外源海藻糖对粳稻品系W1844籽粒灌浆特性及产量形成的影响

doi:10.16819/j.1001-7216.2023.220712

摘要/Abstract

摘要：

【目的】 明确外源海藻糖对大穗型粳稻籽粒灌浆结实的影响。【方法】 以大穗型粳稻品系W1844为试验材料，在孕穗末期叶面喷施浓度分别为0、15、30、50、70、100 mmol/L的海藻糖，分别标记为T₀、T₁₅、T₃₀、T₅₀、T₇₀、T₁₀₀。研究不同浓度海藻糖对大穗型粳稻W1844光合特性、茎鞘物质积累与转运、籽粒灌浆特性和产量的影响。【结果】 1）喷施不同浓度海藻糖均能提高水稻产量，以T₅₀处理的水稻产量最高。与T₀相比，T₅₀处理下的产量平均增加7.71%，强势粒的结实率与千粒重分别提高4.36%和5.92%，弱势粒的结实率与千粒重分别提高11.98%和10.01%。由此可见，海藻糖对弱势粒灌浆结实的改善效果更好。2）海藻糖处理改变了籽粒灌浆特性。与T₀相比，T₅₀处理下强、弱势粒达到最大灌浆速率的时间分别缩短了3.70 d与3.93 d；灌浆前期强、弱势粒的平均灌浆速率分别提高了16.57%、28.03%；灌浆中期强、弱势粒平均灌浆速率分别提高了20.74%、24.54%。此外，T₅₀处理下，强、弱势粒达到最大灌浆速率时的粒重分别比T₀增加了6.74%和7.36%。3）海藻糖处理显著提高水稻剑叶的光合特性。T₅₀处理下水稻叶片的净光合速率、气孔导度和蒸腾速率分别比T₀平均提高了31.08%、42.58%和10.42%。4）海藻糖处理促进了抽穗期茎鞘非结构性碳水化合物(NSC)的积累与转运。与T₀处理相比，T₅₀处理下抽穗期茎鞘NSC含量平均增加11.63%，茎鞘物质转运量平均提升28.52%，且对籽粒贡献率增加18.18%。这可能是由于茎鞘蔗糖转运基因OsSUT2、OsSUT4和OsSUT5上调表达，促进了茎鞘的物质转运。【结论】 外源海藻糖可有效地改善大穗型粳稻的灌浆结实，这可能是由于海藻糖处理能提高剑叶净光合速率，上调蔗糖转运基因的表达，促进茎鞘NSC的积累与转运，从而加快了强、弱势粒的灌浆，提高了其结实率和粒重。在本研究条件下，50 mmol/L海藻糖处理对大穗型水稻W1844籽粒灌浆结实的改善效果最好。

关键词: 大穗型粳稻, 外源海藻糖, 灌浆特性, NSC的积累与转运, 产量形成

Abstract:

【Objective】 It is very important to clarify the effect of exogenous trehalose on grain filling and seed setting of large-panicle japonica rice. 【Method】 The effects of different concentrations of trehalose on photosynthetic characteristics, dry matter accumulation and translocation of stems and sheath, grain filling characteristics and yield of large-panicle japonica rice W1844 were investigated by foliar spraying of various concentrations of trehalose (T₀, T₁₅, T₃₀, T₅₀, T₇₀ and T₁₀₀, 0, 15, 30, 50, 70, 100mmol/L, respectively) at the end of booting stage. 【Result】 1) Trehalose exposure increased grain yield, and the highest yield was obtained under T₅₀ treatment. Compared with T₀, the yield under T₅₀ treatment increased by 7.71% on average, and the yield and thousand grain weight of superior spikelets increased by 4.36% and 5.92%, respectively, while the yield and thousand grain weight of inferior spikelets increased by 11.98% and 10.01%, respectively. Trehalose had a better effect on the improvement of grain filling and seed-setting of the inferior spikelets. 2) Trehalose treatment changed the grain filling characteristics. Compared with T₀, the time to the maximum filling rate of superior and inferior spikelets was shortened by 3.70 and 3.93 d under T₅₀ treatment; the average filling rates of superior and inferior spikelets were increased by 16.57% and 28.03% in the early stage of filling; the average filling rates of superior and inferior spikelets were increased by 20.74% and 24.54% in the middle of grain filling. In addition, compared with T₀, the grain weight of superior and inferior spikelets at maximum filling rate increased by 6.74% and 7.36%, respectively. The net photosynthetic rate, stomatal conductance and transpiration rate of rice leaves under T₅₀ treatment increased by 31.08%, 42.58% and 10.42% on average, respectively as compared with T₀. 4) Trehalose treatment promoted the accumulation and translocation of non-structural carbohydrates (NSC) in stem and sheath during the heading stage. Compared with T₀, the NSC content in stem and sheath at the heading stage increased by 11.63% on average under T₅₀ treatment, and the translocation of dry matter in stems and sheaths increased by 28.52% on average, and the contribution rate to seeds increased by 18.18%. This might be due to the up-regulated expression level of sucrose transporter genes OsSUT2, OsSUT4 and OsSUT5 in stems and sheaths, which promoted dry matter transport in stems and sheaths.【Conclusion】 Exogenous trehalose could effectively improve grain filling and seed setting of large-panicle japonica rice, which might be due to the fact that trehalose treatment could increase the net photosynthetic rate of flag leaf, up-regulate the expression of sucrose transporter genes, and promote the accumulation and translocation of NSC in the stems and sheaths, thus increasing the grain filling rate, seed-setting rate and grain weight of superior and inferior grains. Under the present experimental conditions, 50 mmol/L is the most effective in improving grain filling and seed setting in large-panicle rice W1844.

Key words: large-panicle japonica rice, exogenous trehalose, grain filling characteristics, stem and sheath NSC accumulation and translocation, yield formation

黄亚茹, 徐鹏, 王乐乐, 贺一哲, 王辉, 柯健, 何海兵, 武立权, 尤翠翠. 外源海藻糖对粳稻品系W1844籽粒灌浆特性及产量形成的影响[J]. 中国水稻科学, 2023, 37(4): 379-391.

HUANG Yaru, XU Peng, WANG Lele, HE Yizhe, WANG Hui, KE Jian, HE Haibing, WU Liquan, YOU Cuicui. Effects of Exogenous Trehalose on Grain Filling Characteristics and Yield Formation of japonica Rice Cultivar W1844[J]. Chinese Journal OF Rice Science, 2023, 37(4): 379-391.

