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��니카 벼�서 노화촉진 처리� 따른 종� 활력 � 미질 관련 형질 변화 분�

강경민1,2, �승�1, 하수경1, �길�1, 박재령1, 진민아1, 서정필1, 모�준2, �현숙1,*

Analysis of Seed Vigor and Grain Quality Traits under Accelerated Aging Treatment in japonica Rice

Korean Journal of Breeding Science 2025;57(3):217-230.
Published online: September 1, 2025

1�촌진�청 국립�량과학� 기초�량작물부 품종개발과

2전�대학� 작물�명과학과

1Crop Breeding Division, Department of Crop Sciences, National institute of Crop and Food Science, Rural Development Administration, Wanju, 55365, Republic of KoreaÿØÿàJFIFddÿìDuckydÿáqhttp://ns.adobe.com/xap/1.0/ ÿîAdobedÀÿÛ„ÿÀŠÿÄ     ! 1AQa"q� 2#w8ð‘¡ÁÑB3´µ6v¶7×X±Rr²$˜9áñb’C³Ãt%u&–ÖW·‚s'(xy¢ÂÓ4T”5f‡—ÇH¨¸Ø  !1AQaq"�‘¡2B ±ÁRbr#³´u67ðѲ3sTt5v8áñâSc$4Ä‚’¢C“ÔÔ%UÕÂÒÓ…•FVÅÿÚ ?ßàêÓ_A÷ú‹× ­j- ŠHÌûû>‚>Ì,šm*>½ôßÈÞfzp"áõö—£Téõ‹£—Ær÷Kkr¾ÒÙî��âœ^× »Òr.|_&€ã]¿|*œ™v¿P¶²uܶv”o‡±¿QÇ1Ÿ ¾�ŒŸmÍëw¾VJUöhu-ãIð"=L‹‡n§õ‘iù½AÆ•8Úö�"ûÛ²½þ„� ák�*Ò¿[yùu:ý.Ž‚v«U¤ÒÚQ+•)Ñá%F� D•Hy²V‡è×B훩kš>‘Hêy8øÓj©Ý¹ q~§9Dúñ4íKR‚þm±zõÚíQ„¥ßö)µ^Ê”Ó.J%k_tVi5NTê¥Ójg!'¡k©Çy|5as…¨ðÃOÏÁÈ»)R–¯Û¸ß©FMÔ¿3LÕ4çj¹6dÖùÚœ#êNIwUF©•]J ûq¡BŒô¹/(ËF™aãŠâfJöRzq3à\GÕ›™ ?ö~\£ 6å)6èý•Wµµ4¶m[O›ö^L0ìEË&r¤RMïØ¶*º×ÑC .ÍÂ]Un·l¥Š-1— 1ßr#Rj/¥´ø¯&QɈ׉˩·ás6­Rj=5Tgæ.´êy™.·PÓ¡:ÖJJíS«…÷:îßC8èÝ-Äù„2u]–d�&¢é»Òvøª·Uºz;¾»7p·9‘˜» 5Všã‰nLÖ¢µ£”¢"y)Ц">Ói×™Ñ(ûIDDd|œ¹Ö Yj0·­; L›§ãR¯fÙÞàÝS�˜:k¥ÚÃñtYäK%*ÒN2^ômñ|&dÄŸthó":‡ãÊe§Øy¾)u§P—¶ôQó¡ZöðW)gcC3Pv&“M¾ÇéM¯„ÂWìÏì¬d&¯Å´ãÛ²½�mvŸT¹èRoÞÕªM0Ü3ä*…F3Yº›dûèæãýÃ6¡®èÚ\8õ,¼\h÷Ý» kß”‘õâi¶¡<k× žÞNTõð¦wSëÔªº­Ñmljö•’j[>->±p£tU©Ôâ�%¯'§L‹R>&EÇBïôH$ßÝý�§ÉMQAU¶x[›$Z6’ÒÍv®Ôi&’_“a.K˜IQ{äh©yÆ’½ ’j"JùOCá ê×9òö—ìeäFŒªÒ�´ÞÿfŽjµîí;™ËšÎ©<[3”_m%ÃïÑ÷ ·•l‹¦Q@¥Ö4œ³Ñg²ü�=5$°™(J]YëÁ®c-OMqé<÷ÊÚõÇ�‡“ wSø²�¨¹z’œ«ïÚ—)kš$ÜË7ªV©I¥ïÅPý붾ͯnV+卵*t]i‚ÐŽD¥31ñí~S¨Añí1ûéÃCÐ2u)ÑÂʼnΛãQn-UÅoîi÷3ò´�:®«gß»rI8ý¯ÊØ“Wm*G zà«Ü•)ZÌ×§J}Åê§Yµ Y“lGeJ6ØcB2I ‹NS3×Qµ>kÔõÞ=KT›½ÅBëœâ—ÆÙÎT]ŠŽž‚W鿇¢ÂÞ6œ¬«Ü+ãSÓÛ¦ööï÷Ož—X©QgG©R¦? 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*Corresponding to Hyun-Sook LeeTEL. +82-63-238-5273E-mail. leehs0107@korea.kr
• Received: June 15, 2025   • Revised: August 8, 2025   • Accepted: August 11, 2025

