超级稻南粳5718茎秆特性及其抗倒分子机制

doi:10.7668/hbnxb.20196364

摘要/Abstract

摘要：

超级稻品种南粳5718被列为农业农村部主导品种,在大面积生产中抗倒性表现出色。研究南粳5718的茎秆特性,解析其抗倒伏的分子机制,能够为抗倒伏水稻育种提供新的思路。以抗倒性较强的超级稻品种南粳5718和2个抗倒性较弱品种南粳518和南粳58为材料,比较农艺性状、茎秆特性与抗倒伏的相关性,发现南粳5718抽穗后20 d的株高、重心高度显著低于南粳518和南粳58,倒二节间长度、倒二和倒三节间的直径和茎壁厚度均高于南粳518和南粳58。茎秆结构性和非结构性碳水化合物含量测定结果表明,倒二和倒三节间的纤维素和木质素含量高是导致南粳5718强抗倒性的重要因素。进一步分析发现,3个品种倒二节间纤维素酶活性无显著差异。南粳5718和另外2个品种相比,102个纤维素和木质素合成代谢途径相关基因中只有pPLAⅢα(编码糖蛋白相关磷脂酶A)、CBF诱导因子bHLH002、ILA1(编码丝裂原活化蛋白激酶)和转录因子基因NAC5在编码区存在错义突变。正调控纤维素合成的转录因子MYB61、负调控纤维素合成的细长秆基因SLR1和PIL13、多半乳糖醛酸酶PG3、乙烯响应因子ERF34、矮秆基因DPH1、转录因子MYB7等基因的相对表达量在南粳5718和南粳58之间存在显著差异。木质素代谢过程中的4-香豆酸:辅酶A连接酶4CL3、MYB转录因子RLM1、对香豆素酯3-羟化酶CYP98A4、NAC转录因子SWN1和苯丙氨酸脱氨酶基因PAL1等基因的相对表达量在南粳5718和南粳58之间存在显著差异。关键基因在节间的表达差异是南粳5718茎秆高纤维素和木质素含量导致强抗倒性的原因。综上,超级稻南粳5718强抗倒性在茎秆特性上表现为具有较粗的节间直径和较大的茎壁厚度,节间较高的纤维素与木质素含量为其抗倒性提供了物质基础,纤维素和木质素合成通路中关键基因表达差异可能是南粳5718茎秆强抗倒性的主要原因。

关键词: 水稻, 抗倒伏, 茎秆, 纤维素, 分子机制

Abstract:

The super rice variety Nanjing 5718 has been designated as a leading variety by the Ministry of Agriculture and Rural Affairs and has demonstrated outstanding lodging resistance in large-scale production.Studying the stem characteristics of Nanjing 5718 and elucidating the molecular mechanisms underlying its lodging resistance can provide new insights for breeding lodging-resistant rice varieties.The lodging-resistant super rice Nanjing 5718 and two varieties,Nanjing 518 and Nanjing 58,with weak lodging resistance,were used as materials to compare agronomic traits,stem characteristics and their correlation with lodging resistance.The plant height and center of gravity of Nanjing 5718 were significantly lower than those of other varieties at 20 days after heading.The length of the second basal internode,stem diameter and stem wall thickness of the second and third basal internodes of Nanjing 5718 were significantly higher than other varieties at 20 days after heading.High content of cellulase and lignin in the stem was the important factor leading to strong lodging resistance.No significant differences in cellulase activity were observed in the second basal internodes among the three varieties.Compared to the other two varieties,only the pPLAⅢα,bHLH002,ILA1 and NAC5 among the 102 genes related to the cellulose and lignin biosynthetic pathways had missense mutations in their coding regions in Nanjing 5718.The relative expression levels of key cellulose biosynthesis-regulating genes,such as MYB61, SLR1,PIL13, PG3, ERF34,DPH1 and MYB7 were significantly different between Nanjing 5718 and Nanjing 58.Significant differences were also observed in the relative expression levels of some lignin metabolism genes between Nanjing 5718 and Nanjing 58:4CL3,RLM1,CYP98A4,SWN1,PAL1,and so on.The differential expression of those genes in basal internodes likely underlies the high cellulose and lignin content of stem and consequent strong lodging resistance in Nanjing 5718.In conclusion,the strong lodging resistance of the super rice variety Nanjing 5718 results from its increased internode diameter and stem wall thickness.The higher contents of cellulose and lignin in the internodes provide a structural basis for this enhanced lodging resistance.Variations in the expression of key genes involved in the cellulose and lignin biosynthesis pathways may be the primary factor contributing to the superior lodging resistance observed in Nanjing 5718.