图/表 11

参考文献 49

[1]	Khush G S. What it will take to feed 5.0 billion rice consumers in 2030[J]. Plant Molecular Biology, 2005, 59(1): 1-6.
[2]	谢光辉, 杨建昌, 王志琴, 朱庆森. 水稻籽粒灌浆特性及其与籽粒生理活性的关系[J]. 作物学报, 2001, 27(5): 557-565.
	Xie G H, Yang J C, Wang Z Q, Zhu Q S. Grain filling characteristics of rice and their relationships to physiological activities of grains[J]. Acta Agronomica Sinica, 2001, 27(5): 557-565. (in Chinese with English abstract)
[3]	黄农荣, 钟旭华, 王丰, 郑海波. 超级杂交稻结实期根系活力与籽粒灌浆特性研究[J]. 中国农业科学, 2006, 39(9): 1772-1779.
	Huang N R, Zhong X H, Wang F, Zheng H B. Root vigor and grain-filling characteristics in super hybrid rice[J]. Scientia Agricultura Sinica, 2006, 39(9): 1772-1779. (in Chinese with English abstract)
[4]	Tsutomu I, Toshiaki M, Ryu O, Tohru Y. Morphological development of rice caryopses located at the different positions in a panicle from early to middle stage of grain filling[J]. Functional Plant Biology, 2003, 30(11): 1139-1149.
[5]	Mohapatra P K, Patel R, Sahu S K. Time of flowering affects grain quality and spikelet partitioning within the rice panicle[J]. Australian Journal of Plant Physiology, 1993, 20(2): 231-241.
[6]	Naik P K, Mohapatra P K. Ethylene inhibitors promote male gametophyte survival in rice[J]. Plant Growth Regulation, 1999, 28(1): 29-39.
[7]	You C C, Zhu H L, Xu B B, Huang W X, Wang S H, Ding Y F, Liu Z H, Li G H, Chen L, Ding C Q, Tang S. Effect of removing superior spikelets on grain filling of inferior spikelets in rice[J]. Frontiers Plant Science, 2016, 7: 1161.
[8]	李哲, 姜沣益, 代梦雪, 李宇星, 张文静, 马尚宇, 黄正来, 樊永惠. 外源海藻糖对高温胁迫小麦干物质积累和籽粒灌浆的影响[J]. 麦类作物学报, 2022, 42(5): 614-622.
	Li Z, Jiang F Y, Dai M X, Li Y X, Zhang W J, Ma S Y, Huang Z L, Fan Y H. Effect of exogenous trehalose on dry matter accumulation and grain filling of wheat under heat stress[J]. Journal of Triticeae Crops, 2022, 42(5): 614-622. (in Chinese with English abstract)
[9]	Liang Z M, Luo J, Wei B, Liao Y C, Liu Y. Trehalose can alleviate decreases in grain number per spike caused by low-temperature stress at the booting stage by promoting floret fertility in wheat[J]. Journal of Agronomy and Crop Science, 2021, 207(4): 717-732.
[10]	Luo Y, Xie Y, He D, Wang W, Yuan S. Exogenous trehalose protects photosystem II by promoting cyclic electron flow under heat and drought stresses in winter wheat[J]. Plant Biology, 2021, 23(5): 770-776.
[11]	Luo Y, Gao Y M, Wang W, Zou C J. Application of trehalose ameliorates heat stress and promotes recovery of winter wheat seedlings[J]. Biologia Plantarum, 2014, 58(2): 395-398.
[12]	李金花, 张春艳, 刘浩, 李永春. 海藻糖的特性及其在植物抗逆性中的应用[J]. 江西农业学报, 2011, 23(6): 25-27.
	Li J H, Zhang C Y, Liu H, Li Y C. Characteristics of trehalose and its application in improving stress tolerance of plants[J]. Acta Agriculturae Jiangxi, 2011, 23(6): 25-27. (in Chinese with English abstract)
[13]	Wingler A, Fritzius T, Wiemken A, Boller T, Aeschbacher R A. Trehalose induces the ADP-glucose pyrophosphorylase gene, ApL3, and starch synthesis in Arabidopsis[J]. Plant Physiology, 2000, 124(1): 105-114.
[14]	Kolbe A, Tiessen A, Schluepmann H, Paul M, Ulrich S, Geigenberger P. Trehalose 6-phosphate regulates starch synthesis via posttranslational redox activation of ADP-glucose pyrophosphorylase[J]. Proceedings of the National Academy of Sciences, 2005, 102(31): 11118-11123.
[15]	Iturriaga G, Suárez R, Nova-Franco B. Trehalose metabolism: From osmoprotection to signaling[J]. International Journal of Molecular Sciences, 2009, 10(9): 3793-3810.
[16]	Yadav U P, Ivakov A, Feil R, Duan G Y, Walther D, Giavalisco P, Piques M, Carillo P, Hubberten H, Stitt M, Lunn J E. The sucrose-trehalose 6-phosphate (Tre6P) nexus: Specificity and mechanisms of sucrose signalling by Tre6P[J]. Journal of Experimental Botany, 2014, 65(4): 1051-1068.