Copyright © 2025 by the Korean Society of Breeding Science

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

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  • The decrease in seed vigor and grain quality during storage has become an increasingly critical issue due to the extended storage periods resulting from declining rice consumption and climate change in Korea. Despite its importance, few studies have investigated rice seed aging in a large number of Korean rice cultivars. In this study, 53 japonica rice cultivars were evaluated for seed germination, vigor, and grain quality-related traits under accelerated-aging conditions. Seed germination rate was evaluated 7 days after imbibition, following accelerated aging treatments at 42℃ and 95% relative humidity for 8, 12, and 16 days. The average germination rates were 72.9% in the control (0 d), 71.7% after 8 days, 59.0% after 12 days, and 5.6% after 16 days of treatment. Grain quality-related traits, including texture and pasting properties, were also evaluated in rice subjected to the accelerated aging treatment. Adhesiveness and stickiness decreased, whereas hardness and toughness increased, as seed aging progressed. Cluster analysis based on germination rates after accelerated aging identified three distinct clusters, with cultivars in Cluster 3 maintaining a high germination rate of 70.6% even after 12 days of aging, showing clear differences from the other clusters. A principal component analysis (PCA) was conducted to investigate the relationship between germination rate and grain quality-related traits. The results showed that the germination rate and hardness tended to be negatively associated across all three clusters. These results suggested that cultivars with higher germination rates tend to maintain a softer cooked rice texture after aging. Our results provide insight into the relationship between seed aging and grain quality, highlighting elite cultivars that maintain both germination ability and grain quality during storage as valuable resources for breeding programs.
벼(Oryza sativa. L)는 전 세계 �구� 절반 ��� 주�으로 소비하는 중요한 �량작물�다(Han et al. 2018). 그러나 ��활� 서구화 � 대체� 선호 등� 사회� 변화� 따� 국내 쌀 소비량� 지��으로 �소하고 있으며, 2005년� 80 kg ���� 1�당 쌀 소비량� 2023년�는 56.4 kg까지 �소하여 쌀 �산량� 소비량� 초과하는 구조�� 공급 과잉 �태가 지��고 있다(USDA Foreign Agricultural Service 2024). �로 �해 매년 30-40만 톤 규모� 쌀� 공공비축미로 매입�어 저장�고 있으며, 국내� 저온 저장 시설 부족으로 �해 대부분� �온 창고�서 톤백 형태로 보관�고 있다(Kim & Kim 2019). �러한 �온 장기 저장� 종� 활력 �소와 밥맛� �함한 미질� 저하�는 문제� ��� �는 것으로 보고�고 있다(Ha et al. 2006, Han et al. 2024, Kim et al. 2007, Tian et al. 2024). �한, 기후변화로 �한 기온 �승으로 �온 장기 저장 시 종� 활력 � 미질 저하 문제가 �욱 심화� 가능성� 제기�고 있다.
종�� 활력� 다양한 환경조건�서 종�가 빠르고 균�하게 발아하여, 건전한 유묘로 성장할 수 있는 능력� �미하고, 수명� 저장 기간 �안 종�가 발아 가능한 �존 �태를 유지하는 능력� �미한다(Finch-Savage & Bassel 2016, Nadarajan et al. 2023). 종�� 활력� 수명�만 아니� 발아력, 초기 �장력 등� �함하는 � 넓� 개�으로 �해�다. 종�� 활력 � 수명� 온�, 습� 등� 저장 환경� �게 �향� 받는다. �반�으로 온�가 높고 종�� 수분 함량� 많�수� 노화가 빠르게 진행�어 활력� 급격히 저하�다. Kawamura et al. (2004)� 연구� 따르면, 수분 함량� 약 15%� ��니카 품종� 정조를 25℃ �온�서 저장한 경우, 약 12개월 후 발아율� 0%까지 �소하였으나, 15℃ 저온 저장�서는 약 48개월, 5℃ 저장 조건�서는 4년 후�� 90% ��� 발아율� 유지하였다. �한, �태형� 따� 종� 수명� 다른�, �반�으로 �디카 품종� ��니카 품종보다 수명� 길다고 알려져 있다(Ellis et al. 1992, Na et al. 2014, Wu et al. 2021). ��니카와 �디카 품종 456개� 유전��� 대�으로 42℃, �대습� 88% 조건�서 20�간 노화촉진 처리 후 발아율� 비�한 결과, ��니카 품종� �균 33.5%, �디카 품종� �균 60.4%로 �태형 간� 유�한 차�를 보였다(Wu et al. 2021). �한, 같� �태형 내�서� 품종� 따� 수명� 차�가 확��었으며, 국내 육성 ��니카 품종�서� 유�한 차�가 보고�었다(Shon et al. 2014). 30개월간 저장한 국내 육성 29개 품종� 발아율� 비�한 결과, �부 품종� 30개월 후�� 80% ��� 발아율� 유지하였으나, �부는 10% �하로 급�하여 품종 간 뚜렷한 차�를 보였다.