Key words: Rice, Lodging resistance, Stem, Cellulose, Molecular mechanism

中图分类号:

宋悦, 安红强, 陈盈盈, 赵庆勇, 李程, 朱镇, 梁文化, 周丽慧, 赵凌, 朱克明, 张亚东. 超级稻南粳5718茎秆特性及其抗倒分子机制[J]. 华北农学报, 2026, 41(4): 1-11. doi: 10.7668/hbnxb.20196364.

SONG Yue, AN Hongqiang, CHEN Yingying, ZHAO Qingyong, LI Cheng, ZHU Zhen, LIANG Wenhua, ZHOU Lihui, ZHAO Ling, ZHU Keming, ZHANG Yadong. Stem Characteristics of Super Rice Nanjing 5718 and the Molecular Mechanism of Its Lodging Resistance[J]. Acta Agriculturae Boreali-Sinica, 2026, 41(4): 1-11. doi: 10.7668/hbnxb.20196364.

http://www.hbnxb.net/CN/Y2026/V41/I4/1

图/表 8

表1 荧光定量PCR所用引物序列

Tab.1 Primer sequences for fluorescence quantitative PCR

基因名称 Gene name	基因功能 Gene function	正向引物序列(5'—3') Forward primer sequence(5'—3')	反向引物序列(5'—3') Reverse primer sequence(5'—3')
UBQ	内参基因	CAAGATGATCTGCCGCAAATGC	TTTAACCAGTCCATGAACCCG
SLR1	阻止GA信号向下传导	GATCGTCACCGTGGTAGAGC	GAGGGAATCGAACATGGTGG
PIL13	在GA信号调节中发挥作用	GCAAACAGTGCCACCACAGG	CTAAATTCCATCAGAGGTTGG
CesA7	负责催化纤维素的合成	AAGCCATGCGGGGTCTCGTG	CATCCATCCGGTCATCCCTCTTG
MYB61	激活纤维素合成酶基因表达	ATATCTTGAGCCACTTGT	ATCCTCTTGGTGATTCAG

图1 3个品种抽穗后20 d茎秆形态学及力学指标图中不同小写字母表示不同水稻品种的农艺性状在P<0.05水平差异显著。图3—5同。

Fig.1 Stem morphology and mechanical indexes of three varieties at 20 days after heading Different lowercase letters in the figure indicate that there are significant differences in agronomic traits among different rice varieties at P<0.05.The same as Fig.3—5.

表2 农艺性状、形态性状和倒伏指数的相关性分析

Tab.2 Correlation analysis between lodging index and agronomic and morphological traits

性状 Traits	相关系数 Correlation coefficient
株高 Plant height	-0.23
穗数 Panicles	0.19
实粒数 Filled grain number	-0.51^**
总粒数 Total grains	-0.65^**
结实率 Setting rate	0.33
千粒质量 1 000 grains weight	0.57^**
产量 Yield	0.34
重心高度 Height of center of gravity	-0.70^**
倒二节间长度 Basal second internodes length	0.83^**
倒三节间长度 Basal third internodes length	0.32
倒二节间直径 Basal second internodes diameter	0.52^**
倒二节间厚度 Basal second internodes thickness	0.22
倒三节间直径 Basal third internodes diameter	0.40^*
倒三节间厚度 Basal third internodes thickness	0.43^*