[17]	Richards F J. A flexible growth function for empirical use[J]. Journal of Experimental Botany, 1959, 10(2): 290-301.
[18]	Zhang C X, Fu G F, Yang X Q, Yang Y J, Zhao X, Chen T T, Zhang X F, Jin Q Y, Tao L X. Heat stress effects are stronger on spikelets than on flag leaves in rice due to differences in dissipation capacity[J]. Journal of Agronomy and Crop Science, 2016, 202(5): 394-408.
[19]	Yoshida S. Physiological aspects of grain yield[J]. Annual Review of Plant Physiology, 1972, 23(1): 437-464.
[20]	Yang J C, Zhang J H. Grain filling of cereals under soil drying[J]. The New Phytologist, 2006, 169(2): 223-236.
[21]	Aoki N, Hirose T, Scofield G N, Whitfeld P R, Furbank R T. The sucrose transporter gene family in rice[J]. Plant and Cell Physiology, 2003, 44(3): 223-232.
[22]	Mohapatra P, Patel R, Sahu S. Time of flowering affects grain quality and spikelet partitioning within the rice panicle[J]. Functional Plant Biology, 1993, 20(2): 231-241.
[23]	Ishimaru T, Matsuda T, Ohsugi R, Yamagishi T. Morphological development of rice caryopses located at the different positions in a panicle from early to middle stage of grain filling[J]. Functional Plant Biology, 2003, 30(11): 1139-1149.
[24]	Ao H J, Wang S H, Zou Y B. Study on yield stability and dry matter characteristics of super hybrid rice[J]. Scientia Agricultura Sinica, 2008, 41(7): 1927-1936.
[25]	Yang J C, Zhang J H. Grain-filling problem in ‘super’ rice[J]. Journal of Experimental Botany, 2010, 61(1): 1-5.
[26]	Khater M A, Dawood M G, Sadak M S, Shalaby M A, El-Awadi M E, El-Din K G. Enhancement the performance of cowpea plants grown under drought conditions via trehalose application[J]. Middle East Journal of Agriculture Research, 2018, 7(3): 782-800.
[27]	Luo Y, Xie Y, Li W, Wei M, Dai T, Li Z, Wang B. Physiological and transcriptomic analyses reveal exogenous trehalose is involved in the responses of wheat roots to high temperature stress[J]. Plants, 2021, 10(12): 2644.
[28]	Luo Y, Wang Y, Xie Y, Gao Y, Li W, Lang S. Transcriptomic and metabolomic analyses of the effects of exogenous trehalose on heat tolerance in wheat[J]. International Journal of Molecular Sciences, 2022, 23(9): 5194.
[29]	Zhao D Q, Li T T, Hao Z J, Cheng M L, Tao J. Exogenous trehalose confers high temperature stress tolerance to herbaceous peony by enhancing antioxidant systems, activating photosynthesis, and protecting cell structure[J]. Cell Stress and Chaperones, 2019, 24(1): 247-257.
[30]	Chen L, Deng Y, Zhu H L, Hu Y X, Jiang Z G, Tang S, Wang S H, Ding Y F. The initiation of inferior grain filling is affected by sugar translocation efficiency in large panicle rice[J]. Rice, 2019, 12(1): 1-13.
[31]	Islam M N, Masud A A C, Alam M M, Islam M N, Rahman M L, Hasanuzzaman M. Osmolyte-induced water deficit stress mitigation during panicle initiation stage in transplanted rice (Oryza sativa L.)[J]. Plant Science Today, 2022, 9(1): 9-20.
[32]	李敏, 罗德强, 江学海, 蒋明金, 李树杏, 姬广梅, 李立江, 周维佳. 高产氮高效型灿稻品种的籽粒灌浆特性[J]. 中国农业科技导报, 2020, 22(9): 22-30.
	Li M, Luo D Q, Jiang X H, Jiang M J, Li S X, Ji G M, Li L J, Zhou W J. Grain filling characteristics of the ricecultivar with high yield and high nitrogen use efficieney[J]. Journal of Agricultural Science and Techrology, 2020, 22(9): 22-30. (in Chinese with English abstract)
[33]	唐益平, 李向峰, 王辉, 胡王琴, 任楚婷, 黄亚茹, 徐鹏, 尤翠翠, 柯健, 何海兵, 武立权. 茎鞘非结构性碳水化合物对大穗型粳稻强、弱势粒灌浆与品质的影响[J]. 华北农学报, 2021, 36(5): 107-117.
	Tang Y P, Li X F, Wang H, Hu W Q, Ren C T, Huang Y R, Xu P, You C C, Ke J, He H B, Wu L Q. Effect of non-structural carbohydrate in stem and sheath on grain filling and quality of superior and inferior spikelets in large-panicle japonica rice[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(5): 107-117. (in Chinese with English abstract)
[34]	顾俊荣, 韩立宇, 董明辉, 陈培峰, 乔中英. 不同穗型粳稻干物质运转与颖花形成及籽粒灌浆结实的差异研究[J]. 扬州大学学报: 农业与生命科学版, 2017, 38(4): 68-73+88.