종� 저장 기간 �안 온�, 습� 등� 환경조건� 따� 활력 저하�만 아니�, 밥맛� �함한 미질� 저하�는 품질 변화� 수반�다(Han et al. 2024, Kim et al. 2007, Park et al. 2012, Tian et al. 2024). 저장 기간� 따� 변화하는 물리� 특성으로는 경�(hardness), 찰기(stickiness) 등 밥� 물성 변화가 있으며, 보통 저장 기간� 길어질수� 경�는 �가하고, 찰기는 �소하는 경향� 보�다(Gujral & Kumar 2003, Park et al. 2012). 화학� 특성으로는 전분, 단백질, 지질 등 주요 성분� 구조� 변화가 있다(Gu et al. 2025, Hu et al. 2022, Zhao et al. 2021). �한 �미 저하� 주요 �� 중 하나� 고미취 발�� 지방산� 산화� �한 것으로, 고온 조건�서 � 빠르게 진행�는 것으로 보고�었다(Kim et al. 2011). 한편, 아밀로스 � 단백질 함량� 품종 고유� 특성으로, �반�으로 저장 중 변화가 �지 않� 것으로 보고�었다(Kim et al. 2023, Thanompolkrung et al. 2017, Zhou et al. 2002). �러한 물리�, 화학� 변화는 밥� 조��, �미, 호화특성 등� �향� 미� 저장 중 쌀� 품질과 �미를 결정하는 핵심 요�으로 작용한다. �한, �온 장기 저장� 따른 발아율과 �미 사�� 높� �관관계가 보고� 바 있다(Jang et al. 2009, Kim et al. 2023). �와 같� 선행 연구들� 저장 중 종�� 물리�⋅화학� 특성 변화와 그� 따른 미질 저하 현�� 보고하고 있지만, 대부분 �디카 품종 �는 국외 품종� 중심으로 수행�어, 국내 ��니카 품종� 대�으로 한 연구가 미�한 실정�다.
실제 저장 환경�서 장기�으로 종� 활력� �가하기 위해서는 많� 시간과 다양한 환경 요�� �향� 고려해야 하므로, �를 대체할 수 있는 노화촉진(accelerated aging; AA) 처리 기법� �리 사용�고 있다(Delouche & Baskin 1973). 가장 보편�으로 사용�는 방법� 고온다습 조건(40-45℃, �대습� 90-100%)� 배양기�서 10� �내로 종�를 처리하는 방��다(Bijanzadeh et al. 2017, Delouche & Baskin 1973, Kapoor et al. 2011). 고온다습 조건으로 16개 작물� 노화촉진 처리한 후� 발아율� 3년간� 저장 기간 �안� 발아율과 높� �관관계를 보� 바 있다(Delouche & Baskin 1973). �� 본 연구는 국내 ��니카 품종들� 대�으로 고온다습 조건� 노화촉진 처리� 따� 변화하는 종� 활력과 미질 관련 형질� 분�함으로�, 향후 장기저장 조건�서 종� 활력과 미질 관련 형질� 우수한 벼 육종 소재 개발� 위한 기초 �료를 제공하고� 한다.
시험재료 � 노화촉진 처리
본 연구� 사용� 시험재료는 �본�서 육성� 고시히카리, 히토메보레, 추청과 국내�서 육성� ��니카 벼 50개 품종(조�종 9, 중�종 8, 중만�종 36)� 선정하였다(Table 1). �들 품종� 2024년 �촌진�청 국립�량과학� 벼 시험 �장� 4월 30� 파종하여, 5월 27�� 30×15 cm 재�거리로 �앙하였다. 시비량� N-P2O5-K2O=9-4.5-5.7 kg/10a로 하였고, �앙 후� 관리는 �촌진�청 표준 재배법� 준하여 수행하였다. 종�는 출수 �태형� 따� 조�종� 출수 후 40-45�, 중�종� 45-50�, 중만�종� 50-55�� 수확하였다. 종�는 수확 �후 �연�태�서 수분함량 11-13% 수준까지 건조 후, 실험 전까지 �온� 보관하였다. 종� 노화촉진 처리는 Delouche & Baskin (1973) � Qu et al. (2020)� 연구를 참고하여 수확 후 3개월 시�� 정조를 42℃, �대습� 95% 조건�서 8, 12, 16� �안 보관 후 실험� 사용하였다.
종� 발아율 � 활력 조사
노화 촉진 처리 조건�서 8�, 12�, 16� 보관한 종�� 발아율� 조사하였다. 건전한 종�는 tissue embedding cassette� 35립씩 3반복으로 담아 처리 기간별로 보관하였으며, 종� 소�� Ji et al. (2019)� 방법� 참고하였다. 본 연구�서는 70% �탄올� 1분, 0.6% 차아염소산나트륨(NaClO)� 15분간 처리하는 방법으로 수정하여 진행하였다. 소�한 종�는 30℃ 항온기�서 7�간 발아시킨 후, 최종 발아율(%)� 조사하였다.
종� 활력 검정� AOSA (2010)� 테트�졸리움 검사(Tetrazolium-test) 지침서를 참고하여 수행하였다. TZ 검정� 살아있는 세��서 호� 과정 중 발�하는 산화환� 반�� �용하여, triphenyl-tetrazolium chloride (TTC)가 환��어 배 조�� 붉게 염색�는 �리를 기반으로 한다(Porter et al. 1947). TZ 검정� 종�를 절단한 후 염색하여 �가하는 과정� �함하므로, 발아 실험보다 시료당 � 많� 작업 시간� 소요�다(França-Neto & Krzyzanowski 2019). 따�서 본 연구�서는 시료당 작업량� 고려하여 단반복으로 수행하였다. 품종별로 정조 50립씩� �류수� 침지하고 20℃ 항온기�서 18시간 �수 처리한 후, 종�를 절단하여 1% 2,3,5-triphenyl tetrazolium chloride (TTC) 용액� 침지하고 30℃�서 2시간 염색한 후, 배 부위� 염색 여부를 기준으로 전체 종� 중�서 염색� 종�� 비율(%)� 조사하였다.
밥� 물성과 쌀가루� 호화�� 특성 조사