图2 南粳5718和南粳58倒二节间组织化学染色

Fig.2 Histochemical staining of the basal second internodes of Nanjing 5718 and Nanjing 58

图3 3个品种抽穗后20 d茎秆倒二和倒三节间结构性及非结构性碳水化合物含量

Fig.3 The content of structural and non-structural carbohydrates in the basal second and third internodes of the three varieties at 20 days after heading

图4 3个品种抽穗后20 d倒二和倒三节间纤维素酶活性

Fig.4 The cellulase activity in the basal second and third internodes of the three varieties at 20 days after heading

图5 不同品种抽穗后20 d茎秆中纤维素代谢通路关键基因的相对表达量

Fig.5 The relative expression of key genes in cellulose metabolic pathway in stems of different varieties at 20 days after heading

图6 南粳5718和南粳58茎秆中纤维素(A)和木质素(B)代谢通路相关基因表达热图

Fig.6 Expression heatmap of genes related to cellulose(A)and lignin(B) metabolic pathway in stems of Nanjing 5718 and Nanjing 58

参考文献 46

[1]	王睿智, 李涛, 钱相玲, 张丫, 杨晓琢, 李光彦, 魏海燕, 张洪程, 刘国栋. 氮素穗肥对南方香粳稻南粳9108产量、品质和香气的影响[J]. 中国农业科学, 2025, 58(12):2316-2332.doi:10.3864/j.issn.0578-1752.2025.12.004.
	Wang R Z, Li T, Qian X L, Zhang Y, Yang X Z, Li G Y, Wei H Y, Zhang H C, Liu G D. Effects of nitrogen panicle fertilizer on yield,quality and aroma of southern japonica rice Nanjing 9108[J]. Scientia Agricultura Sinica, 2025, 58(12):2316-2332.
[2]	Mullangie D P, Thiyagarajan K, Swaminathan M, Ramalingam J, Natarajan S, Govindan S. Breeding resilience:exploring lodging resistance mechanisms in rice[J]. Rice Science, 2024, 31(6):659-672.doi:10.1016/j.rsci.2024.08.002.
[3]	陆展华, 王晓飞, 刘维, 卢东柏, 王石光, 薛皦, 何秀英. 优质稻粤农丝苗抗倒伏影响因素和遗传分析[J]. 植物遗传资源学报, 2021, 22(3):638-645.doi:10.13430/j.cnki.jpgr.20200918003.
	Lu Z H, Wang X F, Liu W, Lu D B, Wang S G, Xue J, He X Y. Influencing factors and genetic analysis of lodging resistance of high-quality rice Yuenong Simiao[J]. Journal of Plant Genetic Resources, 2021, 22(3):638-645.
[4]	田文涛, 邵平, 王燚, 魏中伟, 王晓玲, 马国辉. 超级杂交稻茎秆形态结构及其与抗倒性的关系研究[J]. 杂交水稻, 2017, 32(2):67-71,81.doi:10.16267/j.cnki.1005-3956.201702019.
	Tian W T, Shao P, Wang Y, Wei Z W, Wang X L, Ma G H. Stem morphological structure and its relation to lodging resistance of super hybrid rice[J]. Hybrid Rice, 2017, 32(2):67-71,81.
[5]	汪清焰, 刘斌美, 杨阳, 叶亚峰, 任艳, 吴跃进. 水稻脆茎突变体细胞壁组分与茎秆力学性能的研究[J]. 生物学杂志, 2020, 37(1):26-29.doi:10.3969/j.issn.2095-1736.2020.01.026.
	Wang Q Y, Liu B M, Yang Y, Ye Y F, Ren Y, Wu Y J. Cell wall components and mechanical properties in brittle culm mutants of rice[J]. Journal of Biology, 2020, 37(1):26-29.
[6]	潘俊峰, 李国辉, 崔克辉. 水稻茎鞘非结构性碳水化合物再分配及其在稳产和抗逆中的作用[J]. 中国水稻科学, 2014, 28(4):335-342.doi:10.3969/j.issn.1001-7216.2014.04.001.
