c/li\u003e\n\u003cli\u003eWu, H., He, Q., Wang, Q., 2023. Advances in rice seed shattering. Int. J. Mol. Sci. 24, 8889.\u003c/li\u003e\n\u003cli\u003eXu, L.M., Zhou, L., Zeng, Y.W., Wang, F.M., Zhang, H.L., Shen, S.Q., Li, Z.C., 2008. Identification and mapping of quantitative trait loci for cold tolerance at the booting stage in a \u003cem\u003ejaponica\u003c/em\u003e rice near-isogenic line. Plant Science 174, 340\u0026ndash;347.\u003c/li\u003e\n\u003cli\u003eYang, L., Lei, L., Wang, J., Zheng, H., Xin, W., Liu, H., Zou, D., 2023. qCTB7 positively regulates cold tolerance at booting stage in rice. Theor. Appl. Genet. 136, 135.\u003c/li\u003e\n\u003cli\u003eZewdu, Z., Atnaf, M., Sato, R., Ishikawa, R., Abe, A., Matsunami, M., Shimono, H., 2025. Comparison of rice yield potential in Ethiopia and in northern Japan: an experimental and modeling study. Journal of Agricultural Meteorology 81, 36\u0026ndash;43.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"euphytica","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"euph","sideBox":"Learn more about [Euphytica](https://www.springer.com/journal/10681)","snPcode":"10681","submissionUrl":"https://submission.springernature.com/new-submission/10681/3","title":"Euphytica","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Cold tolerance, Ethiopia, QTL-seq, Rice, Shattering","lastPublishedDoi":"10.21203/rs.3.rs-8244416/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8244416/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eGenetic improvement is essential for improved rice production in the high-elevation, cool-climate regions of Ethiopia. We conducted QTL-seq analysis using 846 RIL F\u003csub\u003e5\u003c/sub\u003e populations of \u0026lsquo;X-Jigna\u0026rsquo;, a dominant rice cultivar in Ethiopia, \u0026times; \u0026lsquo;Hitomebore\u0026rsquo;, a cold-tolerant elite Japanese cultivar, in field trials in Japan. We detected three QTLs for cold tolerance for inducing spikelet sterility under cold at reproductive stage (\u003cem\u003eqSFC5\u003c/em\u003e, \u003cem\u003eqSFC8\u003c/em\u003e, and \u003cem\u003eqSFC10\u003c/em\u003e) and one for seed shattering habit (\u003cem\u003eqSHT1\u003c/em\u003e). In haplotype analysis using indel markers in an F\u003csub\u003e6\u003c/sub\u003e population in Japan and Ethiopia, \u003cem\u003eqSFC5\u003c/em\u003e and \u003cem\u003eqSFC8\u003c/em\u003e were confirmed in Japan and \u003cem\u003eqSHT1\u003c/em\u003e was confirmed in both Japan and Ethiopia. Markers for these will be useful to speed up breeding in Ethiopia.\u003c/p\u003e","manuscriptTitle":"Identification of loci for cold tolerance at reproductive stage and seed shattering in Ethiopian rice cultivar ‘X-Jigna’","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-24 18:35:03","doi":"10.21203/rs.3.rs-8244416/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-03-12T10:37:45+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-12T03:09:19+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-04T15:04:46+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"65752220239403739695064047720964177576","date":"2025-12-30T12:36:04+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"304250143164376085583717006093859719169","date":"2025-12-23T00:59:27+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-12-22T14:51:59+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-12-03T05:34:41+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-12-03T05:34:22+00:00","index":"","fulltext":""},{"type":"submitted","content":"Euphytica","date":"2025-11-30T20:53:01+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"euphytica","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"euph","sideBox":"Learn more about [Euphytica](https://www.springer.com/journal/10681)","snPcode":"10681","submissionUrl":"https://submission.springernature.com/new-submission/10681/3","title":"Euphytica","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b6c8791d-5cb1-4832-8cd4-edb3eceae869","owner":[],"postedDate":"December 24th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-29T12:09:49+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-24 18:35:03","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8244416","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8244416","identity":"rs-8244416","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}