취반 후 밥� 물성 관련 특성� �시프레서(TensipresserTM, My Boy Ⅱ System, Taketomo Electric Inc., Tokyo, Japan)를 �용하였다. 백미 시료 30 g� 물� 첨가하여 가�게 섞어준 뒤 실온(약 25℃)� 30분간 침지 후 전기밥솥(CR-0313V, Cuckoo Electronics Co. Ltd., Yangsan, Korea)� 넣고 취반하였다. 취반한 밥� 10 g씩 정량 한 후, 샬레� 담아 압축 성형 한 후 2분간 �온�서 방치한 후 측정하였다. 측정� two-bite 압착법으로 수행하였으며, 첫 번째 압착� 시료 � �께� 25%까지 눌러 표면층� 물성 특성�, � 번째 압착� 90%까지 압착하여 전체 밥� 물성 특성� 측정하였다. 모든 시료는 3반복으로 측정하였으며, 물성 특성으로는 경�(hardness), 탄력성(toughness), 찰기(stickiness), 부착성(adhesiveness) � 4개 항목� 측정하였다(Lee et al. 2023, Natio & Ogawa 1998, Takahashi et al. 2000).
호화�� 특성� 신���측정기 RVA4500 (Perten Instruments, New South Wales, Australia)� �용하여 측정하였다. RVA 분� 전용 용기� 쌀가루 시료 3 g과 �류수 25 mL를 넣고 온�를 95℃까지 �승시킨 후, 50℃까지 냉�하고 유지하며 �� 변화를 측정하였다. 해당 과정�서 호화개시온�(pasting temperature), 최고��(peak viscosity), 최저��(trough viscosity), 최종��(final viscosity)를 측정하였다. �한, 최고���서 최저��를 뺀 값� 강하��(breakdown viscosity)로, 최종���서 최고��를 뺀 값� 치반��(setback viscosity)로 계산하였다. � 항목� RVA ��는 3회 측정한 값� �균으로 나타내었으며, 단위는 RVU (rapid viscosity unit)로 표기하였다(Park et al. 2021).
통계분�
통계 분�� R (Version 4.0.2, The R Foundation for Statistical Computing Platform)� �용하여 수행하였다. 발아율, 활력, 미질 관련 형질 등� 유�성 검정� 위해 base R� aov 함수를 사용하여 ��분산분�(one-way ANOVA)� 수행하였으며, 처리 �균 간� 유�성 검정� agricolae 패키지� Duncan 다중범위시험법(Duncan’s Multiple Range Test)를 �용하여 5% 유�수준�서 실시하였다. 발아율 변화 패턴� 시�� 분류를 위해 R� umap 패키지를 활용하여 UMAP (Uniform Manifold Approximation and Projection)분�� 수행하였으며, 주요 파�미터는 기본값(n_neighbors=15, min_dist=0.1, metric=‘euclidean’)� 사용하였다. 발아율 특성� 연�성과 군집간 경계� 불명확성� 고려하여 UMAP� 통해 차�� 축소하고, fclust 패키지를 �용한 �지 C-�균 군집화(Fuzzy C-Means Clustering, FCM)를 통해 53개 벼 품종� 군집화하여 � ��터� 군집 소��를 수치화 하였다. FCM� ��터� 군집 소��를 수치화하는 �러스터� 기법으로, 군집화� 신뢰�를 정량�으로 확�하기 위해 사용하였다. 군집 간 발아율과 미질 관련 형질 간� 관계를 분�하기 위해 주성분 분�(Principal Component Analysis, PCA)� 수행하였다. 분�� R� FactoMineR 패키지� PCA 함수를 사용하였다.
노화촉진 처리� 따른 종� 발아율
��니카 벼 53개 품종� 노화촉진 처리(42℃, �대습� 95%) 후 발아율 조사 결과는 Fig. 1A와 Supplementary Table 1� 제시하였다. 전체 시험 품종� 무처리(0�) �균 발아율� 72.9% (범위 36.2-93.3%)였다(Supplementary Table 1). 신�진, 수광 등� 90% ��� 높� 발아율� 보였으나, 해들, 온누리, �호진미 등� 50% �하로 무처리 조건�서� 품종 간 발아율 차�를 보였다. 8� 처리 후�는 �균 71.7% (45.7-88.6%)로 무처리와 유�한 차�는 없었으나, 12� 처리 후�는 59.0% (30.5-83.8%)로 �소하였다. 16� 처리 후�는 �균 5.6% (0.0-27.6%)로 급격히 �소하였다. 한편, 12� 처리 후�� 참�진(83.8%), 신�진1(81.9%), 미소진품(81.9%), 고시히카리와 수광(78.1%), 신�진과 �미벼(75.2%), 히토메보레(73.3%), 현품과 삼광(72.4%), 드래향(71.4%)� 70% ��� 발아율� 유지하였다. 16� 처리 후�는 45개 품종들� 10% �하� 발아율� 보였으나, 참�진(27.6%), 아미쌀(18.1%), 청품(15.2%)� 15% ��� 발아율� 유지하였다.
노화촉진 처리 기간� 길어질수� 발아율� 전반�으로 �소하였으며, 특히 12� 처리 후와 비�했� 때, 16� 처리 후 발아율� 급격하게 �소하였다. �는 고온다습 조건� 노화촉진 처리� 따� 발아율� �소하고, 품종 간 반�� 차�가 있다는 선행 연구 결과와 �치한다(Bijanzadeh et al. 2017, Govindaraj et al. 2017, Kapoor et al. 2011). ��한 처리 조건�서� 품종 간 발아율 차�는 품종 고유� 종� 특성으로 �는 종� 활력 유지력� 우수한 품종� 선발할 수 있는 가능성� 시사한다. 본 연구 결과를 바탕으로 수확 당시 종�� 활력 �태� �향� 받� 수 있지만, 국내 벼 종�� 수명 �가를 위한 노화촉진 처리 기간� 품종 간 변�를 확실히 보�는 8��서 16� 사�가 �절하다고 �단�다. 한편, �디카 품종� 대�으로 40℃, �대습� 100% 조건�서 20�간 노화촉진 처리한 연구�서는 15� 처리가 �가� �합하다는 결과가 있었으나(Govindaraj et al. 2017), �는 �반�으로 �디카 품종� 활력� � 높기 때문으로 보�다. �한, 종� 수명� 유전� 요� 외�� �육 기간, 종� 등숙기, 수확 �후� �르는 전 �육 과정� 환경� 요�� �향� 받는다고 보고�고 있다(Wu et al. 2021). 본 연구�서 무처리 조건� �균 발아율� 낮거나 노화촉진 16� 처리 후 발아율� 급�한 ��� 2024년 여름철 고온� �한 종� 품질 저하� �한 것으로 사료�나, �� 대해서는 추가 연구가 필요하다. 실제로 �부 연구�서는 등숙기� 고온 스트레스가 발아 관련 호르몬 대사 � 전분 분해 효소 유전�발현� 조절하여 발아율� �소시킨다는 보고가 있다(Suriyasak et al. 2020). �, 등숙 초기� 고온� 내배유 발달과 전분 합성� �향� 주어 종�� 충실�를 저하시킬 수 있다는 보고가 있다(Tayade et al. 2018).
노화촉진 처리� 따른 종� 활력