	Pan J F, Li G H, Cui K H. Re-partitioning of non-structural carbohydrates in rice stems and their roles in yield stability and stress tolerance[J]. Chinese Journal of Rice Science, 2014, 28(4):335-342.
[7]	Zhao Y, Gao J, Wang X H, Rashid M A R, Wu Z W, Ma Z Q, Wu H, Xu B X, Wu Z Y, Gu Y S, Pan Y H, Li D T, Wang R Y, Guo Z H, Ma W D, Sun X M, Li J J, Zhang H L, Mao T L, Zhang Z Y, Li Z C. Elite haplotype of STRONG1 enhances rice yield by improving lodging resistance,panicle and plant architecture[J]. Nature Communications, 2025, 16(1):5894.doi:10.1038/s41467-025-60604-9.
[8]	李刚, 高清松, 李伟, 张雯霞, 王健, 程保山, 王迪, 高浩, 徐卫军, 陈红旗, 纪剑辉. 定向敲除SD1基因提高水稻的抗倒性和稻瘟病抗性[J]. 中国水稻科学, 2023, 37(4):359-367.doi:10.16819/j.1001-7216.2023.221113.
	Li G, Gao Q S, Li W, Zhang W X, Wang J, Cheng B S, Wang D, Gao H, Xu W J, Chen H Q, Ji J H. Directed knockout of SD1 gene improves lodging resistance and blast resistance of rice[J]. Chinese Journal of Rice Science, 2023, 37(4):359-367.
[9]	Tu B, Tao Z, Wang S G, Zhou L, Zheng L, Zhang C, Li X Z, Zhang X Y, Yin J J, Zhu X B, Yuan H, Li T, Chen W L, Qin P, Ma B T, Wang Y P, Li S G. Loss of Gn1a/OsCKX2 confers heavy-panicle rice with excellent lodging resistance[J]. Journal of Integrative Plant Biology, 2022, 64(1):23-38.doi:10.1111/jipb.13185. URL
[10]	Yano K, Ookawa T, Aya K, Ochiai Y, Hirasawa T, Ebitani T, Takarada T, Yano M, Yamamoto T, Fukuoka S, Wu J Z, Ando T, Ordonio R L, Hirano K, Matsuoka M. Isolation of a novel lodging resistance QTL gene involved in strigolactone signaling and its pyramiding with a QTL gene involved in another mechanism[J]. Molecular Plant, 2015, 8(2):303-314.doi:10.1016/j.molp.2014.10.009. pmid: 25616386
[11]	Zhong R Q, Lee C, McCarthy R L, Reeves C K, Jones E G, Ye Z H. Transcriptional activation of secondary wall biosynthesis by rice and maize NAC and MYB transcription factors[J]. Plant & Cell Physiology, 2011, 52(10):1856-1871.doi:10.1093/pcp/pcr123.
[12]	He J, Liu Y Q, Yuan D Y, Duan M J, Liu Y L, Shen Z J, Yang C Y, Qiu Z Y, Liu D M, Wen P Z, Huang J, Fan D J, Xiao S Z, Xin Y Y, Chen X N, Jiang L, Wang H Y, Yuan L P, Wan J M. An R2R3 MYB transcription factor confers brown planthopper resistance by regulating the phenylalanine ammonia-lyase pathway in rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2020, 117(1):271-277.doi:10.1073/pnas.1902771116. pmid: 31848246
[13]	周帅. 江苏射阳县南粳5718种植表现及优质高产栽培技术[J]. 农业工程技术, 2023, 43(16):64,66.doi:10.16815/j.cnki.11-5436/s.2023.16.038.
	Zhou S. Planting performance and high-yielding cultivation techniques of Nanjing 5718 in Sheyang,Jiangsu[J]. Applied Engineering Technology, 2023, 43(16):64,66.