	Gu J R, Han L Y, Dong M H, Chen P F, Qiao Z Y. Studies on the difference of dry matter accumulation and transportation, spikelets formation and the grain filling of Japonica rice varieties with different panicle types[J]. Journal of Yangzhou University: Agricultural and Life Science Edition, 2017, 38(4): 68-73+88. (in Chinese with English abstract)
[35]	朱庆森, 曹显祖, 骆亦其. 水稻籽粒灌浆的生长分析[J]. 作物学报, 1988, 14(3): 182-193.
	Zhu Q S, Cao X Z, Luo Y Q. Growth analysis on process of grain filling in rice[J]. Acta Agronomica Sinica, 1988, 14(3): 182-193. (in Chinese with English abstract)
[36]	Teng Z, Yu H, Wang G, Meng S, Liu B, Yi Y, Chen Y, Zheng Q, Liu L, Yang J, Duan M, Zhang J, Ye N. Synergistic interaction between ABA and IAA due to moderate soil drying promotes grain filling of inferior spikelets in rice[J]. The Plant Journal, 2022, 109(6): 1457-1472.
[37]	Okamura M, Arai-Sanoh Y, Yoshida H, Mukouyama T, Adachia S, Yabe S, Nakagawa H, Tsutsumi K, Taniguchi Y, Kobayashia N, Kondo M. Characterization of high-yielding rice cultivars with different grain-filling properties to clarify limiting factors for improving grain yield[J]. Field Crops Research, 2018, 219: 139-147.
[38]	Cock J H, Yoshida S. Accumulation of ¹⁴C-labelled carbohydrate before flowering and its subsequent redistribution and respiration in the rice plant[J]. Japanese Journal of Crop Science, 1972, 41(2): 226-234.
[39]	Tsukaguchi T, Horie T, Ohnishi M. Filling percentage of rice spikelets as affected by availability of non-structural carbohydrates at the initial phase of grain filling[J]. Japanese Journal of Crop Science, 1996, 65(3): 445-452.
[40]	Liang J S, Cao X Z, Zhang H Y, Song P, Zhu Q S. The changes and affecting factors of stem-sheath reserve contents of rice during grain filling[J]. Chinese Journal of Rice Science. 1994, 8(3): 151-156.
[41]	Xie G H, Yang J C, Wang Z Q. Grain filling characteristics of rice and their relationships to physiological activities of grains[J]. Acta Agronomica Sinica. 2001, 27(5): 557-565.
[42]	Murty P S S, Murty K S. Spikelet sterility in relation to nitrogen and carbohydrate contents in rice[J]. Indian J Plant Physiology, 1982, 25: 40-48.
[43]	Venkateswarlu B, Visperas R M. Source-sink relationships in crop plants: A review[J]. IRRI Research Paper Series, 1987, 125:1-19.
[44]	Garg A K, Kim J K, Owens T G, Ranwala A P, Choi Y D, Kochian L V, Wu R J. Trehalose accumulation in rice plants confers high tolerance levels to different abiotic stresses[J]. Proceedings of the National Academy of Sciences, 2002, 99(25): 15898-15903.
[45]	Shahbaz M, Abid A, Masood A, Waraich E A. Foliar-applied trehalose modulates growth, mineral nutrition, photosynthetic ability, and oxidative defense system of rice (Oryza sativa L.) under saline stress[J]. Journal of Plant Nutrition, 2017, 40(4): 584-599.
[46]	Fu J, Huang Z, Wang Z, Yang J, Zhang J. Pre-anthesis non-structural carbohydrate reserve in the stem enhances the sink strength of inferior spikelets during grain filling of rice[J]. Field Crops Research, 2011, 123(2): 170-182.
[47]	Liang X G, Gao Z, Shen S, Panl M J, Zhang L, Zhao X, Lin S, Wu G, Chen X M, Zhou S L. Differential ear growth of two maize varieties to shading in the field environment: effects on whole plant carbon allocation and sugar starvation response[J]. Journal of Plant Physiology, 2020, 251: 153194.
[48]	Eom J S, Cho J I, Reinders A, Lee S W, Yoo Y, Tuan P Q, Choi S B, Bang G, Park Y I, Cho M H, Bhoo S H, An G, Hahn T R, Ward J M, Jeon J S. Impaired function of the tonoplast-localized sucrose transporter in rice, OsSUT2, limits the transport of vacuolar reserve sucrose and affects plant growth[J]. Plant Physiology, 2011, 157(1): 109-119.
[49]	Aoki N, Hirose T, Scofield G N, Whitfeld P R, Furbank R T. The sucrose transporter gene family in rice[J]. Plant and Cell Physiology, 2003, 44(3): 223-232.