노화촉진 처리� 무처리 종��서 발아율� 낮게 나타난 품종� 활력� 확�하기 위해 TZ 활력 검정� 수행하였다(Fig. 1B, Supplementary Table 1). 무처리(0�)�서는 염색� 종�가 �균 93.5% (범위 82-98%)로 발아율 72.9%보다 높� 활력� 보여, 종� 휴면�나 고온� �한 종� �태 불량으로 �단하였다. 노화촉진 8� 처리 후�는 �균 77.7% (38-90%), 12� 처리 후 72.9% (50-90%), 16� 처리 후 14.3% (0-42%)로 발아율과 유사하게 처리기간� 길어�� 따� 종� 활력� �소하는 것� 확�하였다. 다만, 발아율과 달리 종� 활력� 노화촉진 8� 처리 후 유�한 �소를 보였고, 8� 처리 결과와 12� 처리 결과� 차�는 �대�으로 작았다. 노화촉진 12� 처리 후 활력� 80% ��으로 유지� 품종으로는 케�롱(90.0%), 진미벼(88.0%), �안, 안�, 신�진(86.0%), 한가�, �미벼, 아미쌀(84.0%), 예찬, 신�진, 참누리(82.0%), 참�진, 새누리, 미소진품, 호�, 현품(80.0%) 등 16개 품종� 있었다. 노화촉진 16� 처리 후�� 종� 활력� 25% ��으로 유지� 품종으로는 케�롱(42.0%), 예찬(40.0%), 보르미, 청품(34.0%), 해품, 수광(32.0%), 참�진(26.0%) 등 7개 품종� 있었다. 본 연구�서는 테트�졸리움 염색률� 종� 발아율과 �치하지는 않았지만, 무처리 조건� 낮� 발아율 결과를 해�하기 위한 참고�료로 활용�었으며, 발아율과 높� �관관계를 보�는 선행 연구 결과를 통해 종� 활력� 신�하게 �가할 수 있는 유효한 지표임� 확�하였다(Sukkaew et al. 2023).
노화촉진 처리� 따른 밥� 물성
노화 촉진 처리� 종�� 취반 후 밥 물성 검정� 위하여 �시프레서(Tensipresser)로 부착성(adhesiveness), 탄력성(toughness), 찰기(stickiness), 경�(hardness)� 네 가지 항목� 측정하였다(Table 2, Fig. 2). 노화 촉진 처리� 따른 발아율과 활력 �소 정�를 고려하여 8� 처리 간격� 종�를 활용하였다. 12� 처리 종�는 8� 처리 종�� 발아율, 종� 활력 대비 � 변화가 작아 제외하였다(Fig. 1).
밥� 입안� 달�붙는 정�를 나타내는 부착성� 무처리(0�)�서 51.1, 노화촉진 8� 처리 후 47.7, 16� 처리 후 42.9로 �소하였다(Fig. 2A). 밥� 씹� 후 �래 형태로 �아가는 복�력� 나타내는 탄력성� 무처리�서 39.3, 노화촉진 8� 처리 후 41.0, 처리 16� 후 44.5로 �가하였다(Fig. 2B). 밥알�리 서로 달�붙는 정�를 나타내는 찰기는 무처리�서 65.9, 노화촉진 8� 처리 후 62.7, 16� 처리 후 53.9로 �소하였다(Fig. 2C). 밥� 씹으면서 압착하는� 드는 힘� 나타내는 경�는 무처리�서 55.3, 노화촉진 8� 처리 후 61.8, 16� 처리 후 67.8로 �가하였다(Fig. 2D). 노화촉진 처리 기간� 길어질수� 탄력성과 경�는 �가하고, 부착성과 찰기는 �소하였다. �러한 결과는 고온 장기 저장시 취반 후 밥� 경�가 �가하고 부착성� �소한다는 선행 연구 결과와 �치하였다(Park et al. 2012, Zhou et al. 2007). �반�으로 종�� 노화� 따� 전분� 구조가 변화하고, 수분 보유력� 저하�면서 밥� 물성� 달�지는 경향� 알려져 있으며(Zhou et al. 2007), 본 실험 결과�서� 노화 처리� 따� 밥� 경�가 �가하고 찰기가 �소하는 등 밥� 물성 변화를 확�하였다. �본과 한국 등 �아시아 국가�서는 찰지고 부드러운 ��� 선호하는 경향� 있으며(Chen et al. 2012), 본 실험�서 관찰� 물성 변화는 �러한 선호 경향과는 반대 방향으로 변화하였다. �� 따� 본 연구�서는 53개� 품종� 사용하여 종� 노화 단계� 따른 밥� 물성 변화를 분�함으로� 종� 노화와 �미 저하� 연관성� 시사하였다.
노화촉진 처리� 따른 호화�� 특성
호화�� 특성� 신���측정기(RVA 4500)를 �용하여 조사하였다(Table 2, Fig. 3). RVA 곡선� 전분� 호화를 나타내는 곡선으로, 쌀� 취반 � �미 특성� �가� 유용한 것으로 알려져있다(Cho et al. 2013). 전분� 호화온� ��으로 가열할 경우, 전분 입�가 팽윤하고, 내부� 아밀로스와 아밀로펙틴� 용해�며 ��가 �가하게 �다. 전분� 결정성 구조가 열� �해 붕괴�고, 아밀로스와 아밀로펙틴 분�간� 수소결합� �어지면서, ��� �향� 미치게 �는 과정� 전분� 호화(gelatinization)��고 하며, � 시��서 ��는 최고� �달한다. 전분 입�� 팽윤� 최대� �달한 뒤�는 입�� 구조가 파괴�기 시작하고, �로 �해 ��가 다시 �소하는 현�� 나타난다. �후 냉� 단계�서 가열 중 용출� 아밀로스 분�들� 서로 얽혀 �� 구조를 형성하게 �는 과정�서 수소결합� 통한 분� 간 �호작용으로 전분 입�들� 재결합하면서 �성� 다시 �가하게 �다. �때 RVA 곡선�서 최종�으로 나타나는 ��를 최종��(final viscosity)�고 한다(Lee et al. 2004, Tako et al. 2014).
본 실험�서 노화촉진 처리� 따� 최고��(peak viscosity)는 무처리(0�)�서 245.7 RVU, 8� 처리 후 297.6 RVU, 16� 처리 후 293.6 RVU로 �가하였다(Fig. 3A). 최종��는 무처리�서 129.2 RVU, 노화촉진 8� 처리 후 151.1 RVU, 16� 처리 후 147.2 RVU로 �가하였다(Fig. 3B). �� 따� 최종��와 최고��� 차�를 나타내는 치반��(setback)는 무처리�서 -38.6 RVU, 노화촉진 8� 처리 후 -62.4 RVU, 16� 처리 후 -58.7 RVU로 �소하였다(Fig. 3C). 최고��와 최저��� 차�를 나타내는 강하��(breakdown)는 호화중� 열, 전단(shear)� 대한 저항 정�를 �미하며(Sim et al. 2015), 무처리�서 114.8 RVU, 노화촉진 8� 처리 후 146.5 RVU, 16� 처리 후 146.3 RVU로 �가하는 양�� 보였다(Fig. 3D). 본 실험�서� 호화�� 특성� 종�활력, 물성과 달리 무처리(0�)�서 노화촉진 8� 처리 사�� 급격하게 변화하였으나, �후 기간�는 �게 변화하지 않았다. �는 호화�� 특성� 노화 진행� 따� 가장 빠르게 변화하는 형질� 가능성� 시사한다. 노화처리� 따� 최고��, 최종��, 강하��는 �가, 치반��는 �소하였으며, 모든 항목�서 통계�으로 유�한 차�가 확��었다. �러한 결과는 Shi et al. (2022)� 7년간 장기 저장� 쌀�서 높� 최고��와 강하��, 낮� 치반��를 보였다는 결과와 유사하다. �한 �연조건�서 노화� 쌀�서 최고��가 �가한다는 결과와� 유사하다(Pearce et al. 2001, Soponronnarit et al. 2008, Wu et al. 2016). �러한 변화는 노화 과정�서 전분� 구조 변화, 특히 팽윤력� 변화와 관련� 있다. Park et al. (2012)와 Zhou et al. (2015)는 호화�� 특성� 노화 �태를 민�하게 반�하는 지표가 � 수 있�� 제시하였고, Jacobs (1995) �한 전분� 팽윤력(swelling)과 전단 저항성(anti-shearing ability)� 변화가 호화�� 특성과 밀접한 관련� 있다고 보고하였다. Shu et al. (2021)� 저장 중 쌀� 효소 활성� �소하고 전분과 지질 복합체 형성� �가함� 따� 최고�� � 최종��가 �승하는 경향� 나타난다고 보고하였다. �러한 선행 연구들� 본 연구 결과와 함께 종� 노화가 전분 호화특성 변화와 연관�어 있�� 시사한다.