[14]	徐凯, 黄成裕, 韦军, 霍金兰. 2021年盐都区水稻灌浆结实期灾情调查及成因分析[J]. 大麦与谷类科学, 2022, 39(3):58-63.doi:10.14069/j.cnki.32-1769/s.2022.03.009.
	Xu K, Huang C Y, Wei J, Huo J L. Investigation of the causes of rice production losses in the grain filling stage in Yandu district in 2021[J]. Barley and Cereal Sciences, 2022, 39(3):58-63.
[15]	王夏雯, 王飞飞, 金倩, 陈卫军, 陈春, 于成功. 苏北地区稻虾共作模式粳稻适栽品种(系)的筛选[J]. 北方农业学报, 2021, 49(6):58-64.doi:10.12190/j.issn.2096-1197.2021.06.07.
	Wang X W, Wang F F, Jin Q, Chen W J, Chen C, Yu C G. Selection of suitable Japonica rice varieties(lines)for rice and shrimp co-cropping model in Northern Jiangsu[J]. Inner Mongolia Agricultural Science and Technology, 2021, 49(6):58-64.
[16]	赵凌, 张勇, 朱镇, 陈涛, 赵庆勇, 张亚东, 王才林. 南粳系列品种氮素利用效率初探[J]. 江苏农业学报, 2022, 38(5):1153-1161.doi:10.3969/j.issn.1000-4440.2022.05.001.
	Zhao L, Zhang Y, Zhu Z, Chen T, Zhao Q Y, Zhang Y D, Wang C L. Study on nitrogen use efficiency of Nanjing series japonica rice varieties[J]. Jiangsu Journal of Agricultural Sciences, 2022, 38(5):1153-1161.
[17]	夏敏, 胡群, 梁健, 张洪程, 郭保卫, 曹利强, 陈厚存. 壮秆剂及用量对水稻产量和抗倒伏能力的影响[J]. 作物学报, 2017, 43(2):296-306.doi:10.3724/sp.j.1006.2017.00296.
	Xia M, Hu Q, Liang J, Zhang H C, Guo B W, Cao L Q, Chen H C. Effect of variety and application amount of stalk strengthening agent on yield and lodging resistance in rice[J]. Acta Agronomica Sinica, 2017, 43(2):296-306. doi: 10.3724/SP.J.1006.2017.00296 URL
[18]	陈桂华, 邓化冰, 张桂莲, 唐文帮, 黄璜. 水稻茎秆性状与抗倒性的关系及配合力分析[J]. 中国农业科学, 2016, 49(3):407-417.doi:10.3864/j.issn.0578-1752.2016.03.001.
	Chen G H, Deng H B, Zhang G L, Tang W B, Huang H. The correlation of stem characters and lodging resistance and combining ability analysis in rice[J]. Scientia Agricultura Sinica, 2016, 49(3):407-417.
[19]	张宝, 陈冰炜, 翟胜丞, 潘彪, 魏建强, 勇强. 不同染料标识木栓细胞壁中木栓脂的分布[J]. 西北林学院学报, 2021, 36(2):208-212.doi:10.3969/j.issn.1001-7461.2021.02.31.
	Zhang B, Chen B W, Zhai S C, Pan B, Wei J Q, Yong Q. Distribution of suberin in the phellem cells determined by different dyes[J]. Journal of Northwest Forestry University, 2021, 36(2):208-212.
[20]	卢昆丽, 尹燕枰, 王振林, 李勇, 彭佃亮, 杨卫兵, 崔正勇, 杨东清, 江文文. 施氮期对小麦茎秆木质素合成的影响及其抗倒伏生理机制[J]. 作物学报, 2014, 40(9):1686-1694.doi:10.3724/sp.j.1006.2014.01686.
	Lu K L, Yin Y P, Wang Z L, Li Y, Peng D L, Yang W B, Cui Z Y, Yang D Q, Jiang W W. Effect of nitrogen fertilization timing on lignin synthesis of stem and physiological mechanism of lodging resistance in wheat[J]. Acta Agronomica Sinica, 2014, 40(9):1686-1694. doi: 10.3724/SP.J.1006.2014.01686 URL
