Transcription Factor MYCs Regulating Terpenoids in Tomato Trichomes: Research Progress on Molecular Mechanism

YU Lan, WANG Haoran, ZHANG Ying, XING Hongyun, DING Qi, ZHAO Baozhen, CUI Na

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Chinese Agricultural Science Bulletin ›› 2022, Vol. 38 ›› Issue (6) : 87-93. DOI: 10.11924/j.issn.1000-6850.casb2021-0346

Transcription Factor MYCs Regulating Terpenoids in Tomato Trichomes: Research Progress on Molecular Mechanism

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Abstract

Tomato is an important vegetable widely grown worldwide, and it is susceptible to pests and diseases, resulting in the reduction of yield. Trichome is a crucial barrier covering the epidermis of tomato, which can enhance the adaptability of tomato to biotic and abiotic stresses. In order to further study the functional mechanism of trichomes in tomato stress resistance, the types of tomato trichome inclusions and molecular regulation mechanism were summarized in this review, and the molecular mechanism of the terpenoids’ synthesis regulation in tomato trichomes by the core transcription factor MYC1 and MYC2 in JA signaling pathway was concluded. The problems existing in the current trichome research were analyzed, and the future research direction of trichome was discussed.

Key words

JA signaling / regulating mechanism / tomato trichome / terpenoids / SlMYC1/2 / transcription factor

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YU Lan , WANG Haoran , ZHANG Ying , XING Hongyun , DING Qi , ZHAO Baozhen , CUI Na. Transcription Factor MYCs Regulating Terpenoids in Tomato Trichomes: Research Progress on Molecular Mechanism. Chinese Agricultural Science Bulletin. 2022, 38(6): 87-93 https://doi.org/10.11924/j.issn.1000-6850.casb2021-0346

0 引言

水稻作为全球农业生产的重要组成部分，其产量和品质直接关系到亿万人的饮食健康。优质食味粳稻[1]，因其独特的食味和营养价值，受到了广大消费者的喜爱。然而，随着农业生产环境的变化和消费者对食品品质要求的提高，如何在保证产量的前提下进一步优化稻米品质，成为了农业科研和生产的重要课题。
早在20世纪80年代，学界就已经认识到稻米品质的形成并非仅由遗传因素决定，农业生产管理也对其有深刻影响[2]，其中轮作模式[3]与氮肥管理[4]被认为是调控稻米品质的重要生产环节。江苏地区传统的稻-麦（油）轮作模式，虽然在一定程度上稳定了水稻产量，但长期的单一轮作可能会导致土壤生物活性降低和病虫害风险增加，进而影响稻米品质[5]。为此，科研人员和农业生产者积极探索新的轮作模式，如稻-紫云英[6]和稻-羊肚菌[7]，旨在通过多样化的轮作模式来改善土壤环境并优化稻米品质。吴玉红等[8]在研究中发现，与传统稻-麦或稻-休耕轮作模式相比，稻-紫云英轮作显著提高了稻米的蛋白质含量，降低了直链淀粉含量，改善了胶稠度。氮肥是农业生产中使用最广泛的营养肥料，为了追求高产，生产上普遍存在过度施用氮肥的问题[9]。氮肥的过量施用不仅会对水稻产量造成逆效应，而且会影响稻米品质[10]。有研究表明，随着施氮量的增加，加工品质和蛋白质含量增加，外观品质、直链淀粉含量、胶稠度以及蒸煮品质下降[11]。殷春渊等[12]也提到了施氮量与稻米品质间的非线性关系，适量减少氮肥施用量，可以提高稻米的精米率，降低垩白度。
近年来，随着农业生态学和土壤学的不断研究，揭示了多元化轮作模式与氮肥管理的潜在协同效应，为优化稻米品质提供了新的视角[13-14]。颜志雷等[15]研究发现，稻-紫云英长期轮作模式下，化肥配施紫云英可以显著提高微生物碳量和微生物氮量。林郸等[16]进一步证实，不同的轮作模式和氮肥策略对稻米品质有着显著的影响，结合适宜氮肥用量和氮肥运筹策略，可以显著改善稻米的蒸煮食味与营养品质。尽管其他轮作模式如稻-羊肚菌在实践中也得到应用，但其对稻米品质的潜在影响尚缺乏明确的学术报道。而且，如何在实际生产中整合不同的轮作模式和氮肥策略，以达到最佳的稻米品质，仍是一个未解决的问题。本研究旨在探讨不同轮作模式与氮肥运筹对优质食味粳稻稻米品质的调控效应，为农业生产实践提供科学的指导和建议。

1 材料与方法

1.1 试验地点及供试品种

试验于2021年和2022年在江苏省常熟市古里镇江苏省现代农业（稻麦）产业技术体系坞坵基地进行，试验田土壤类型为黏土。2022年土壤耕作层有机质质量分数32.8 g/kg，速效氮、磷和钾质量分数分别为135.2、8.1、121.6 mg/kg，供试品种为优质食味粳稻品种‘宁香粳9号’（早熟晚粳）。