발아율 변화 패턴� 따른 군집 분�
본 연구� 사용� ��니카 벼 53개 품종� 발아율 변화 패턴� 따� UMAP 차� 축소 후 Fuzzy C-Means (FCM) 알고리즘� �용한 결과, � 세 개� 군집으로 분류�었다(Fig. 4). 모든 품종� 군집 소� 확률 값� 0.8 ��으로 나타나 군집간 경계가 비�� 명확하게 구분�었�� 확�하였다. � 군집� 발아율 특성� Table 3�, 해당 군집� �한 품종� Figs. 5D, 5E, 5F� 제시하였다. 군집 1� 남�벼, 온누리 등 17개 품종으로 구성�었으며, 무처리(0�) �균 발아율� 가장 낮고, 노화촉진 8� 처리 후 �시�으로 �가하여 휴면성� 있는 군집으로 해��었다. 군집 2는 고시히카리, 히토메보레, �품벼 등 18개 품종� �함�었으며, 무처리 �균 발아율� 73.3%로 중간 수준�었고, 노화촉진 처리 전체기간 �안 군집 1과 3� 중간 �균값� 보였다. 군집 3� 삼광, 신�진, 참�진 등 18개 품종으로, 무처리 �균 발아율� 86.4%로 가장 높았으며, 노화촉진 12� 처리 후 �균 발아율� 70.6%로 세 개 군집 중 가장 높아, 종� 활력� 높게 유지�는 군집으로 해��었다. �러한 군집 분� 결과는 처리 기간� 따른 발아율 변화 양�� 고려하여, ��니카 품종 내�서 세 개� 군집� 구분했다는 ��서 �미가 있다. 본 연구 결과�서 선발� 종� 활력 유지 품종들� 추후 육종 소재로 활용� 가능성� 있다.
발아율과 미질 관련 형질� 주성분 분�
발아율과 미질 관련 형질� 관계성� 조사하기 위하여 PCA 분�� 수행하였다. 분� 결과 전체 형질 분��서 발아율 벡터� 기여�가 낮아 발아율 수준� 따른 형질별 구조 파악� 위하여 세 군집별(Fig. 4, Table 3) PCA 분�� 실시하였다(Fig. 5). 그 결과 군집 1�서는 주성분 1과 2를 통해 전체 변�� 51.4%, 군집 2는 46.4%, 군집 3� 34.6%를 설명할 수 있었다. 군집 1�서는 발아율(GR8, GR12)과 탄력성(TH), 경�(HD)가 주성분 2를 기준으로 서로 반대 방향성� 보였고(Fig. 5A), 군집 2�서는 GR8과 GR12가 경�(HD), 치반��(SB), 최종��(FV)와 반대 방향� 보였다(Fig. 5B). 군집 3�서는 발아율과 탄력성(TH), 경�(HD), 치반��(SB), 최종��(FV)가 반대 방향� 보였다(Fig. 5D).
�외� 호화�� 특성 중�서 최고��(PV)와 강하��(BD), 최종��(FV)와 치반��(SB)는 모든 그룹�서 강한 �관관계를 보였으며, 밥� 물성�서는 경�(HD)와 탄력성(TH), 찰기(ST)와 부착성(AD)� 강한 �관관계를 보였다(Figs. 5D, 5E, 5F). �들 특성간� 관계성� Park et al. (2024)와 Kim et al. (2014)� 연구�서� 보고� 바 있다.
군집 1�서� 품종 분�는 전체�으로 산��어 있지만, 진광� 높� 최고��(PV), 강하��(BD)와 낮� 발아율� 보여 다른 품종들과 분리�어 위치하였다(Fig. 5D). 군집 2는 �본 품종� 고시히카리와 히토메보레가 다른 품종들과 구분�며 높� 최고��(PV)와 낮� 치반��(SB)를 가지는 것으로 확��었다(Fig. 5E). 히토메보레는 고시히카리와 하츠보시가 �배� 품종으로 유전� 유사성� 매우 높으며, � 품종 모� 높� 최고��를 보�다는 선행 연구 결과가 있었다(Lestari et al. 2009). 군집 3�서는 노화처리 후� 발아율� 높게 유지�는 참�진, �안, 신�진, 신�진1 등� 가까� 분�하고 있으며 강하��(BD)와 최고��(PV)가 높고, 경�가 낮� 것� 확�하였다. 최고��와 강하��는 �미치와 유�한 양� �관관계를 보�다는 보고가 있으며(Hwang et al. 2007, Kwon et al. 1991, Matsue & Ogata 2000), 국내�서는 부드러운 ��� 선호하기 때문� �들 품종� 종� 활력과 미질� 유지�는 품종� 것으로 기대�다.
본 결과를 통하여 종� 활력 유지 정�와 �부 미질 관련 특성� 관계성� 확�하였고, �는 활력과 미질� 장기간 유지�는 품종 선발 시 중요한 지표로 활용� 수 있다. �한 그룹 3� �함� 종� 활력� 높게 유지�는 품종들� 추후 육종 소재로 활용할 수 있다.
본 연구는 ��니카 벼 품종� 노화촉진 처리� 따� 종� 활력 � 미질 관련 형질� 변화하는 양�� �가하고, 노화촉진 처리 후 종�활력� 유지�는 품종� 선발하여 �를 육종 소재로 활용할 수 있는 가능성� 확�하고� 하였다. ��니카 벼 53개 품종� 대�으로 42℃, �대습� 95%� 조건�서 0, 8, 12, 16�간 종�를 노화촉진 처리한 후, 발아율 � 테트�졸리움 활력 검정, 밥� 물성(부착성, 탄력성, 찰기, 경�), 호화�� 특성(최고��, 최종��, 치반��, 강하��)� 조사하였다. 그 결과, 종� 발아율와 활력� 노화촉진 처리 12� 후 유�하게 �소하였으며, 16� 처리 후�는 발아율 �균� 5.6%로 급격히 �소하였다. 노화촉진 12� 처리 후 결과를 기준으로 발아율과 종� 활력 검정 결과를 종합�으로 고려한 결과, 참�진, 신�진1, 미소진품 등� 노화� 대한 내성� 높� 품종으로 �단�다. 밥� 물성� 노화촉진 처리 기간� �가� 따� 완만하게 변화하였으며, 탄력성과 경�는 �가하고, 부착성과 찰기는 �소하는 경향� 보였다. 호화�� 특성� 노화촉진 처리 8� 후 급격히 변화하였고, �후�는 변화가 �지 않았으며, 노화촉진 처리 기간� 길어질수� 최고��, 최종��, 강하��는 �가하였고, 치반��는 �소하는 경향� 보였다. 노화촉진 처리 기간� 따른 종� 발아율 변화 양�� 기반으로 한 군집 분� 결과, ��니카 벼 53개 품종� � 세 개� 군집으로 나뉘어졌다. 노화촉진 처리� 따른 발아율과 미질 관련 형질� 주성분 분� 결과, 세 개� 군집�서 ��하게 발아율과 밥� 물성 중 경�가 높� 관계성� 보였다. 특히, 군집 3� �한 품종들 중 발아율� 높게 유지�면서 국내�서 선호하는 낮� 경�와, 높� 강하��� 특성� 가진 참�진, �안, 신�진, 신�진1 등� 향후 장기 저장 조건�서 종� 활력과 미질 특성� 우수한 벼 품종� 개발하기 위한 육종 소재로 활용가능할 것으로 기대�다.