[21]	宋安东, 张百良, 吴坤, 黄保. 杂色云芝产木质纤维素酶及对稻草秸秆的降解[J]. 过程工程学报, 2005, 5(4):414-419.doi:10.3321/j.issn:1009-606x.2005.04.013.
	Song A D, Zhang B L, Wu K, Huang B. Production of lignocellulolytic enzymes and rice straw biodegradation by Coriolus versicolor[J]. The Chinese Journal of Process Engineering, 2005, 5(4):414-419.
[22]	Wang T T, Jin Y, Deng L X, Li F, Wang Z Y, Zhu Y Y, Wu Y F, Qu H Y, Zhang S N, Liu Y, Mei H W, Luo L J, Yan M, Gu M, Xu G H. The transcription factor MYB110 regulates plant height,lodging resistance,and grain yield in rice[J]. The Plant Cell, 2024, 36(2):298-323.doi:10.1093/plcell/koad268. URL
[23]	Ma J, Cheng Z J, Chen J, Shen J B, Zhang B C, Ren Y L, Ding Y, Zhou Y H, Zhang H, Zhou K N, Wang J L, Lei C L, Zhang X, Guo X P, Gao H, Bao Y Q, Wan J M. Phosphatidylserine synthase controls cell elongation especially in the uppermost internode in rice by regulation of exocytosis[J]. PLoS One, 2016, 11(4):e0153119.doi:10.1371/journal.pone.0153119.
[24]	Peng S S, Liu Y C, Xu Y C, Zhao J H, Gao P, Liu Q, Yan S Y, Xiao Y H, Zuo S M, Kang H X. Genome-wide association study identifies a plant-height-associated gene OsPG3 in a population of commercial rice varieties[J]. International Journal of Molecular Sciences, 2023, 24(14):11454.doi:10.3390/ijms241411454. URL
[25]	Lu Y Z, Feng Z, Meng Y L, Bian L Y, Xie H, Mysore K S, Liang J S. SLENDER RICE1 and Oryza sativa INDETERMINATE DOMAIN2 regulating OsmiR396 are involved in stem elongation[J]. Plant Physiology, 2020, 182(4):2213-2227.doi:10.1104/pp.19.01008. URL
[26]	Wang F, Cheng Z J, Wang J C, Zhang F, Zhang B C, Luo S, Lei C L, Pan T, Wang Y F, Zhu Y, Wang M, Chen W W, Lin Q B, Zhu S S, Zhou Y H, Zhao Z C, Wang J, Guo X P, Zhang X, Jiang L, Bao Y Q, Ren Y L, Wan J M. Rice STOMATAL CYTOKINESIS DEFECTIVE2 regulates cell expansion by affecting vesicular trafficking in rice[J]. Plant Physiology, 2022, 189(2):567-584.doi:10.1093/plphys/kiac073. pmid: 35234957
[27]	Zhao Z X, Yin X X, Li S, Peng Y T, Yan X L, Chen C, Hassan B, Zhou S X, Pu M, Zhao J H, Hu X H, Li G B, Wang H, Zhang J W, Huang Y Y, Fan J, Li Y, Wang W M. miR167d-ARFs module regulates flower opening and stigma size in rice[J]. Rice, 2022, 15(1):40.doi:10.1186/s12284-022-00587-z.
[28]	Gui J S, Shen J H, Li L G. Functional characterization of evolutionarily divergent 4-coumarate:coenzyme a ligases in rice[J]. Plant Physiology, 2011, 157(2):574-586.doi:10.1104/pp.111.178301. pmid: 21807887