1.2 试验设计

大田试验采用裂区设计，不同轮作模式为主区，氮肥用量为裂区，氮肥比例为裂裂区。试验设置3种轮作模式：稻-麦模式(A1)、稻-紫云英模式(A2)、稻-羊肚菌模式(A3)；氮肥用量设置2种氮肥水平，分别为减氮10%即纯氮243 kg/hm2(B1)和常规施用量纯氮270 kg/hm2(B2)；氮肥比例按照基蘖肥:穗肥设置5:5(C1)、6:4(C2)、7:3(C3)和8:2(C4)4种处理。
供试氮肥为尿素（含N46%），磷肥为过磷酸钙（P2O5含量12%），钾肥为氯化钾（K2O含量60%）。各处理组合磷肥和钾肥用量相同，分别为P2O5 75 kg/hm2和K2O 150 kg/hm2，磷肥作一次性基施，钾肥作基肥与穗肥2次施用。水分、病虫害等田间管理措施保持与大田常规生产一致。

1.3 测定项目与方法

每个处理组合收获的稻谷干燥后储存3个月以确保籽粒品质稳定性，参照中华人民共和国国家标准GB/T 17891—2017《优质稻谷》，测定糙米率、精米率、整精米率、垩白粒率、垩白大小、垩白度、直链淀粉含量、胶稠度和蛋白质含量等关键指标。
稻米淀粉RVA谱特征值分析釆用澳大利亚波通RVA-4500快速粘度分析仪，参照GB/T 24852—2010《大米及米粉糊化特性测定 快速粘度仪法》测定，用TCW(Thermal Cycle for Windows)配套软件进行分析。

1.4 数据统计

相关数据使用Excel 2016进行录入、整理及初步计算。采用SPSS 21.0对数据进行方差分析，多重比较采用SSR新复极差法。2 a数据结果规律和趋势相似，为便于分析和讨论，本研究主要基于2022年数据进行深入分析和解释。

2 结果与分析

2.1 稻米品质分析

2.1.1 对加工品质的影响

对于稻米的加工品质（糙米率、精米率、整精米率）在不同轮作模式和氮肥运筹下的差异，从表1的方差分析结果来看，轮作模式(A)、氮肥用量(B)和氮肥比例(C)均对加工品质指标产生了显著影响，并且三者之间存在着显著的交互作用。具体来说，A、B和C对糙米率、精米率和整精米率的影响均为极显著，说明这些因素在控制稻米的加工品质上起到了关键作用。
表1 不同因素及其互作对加工品质影响的方差分析
因素 糙米率 精米率 整精米率
A ** ** **
B ** ** **
C ** ** **
A×B ** ** **
A×C ** ** **
B×C ** ** **
A×B×C ** ** **
注：*和**分别表示在5%和1%水平上差异显著，ns表示差异不显著。下同。
从表2的数据可以看出，在所有处理组合中，A3B1C2组合达到了最高的糙米率(85.17%)、精米率(80.38%)和整精米率(72.60%)，表明稻-羊肚菌轮作模式、减氮10%的施肥量以及6:4的氮肥运筹策略对于优化稻米的加工品质最为有利。与此相对，A1B2C4组合的糙米率(69.69%)、精米率(58.14%)和整精米率(54.04%)相对较低，这可能说明稻-麦轮作模式、常规氮肥用量和8:2的氮肥运筹策略对稻米的加工品质不甚有利。进一步深入分析数据可以发现，当轮作模式为稻-羊肚菌模式时，糙米率、精米率和整精米率总体高于其他轮作模式；而在氮肥用量方面，减氮10%的施肥策略在多数组合中都能带来较高的加工品质；至于氮肥比例，6:4的策略可能对稻米的加工品质最为有益。综上所述，为了获得最佳的稻米加工品质，建议采用稻-羊肚菌的轮作模式、适量减少氮肥用量，并采取6:4的氮肥运筹策略。
表2 不同处理组合对加工品质的影响
处理组合 糙米率/% 精米率/% 整精米率/%
A1B1C1 72.21 n 65.24 m 51.51 o
A1B1C2 73.19 m 65.72 l 52.97 n
A1B1C3 76.70 fg 70.24 h 63.13 h
A1B1C4 74.89 jk 68.41 j 61.57 i
A1B2C1 74.32 kl 65.80 l 52.76 n
A1B2C2 75.97 hi 67.57 k 56.86 l
A1B2C3 79.85 c 72.28 f 65.60 ef
A1B2C4 69.69 o 58.14 o 54.04 m
A2B1C1 76.55 fgh 68.64 j 60.94 j
A2B1C2 77.04 ef 73.29 e 68.22 c
A2B1C3 76.44 fgh 70.16 h 65.44 f
A2B1C4 75.88 hi 69.90 hi 64.38 g
A2B2C1 67.50 p 63.30 n 59.92 k
A2B2C2 76.12 ghi 71.29 g 66.03 ef
A2B2C3 75.51 ij 69.62 i 65.83 ef
A2B2C4 74.02 l 64.96 m 61.42 ij
A3B1C1 78.44 d 72.18 f 63.02 h
A3B1C2 85.17 a 80.38 a 72.60 a
A3B1C3 80.08 c 78.52 b 69.28 b
A3B1C4 79.96 c 75.70 d 68.45 c
A3B2C1 77.67 e 72.14 f 66.21 e
A3B2C2 83.38 b 78.12 c 68.25 c
A3B2C3 77.75 de 72.46 f 67.11 d
A3B2C4 76.93 f 70.04 h 65.95 ef
注：同列中不同小写字母表示5%水平差异显著。下同。