본 연구는 �촌진�청 국립�량과학�� 시험연구사업(벼 저장성 � �열병 저항성 향� 육종소재 개발, 과제번호: PJ01726902)� 지�으로 수행�었습니다. �촌진�청 국립�량과학�과 전�대학�� 많� �움� �사드립니다.
Fig. 1
Seed vigor assessment after accelerated aging treatment. (A) Germination percentage and (B) Viability of 53 rice cultivars after 0, 8, 12, and 16 days of accelerated aging. Error bars represent standard error (SE).
kjbs-57-3-217-f1.jpg
Fig. 2
Changes in texture properties of cooked rice after accelerated aging treatment. (A) Adhesiveness, (B) Toughness, (C) Stickiness, and (D) Hardness of cooked rice from 53 rice cultivars after 0, 8 and 16 days of accelerated aging. Error bars represent standard error (SE).
kjbs-57-3-217-f2.jpg
Fig. 3
Changes in pasting properties of rice flour after accelerated aging treatment. (A) Peak viscosity, (B) Final viscosity, (C) Setback viscosity, and (D) Breakdown viscosity of rice flour from 53 rice cultivars after 0, 8 and 16 days of accelerated aging. Error bars represent standard error (SE).
kjbs-57-3-217-f3.jpg
Fig. 4
Clustering analysis of germination percentage patterns in 53 rice cultivars under accelerated aging treatment. (A) Projection of high-dimensional germination rate data using UMAP (Uniform Manifold Approximation and Projection) followed by clustering using the Fuzzy C-Means (FCM) algorithm. (B) Germination rate change patterns of each cluster classified by FCM. Thin lines represent individual cultivars, and bold lines indicate the cluster mean.
kjbs-57-3-217-f4.jpg
Fig. 5
Principal component analysis (PCA) of germination and grain quality traits for three rice clusters.(A-C) Score plots of rice cultivars in Cluster 1 (A), Cluster 2 (B), and Cluster 3 (C). (D-F) Corresponding PCA biplots for Cluster 1 (D), Cluster 2 (E), and Cluster 3 (F), showing both sample distributions and trait vectors. The PCA includes germination rate traits: GR0, GR8, GR12, and GR16; pasting properties: PV (peak viscosity), TV (trough viscosity), FV (final viscosity), BD (breakdown viscosity), and SB (setback viscosity); and texture properties: AD (adhesiveness), ST (stickiness), TH (toughness), and HD (hardness). Arrows are color-coded by trait group for clarity: black for germination rate, blue for pasting properties, and red for texture properties. The direction and length of each arrow indicate the magnitude and contribution of the variable to principal components.
kjbs-57-3-217-f5.jpg
Table 1
List of the 53 rice cultivars used in this study, categorized by maturity group.
Table 1
Maturity Group
(Heading date range)
Cultivars
Early
(7/19-7/31)
Boreumi, Dapyeong, Haedamssal, Haedeul, Hitomebore*, Jinkwang, Jinmibyeo, Saeodae1ho, Unkwang
Midium
(8/1-8/10)
Alchanmi, Chamjinmi, Chamnuri, Cheongpum, Daebo, Hangadeuk, Koshihikari*, Samkwang
Mid-Late
(8/10-8/19)
Amissal, Anpyeong, Boramchan, Chamdongjin, Chilbo, Chuchung*, Dangchanjinmi, Deuraehyang, Donganbyeo, Haepum, Haiami, Honong, Hopum, Hopyeong, Hwangkeumnuri, Hwasinbyeo, Hyeonpum, Ilmibyeo, Ilpumbyeo, Jinbaek, Jinsumi, K-Long, Mangeumbyeo, Mipum, Misojinpum, Namchan, Nampyeongbyeo, Onnuri, Pyeongan, Saebonghwang, Saenuri, Sindongjin, Sindongjin1, Sukwang, Yechan, Younghojinmi