[29]	Kishi-Kaboshi M, Seo S, Takahashi A, Hirochika H. The MAMP-responsive MYB transcription factors MYB30,MYB55 and MYB110 activate the HCAA synthesis pathway and enhance immunity in rice[J]. Plant & Cell Physiology, 2018, 59(5):903-915.doi:10.1093/pcp/pcy062.
[30]	Chen Z H, Teng S Z, Liu D, Chang Y, Zhang L Y, Cui X A, Wu J X, Ai P F, Sun X H, Lu T G, Zhang Z G. RLM1,encoding an R2R3 MYB transcription factor,regulates the development of secondary cell wall in rice[J]. Frontiers in Plant Science, 2022, 13:905111.doi:10.3389/fpls.2022.905111. URL
[31]	Chai M F, Bellizzi M, Wan C X, Cui Z F, Li Y B, Wang G L. The NAC transcription factor OsSWN1 regulates secondary cell wall development in Oryza sativa[J]. Journal of Plant Biology, 2015, 58(1):44-51.doi:10.1007/s12374-014-0400-y. URL
[32]	Chen J, Wang L H, Yang Z Y, Liu H B, Chu C L, Zhang Z Z, Zhang Q L, Li X H, Xiao J H, Wang S P, Yuan M. The rice Raf-like MAPKKK OsILA1 confers broad-spectrum resistance to bacterial blight by suppressing the OsMAPKK4-OsMAPK6 cascade[J]. Journal of Integrative Plant Biology, 2021, 63(10):1815-1842.doi:10.1111/jipb.13150. URL
[33]	Huang D B, Wang S G, Zhang B C, Shangguan K K, Shi Y Y, Zhang D M, Liu X L, Wu K, Xu Z P, Fu X D, Zhou Y H. A gibberellin-mediated DELLA-NAC signaling cascade regulates cellulose synthesis in rice[J]. The Plant Cell, 2015, 27(6):1681-1696.doi:10.1105/tpc.15.00015. pmid: 26002868
[34]	Liu Y F, Wu Q, Qin Z L, Huang J L. Transcription factor OsNAC055 regulates GA-mediated lignin biosynthesis in rice straw[J]. Plant Science, 2022, 325:111455.doi:10.1016/j.plantsci.2022.111455. URL
[35]	Zhang H, Liu M, Yin K Q, Liu H H, Liu J Q, Yan Z. A novel OsHB5-OsAPL-OsMADS27/OsWRKY102 regulatory module regulates grain size in rice[J]. Journal of Plant Physiology, 2024, 295:154210.doi:10.1016/j.jplph.2024.154210. URL
[36]	Zhao Q L, Xin C Y, Liu Q H, Jiang X, Yang J, Ma H. Effects of continuous straw return and fertilization on rice lodging resistance[J]. Chilean Journal of Agricultural Research, 2022, 82(1):146-156.doi:10.4067/s0718-58392022000100146. URL
[37]	许轲, 唐磊, 郭保卫, 张洪程, 霍中洋, 戴其根, 魏海燕. 不同水直播方式水稻植株抗倒特性研究[J]. 华北农学报, 2014, 29(6):226-232.doi:10.7668/hbnxb.2014.06.038.
	Xu K, Tang L, Guo B W, Zhang H C, Huo Z Y, Dai Q G, Wei H Y. Lodging resistance of rice under the different pattern of water direct-seeding cultivation method[J]. Acta Agriculturae Boreali-Sinica, 2014, 29(6):226-232. doi: 10.7668/hbnxb.2014.06.038
[38]	兰彩霞, 郭玉华, 赵鑫闻, 赵东霞, 雒鸫. 杂交粳稻及其亲本抗倒伏性的相关分析[J]. 华北农学报, 2010, 25(2):178-181.doi:10.7668/hbnxb.2010.02.036.