基因名称 Gene name	登录号 Accession	上游引物 Up-primer(5’-3’)	上游引物 Down-primer (5’-3’)
OsSUT2	Os12g0641400	AACCTCAAGTCGGCCTTTCT	CTGTGCTAGGGTGATCTGCT
OsSUT3	Os10g0404500	ACATCCAGCCTTGGAAGACG	CTTGCAGTCCTCAGTCGTGT
OsSUT4	Os02g0827200	GACAAGGTCTGGCAACAGGA	CCTCCCCCAAAGAGAGCATC
OsSUT5	Os02g0576600	CGTTCGCTGTGTTGTCGTTA	GCCAATAAACAAGGCAGCCA

基因名称 Gene name	登录号 Accession	上游引物 Up-primer(5’-3’)	上游引物 Down-primer (5’-3’)
OsSUT2	Os12g0641400	AACCTCAAGTCGGCCTTTCT	CTGTGCTAGGGTGATCTGCT
OsSUT3	Os10g0404500	ACATCCAGCCTTGGAAGACG	CTTGCAGTCCTCAGTCGTGT
OsSUT4	Os02g0827200	GACAAGGTCTGGCAACAGGA	CCTCCCCCAAAGAGAGCATC
OsSUT5	Os02g0576600	CGTTCGCTGTGTTGTCGTTA	GCCAATAAACAAGGCAGCCA

年份 Year	处理 Treatment	穗粒数 Grain No. per panicle	结实率Seed setting rate/%		千粒重1000-grain weight/g		产量 Yield/(kg·667m⁻²)
年份 Year	处理 Treatment	穗粒数 Grain No. per panicle	强势粒 SS	弱势粒 IS	强势粒 SS	弱势粒 IS	产量 Yield/(kg·667m⁻²)
2021	T₀	254.33±3.84 b	86.94±1.31 b	74.83±0.74 d	26.94±0.17 b	23.30±0.10 d	682.84±18.27 c
	T₁₅	267.67±7.51 a	87.71±0.60 b	79.44±0.57 c	27.44±0.13 ab	23.86±0.16 c	704.50±9.99 bc
	T₃₀	271.00±4.04 a	90.86±0.45 a	82.23±0.55 ab	27.83±0.10 ab	24.43±0.20 b	727.87±10.60 ab
	T₅₀	267.67±0.88 a	91.47±0.89 a	83.60±1.06 a	28.30±0.52 a	25.12±0.18 a	741.23±1.34 a
	T₇₀	260.00±3.05 a	89.62±0.91 ab	80.64±0.56 bc	27.92±0.25 ab	24.25±0.07 bc	706.43±6.84 bc
	T₁₀₀	266.67±2.67 a	89.96±1.37 ab	79.79±0.57 bc	27.78±0.14 ab	23.79±0.21 cd	706.97±7.67 bc
2022	T₀	258.45±1.24 b	85.83±0.11 b	72.13±0.37 b	25.44±0.13 c	21.87±0.26 c	641.49±4.76 b
	T₃₀	264.57±0.98 a	88.07±0.27 ab	79.13±0.74 a	26.47±0.09 b	23.41±0.31 b	676.06±5.25 a
	T₅₀	268.19±1.58 a	88.77±0.23 a	80.93±0.39 a	27.13±0.01 a	24.54±0.16 a	685.48±1.77 a