*Japanese cultivars

Table 2
Changes in texture properties and pasting properties after accelerated aging treatment.
Table 2
Trait Treatment duration (days)

0 8 16
Texture
properties
Adhesiveness 51.1±0.6 az 47.7±0.8 b 42.9±0.8 c
Toughness 39.3±0.4 c 41.0±0.5 b 44.5±0.5 a
Stickiness 65.9±1.4 a 62.7±1.2 a 53.9±0.7 b
Hardness 55.3±0.9 c 61.8±1.3 b 67.8±0.9 a
Pasting
properties
(RVU)
Peak viscosity 245.7±3.3 b 297.6±3.3 a 293.6±2.5 a
Final viscosity 207.1±2.9 b 235.2±2.8 a 234.9±2.2 a
Setback -38.6±3.3 a -62.4±3.5 b -58.7±2.7 b
Breakdown 114.8±2.7 b 146.5±3.0 a 146.3±2.1 a

Values are means±standard error.

zDifferent letters within a row indicate significant differences within each cluster according to Duncan’s multiple range test (DMRT, p<0.05).

Table 3
Mean germination percentages of three clusters during accelerated aging treatment.
Table 3
Cluster Treatment duration (days)

0 8 12 16
Cluster 1 58.1±2.8 cz 65.8±2.7 a 44.3±2.4 c 4.8±0.8 a
Cluster 2 73.3±2.2 b 71.1±2.3 a 61.2±1.8 b 7.8±1.2 a
Cluster 3 86.4±1.1 a 77.9±1.4 a 70.6±1.9 a 4.0±1.5 a

Values are means±standard error.

zDifferent letters within a column indicate significant differences within each cluster according to Duncan’s multiple range test (DMRT, p<0.05).

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Analysis of Seed Vigor and Grain Quality Traits under Accelerated Aging Treatment in japonica Rice
Korean. J. Breed. Sci.. 2025;57(3):217-230.   Published online September 1, 2025
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Analysis of Seed Vigor and Grain Quality Traits under Accelerated Aging Treatment in japonica Rice
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Analysis of Seed Vigor and Grain Quality Traits under Accelerated Aging Treatment in japonica Rice
Image Image Image Image Image
Fig. 1 Seed vigor assessment after accelerated aging treatment. (A) Germination percentage and (B) Viability of 53 rice cultivars after 0, 8, 12, and 16 days of accelerated aging. Error bars represent standard error (SE).
Fig. 2 Changes in texture properties of cooked rice after accelerated aging treatment. (A) Adhesiveness, (B) Toughness, (C) Stickiness, and (D) Hardness of cooked rice from 53 rice cultivars after 0, 8 and 16 days of accelerated aging. Error bars represent standard error (SE).
Fig. 3 Changes in pasting properties of rice flour after accelerated aging treatment. (A) Peak viscosity, (B) Final viscosity, (C) Setback viscosity, and (D) Breakdown viscosity of rice flour from 53 rice cultivars after 0, 8 and 16 days of accelerated aging. Error bars represent standard error (SE).
Fig. 4 Clustering analysis of germination percentage patterns in 53 rice cultivars under accelerated aging treatment. (A) Projection of high-dimensional germination rate data using UMAP (Uniform Manifold Approximation and Projection) followed by clustering using the Fuzzy C-Means (FCM) algorithm. (B) Germination rate change patterns of each cluster classified by FCM. Thin lines represent individual cultivars, and bold lines indicate the cluster mean.
Fig. 5 Principal component analysis (PCA) of germination and grain quality traits for three rice clusters.(A-C) Score plots of rice cultivars in Cluster 1 (A), Cluster 2 (B), and Cluster 3 (C). (D-F) Corresponding PCA biplots for Cluster 1 (D), Cluster 2 (E), and Cluster 3 (F), showing both sample distributions and trait vectors. The PCA includes germination rate traits: GR0, GR8, GR12, and GR16; pasting properties: PV (peak viscosity), TV (trough viscosity), FV (final viscosity), BD (breakdown viscosity), and SB (setback viscosity); and texture properties: AD (adhesiveness), ST (stickiness), TH (toughness), and HD (hardness). Arrows are color-coded by trait group for clarity: black for germination rate, blue for pasting properties, and red for texture properties. The direction and length of each arrow indicate the magnitude and contribution of the variable to principal components.
Analysis of Seed Vigor and Grain Quality Traits under Accelerated Aging Treatment in japonica Rice

List of the 53 rice cultivars used in this study, categorized by maturity group.

Maturity Group
(Heading date range)
Cultivars
Early
(7/19-7/31)
Boreumi, Dapyeong, Haedamssal, Haedeul, Hitomebore*, Jinkwang, Jinmibyeo, Saeodae1ho, Unkwang
Midium
(8/1-8/10)
Alchanmi, Chamjinmi, Chamnuri, Cheongpum, Daebo, Hangadeuk, Koshihikari*, Samkwang
Mid-Late
(8/10-8/19)
Amissal, Anpyeong, Boramchan, Chamdongjin, Chilbo, Chuchung*, Dangchanjinmi, Deuraehyang, Donganbyeo, Haepum, Haiami, Honong, Hopum, Hopyeong, Hwangkeumnuri, Hwasinbyeo, Hyeonpum, Ilmibyeo, Ilpumbyeo, Jinbaek, Jinsumi, K-Long, Mangeumbyeo, Mipum, Misojinpum, Namchan, Nampyeongbyeo, Onnuri, Pyeongan, Saebonghwang, Saenuri, Sindongjin, Sindongjin1, Sukwang, Yechan, Younghojinmi

Changes in texture properties and pasting properties after accelerated aging treatment.

Trait Treatment duration (days)

0 8 16
Texture
properties
Adhesiveness 51.1±0.6 az 47.7±0.8 b 42.9±0.8 c
Toughness 39.3±0.4 c 41.0±0.5 b 44.5±0.5 a
Stickiness 65.9±1.4 a 62.7±1.2 a 53.9±0.7 b
Hardness 55.3±0.9 c 61.8±1.3 b 67.8±0.9 a
Pasting
properties
(RVU)
Peak viscosity 245.7±3.3 b 297.6±3.3 a 293.6±2.5 a
Final viscosity 207.1±2.9 b 235.2±2.8 a 234.9±2.2 a
Setback -38.6±3.3 a -62.4±3.5 b -58.7±2.7 b
Breakdown 114.8±2.7 b 146.5±3.0 a 146.3±2.1 a

Mean germination percentages of three clusters during accelerated aging treatment.

Cluster Treatment duration (days)

0 8 12 16
Cluster 1 58.1±2.8 cz 65.8±2.7 a 44.3±2.4 c 4.8±0.8 a
Cluster 2 73.3±2.2 b 71.1±2.3 a 61.2±1.8 b 7.8±1.2 a
Cluster 3 86.4±1.1 a 77.9±1.4 a 70.6±1.9 a 4.0±1.5 a
Table 1 List of the 53 rice cultivars used in this study, categorized by maturity group.

*Japanese cultivars

Table 2 Changes in texture properties and pasting properties after accelerated aging treatment.

Values are means±standard error.

zDifferent letters within a row indicate significant differences within each cluster according to Duncan’s multiple range test (DMRT, p<0.05).

Table 3 Mean germination percentages of three clusters during accelerated aging treatment.

Values are means±standard error.

zDifferent letters within a column indicate significant differences within each cluster according to Duncan’s multiple range test (DMRT, p<0.05).