	Lan C X, Guo Y H, Zhao X W, Zhao D X, Luo D. Correlation of lodging resistance traits in Japonica hybrid rice and their parents[J]. Acta Agriculturae Boreali-Sinica, 2010, 25(2):178-181.
[39]	杨惠杰, 房贤涛, 何花榕, 谢祖钦. 福建超级稻品种茎秆结构特征及其与抗倒性和产量的关系[J]. 中国生态农业学报, 2012, 20(7):909-913.doi:10.3724/SP.J.1011.2012.00909.
	Yang H J, Fang X T, He H R, Xie Z Q. Relationship of characteristics of culm construction to lodging resistance and yield of Fujian-bred super-rice cultivars[J]. Chinese Journal of Eco-Agriculture, 2012, 20(7):909-913. doi: 10.3724/SP.J.1011.2012.00909 URL
[40]	Ishida K. Reciprocal regulation of cellulose and lignin biosynthesis by the transcription factor OsTCP19[J]. Journal of Experimental Botany, 2024, 75(1):5-8.doi:10.1093/jxb/erad408. pmid: 38128900
[41]	罗肖郧, 郑兴飞, 彭宣国, 余启芝, 董华林, 殷得所, 王红波, 胡建林, 薛莲, 胡鹏, 徐得泽. 水稻抗倒伏研究:现状、挑战与未来方向[J]. 中国水稻科学, 2026, 40(2):181-195.doi:10.16819/j.1001-7216.2026.240905.
	Luo X Y, Zheng X F, Peng X G, Yu Q Z, Dong H L, Yin D S, Wang H B, Hu J L, Xue L, Hu P, Xu D Z. Research on rice lodging resistance:current status,challenges,and future directions[J]. Chinese Journal of Rice Science, 2026, 40(2):181-195.
[42]	Zhao Y, Wang X P, Gao J, Rehman Rashid M A, Wu H, Hu Q F, Sun X M, Li J J, Zhang H L, Xu P, Qian Q, Chen C, Li Z C, Zhang Z Y. The MYB61-STRONG2 module regulates culm diameter and lodging resistance in rice[J]. Journal of Integrative Plant Biology, 2025, 67(2):243-257.doi:10.1111/jipb.13830. URL
[43]	Wu B, Zhang B C, Dai Y, Zhang L, Shangguan K K, Peng Y G, Zhou Y H, Zhu Z. Brittle culm15 encodes a membrane-associated chitinase-like protein required for cellulose biosynthesis in rice[J]. Plant Physiology, 2012, 159(4):1440-1452.doi:10.1104/pp.112.195529. URL
[44]	Liu G M, Zhang K, Ai J, Deng X J, Hong Y Y, Wang X M. Patatin-related phospholipase A,pPLAⅢα,modulates the longitudinal growth of vegetative tissues and seeds in rice[J]. Journal of Experimental Botany, 2015, 66(21):6945-6955.doi:10.1093/jxb/erv402. URL
[45]	Chen Y, Qi H Y, Yang L J, Xu L, Wang J X, Guo J Z, Zhang L, Tan Y Y, Pan R H, Shu Q Y, Qian Q, Song S Y. The OsbHLH002/OsICE1-OSH1 module orchestrates secondary cell wall formation in rice[J]. Cell Reports, 2023, 42(7):112702.doi:10.1016/j.celrep.2023.112702. URL
[46]	Wu H M, Xie D J, Jia P F, Tang Z S, Shi D Q, Shui G H, Wang G D, Yang W C. Homeostasis of flavonoids and triterpenoids most likely modulates starch metabolism for pollen tube penetration in rice[J]. Plant Biotechnology Journal, 2023, 21(9):1757-1772.doi:10.1111/pbi.14073. URL

选择文件类型/文献管理软件名称

选择包含的内容