年份 Year	处理 Treatment	穗粒数 Grain No. per panicle	结实率Seed setting rate/%		千粒重1000-grain weight/g		产量 Yield/(kg·667m⁻²)
年份 Year	处理 Treatment	穗粒数 Grain No. per panicle	强势粒 SS	弱势粒 IS	强势粒 SS	弱势粒 IS	产量 Yield/(kg·667m⁻²)
2021	T₀	254.33±3.84 b	86.94±1.31 b	74.83±0.74 d	26.94±0.17 b	23.30±0.10 d	682.84±18.27 c
	T₁₅	267.67±7.51 a	87.71±0.60 b	79.44±0.57 c	27.44±0.13 ab	23.86±0.16 c	704.50±9.99 bc
	T₃₀	271.00±4.04 a	90.86±0.45 a	82.23±0.55 ab	27.83±0.10 ab	24.43±0.20 b	727.87±10.60 ab
	T₅₀	267.67±0.88 a	91.47±0.89 a	83.60±1.06 a	28.30±0.52 a	25.12±0.18 a	741.23±1.34 a
	T₇₀	260.00±3.05 a	89.62±0.91 ab	80.64±0.56 bc	27.92±0.25 ab	24.25±0.07 bc	706.43±6.84 bc
	T₁₀₀	266.67±2.67 a	89.96±1.37 ab	79.79±0.57 bc	27.78±0.14 ab	23.79±0.21 cd	706.97±7.67 bc
2022	T₀	258.45±1.24 b	85.83±0.11 b	72.13±0.37 b	25.44±0.13 c	21.87±0.26 c	641.49±4.76 b
	T₃₀	264.57±0.98 a	88.07±0.27 ab	79.13±0.74 a	26.47±0.09 b	23.41±0.31 b	676.06±5.25 a
	T₅₀	268.19±1.58 a	88.77±0.23 a	80.93±0.39 a	27.13±0.01 a	24.54±0.16 a	685.48±1.77 a

年份 Year	处理 Treatment	起始生长势 R₀		最大灌浆速率 GR_max /(mg·grain^-1 d^-1)		达到最大灌浆速率的时间 T_max/d		平均灌浆速率 GR_mean /(mg·grain^-1 d^-1)		最大灌浆速率时粒重 W_max/(mg·grain^-1)		活跃灌浆期 D/d
年份 Year	处理 Treatment	SS	IS	SS	IS	SS	IS	SS	IS	SS	IS	SS	IS
2021	T₀	0.1497	0.0792	1.0842	0.6331	16.96	26.68	0.7208	0.4113	13.02	12.00	37.55	50.59
	T₁₅	0.1783	0.0772	1.1045	0.6595	15.93	25.95	0.7391	0.4259	13.44	12.36	37.46	49.04
	T₃₀	0.2304	0.0890	1.1983	0.7005	14.43	24.28	0.8063	0.4588	13.33	12.59	35.80	49.43
	T₅₀	0.2442	0.0929	1.3176	0.7818	13.94	23.34	0.8864	0.5112	13.79	13.28	33.66	46.41
	T₇₀	0.2254	0.0804	1.1377	0.6794	14.75	25.45	0.7657	0.4411	13.06	12.64	37.05	49.58
	T₁₀₀	0.2098	0.0698	1.1088	0.6514	15.33	27.42	0.7454	0.4174	13.08	12.95	37.64	50.88
2022	T₀	0.1183	0.0842	1.0556	0.6931	18.54	26.95	0.6901	0.4517	12.97	12.34	36.31	49.19
	T₃₀	0.1439	0.1001	1.1450	0.7670	16.58	24.34	0.7580	0.5066	13.52	12.53	35.92	46.63
	T₅₀	0.2335	0.1085	1.2818	0.8326	14.17	22.43	0.8612	0.5494	13.95	12.84	33.08	44.99