2.1.2 对外观品质的影响

从方差分析（表3）来看，轮作模式(A)和氮肥比例(C)对垩白粒率、垩白大小和垩白度的影响均非常显著。相比之下，氮肥用量(B)在垩白粒率上未显示出显著差异，但在垩白大小和垩白度上表现出显著或极显著的差异。在因素之间的互作效应方面，轮作模式(A)与氮肥用量(B)在垩白大小上的影响极为显著。同时，轮作模式(A)与氮肥比例(C)在垩白大小和垩白度上的互作效应也显示出极显著的差异。而当考虑三因素的组合互作(A×B×C)时，对垩白粒率、垩白大小和垩白度的影响均未表现出显著性差异。
表3 不同因素及其互作对外观品质影响的方差分析
因素 垩白粒率 垩白大小 垩白度
A ** ** **
B ns ** *
C ** ** **
A×B ns ** ns
A×C ns ** **
B×C ns ns ns
A×B×C ns ns ns
不同的处理组合对外观品质各指标展现了不同的变化趋势（表4）。整体来看，A1B2C4处理组合下，垩白粒率、垩白大小和垩白度的数值均较高，分别为14.00%、26.33%和3.69%，暗示其外观品质较差。而A3B1C2处理组合下，三者的数值均是最低，分别为7.33%、18.17%和1.33%，代表了较好的外观品质。
表4 不同处理组合对外观品质的影响
处理组合 垩白粒率/% 垩白大小/% 垩白度/%
A1B1C1 13.33 abc 21.83 defg 2.92 bcd
A1B1C2 13.00 abc 20.33 ghijkl 2.64 cdef
A1B1C3 12.00 bcdef 20.17 hijkl 2.42 defg
A1B1C4 14.67 a 23.00 cd 3.37 ab
A1B2C1 12.33 abcde 25.83 a 3.17 abc
A1B2C2 12.00 bcdef 23.17 cd 2.78 cde
A1B2C3 11.67 bcdefg 21.67 defgh 2.52 def
A1B2C4 14.00 ab 26.33 a 3.69 a
A2B1C1 11.00 cdefg 20.50 fghijk 2.26 efg
A2B1C2 9.67 fghi 19.50 jklm 1.88 ghi
A2B1C3 12.00 bcdef 21.83 defg 2.62 cdef
A2B1C4 12.67 abcd 24.83 ab 3.15 abc
A2B2C1 12.00 bcdef 21.67 defgh 2.60 cdef
A2B2C2 11.00 cdefg 21.33 efghi 2.33 efg
A2B2C3 12.33 abcde 22.33 cde 2.77 cde
A2B2C4 13.33 abc 23.83 bc 3.18 abc
A3B1C1 8.33 hi 19.00 klm 1.58 hi
A3B1C2 7.33 i 18.17 m 1.33 i
A3B1C3 10.00 efgh 21.00 efghij 2.10 fgh
A3B1C4 10.33 defgh 22.50 cde 2.33 efg
A3B2C1 9.33 ghi 19.83 ijkl 1.85 ghi
A3B2C2 8.00 hi 18.83 lm 1.51 i
A3B2C3 11.33 cdefg 22.00 def 2.50 def
A3B2C4 12.00 bcdef 23.17 cd 2.78 cde
稻-羊肚菌模式(A3)的外观品质明显优于稻-麦模式(A1)和稻-紫云英模式(A2)，这可能是由于羊肚菌对土壤中的有机质和微生物的积累和改良，提高了稻米的外观品质。氮肥用量在B2（常规施用量）的处理下，特别是在与A1结合时，垩白粒率和垩白大小都呈现出较高的数值，减氮10%(B1)的处理组合的外观品质一般优于常规施用量(B2)，说明适当的减氮可以提高稻米的外观品质。在氮肥比例方面，在某些轮作模式和氮肥用量的组合中，如A1B2，随着基蘖肥:穗肥比例从5:5增至8:2，垩白度从3.17%增加到3.69%，这意味着增加基蘖肥的比例可能会导致外观品质下降。
可见，在提高稻米外观品质方面，稻-羊肚菌模式配合减氮10%的氮肥并采用6:4的氮肥比例(A3B1C2)表现出最佳的外观品质表现，而稻-麦模式配合常规施用量的氮肥并采取8:2的氮肥比例(A1B2C4)则展现出较差的外观品质。

2.1.3 对蒸煮食味与营养品质的影响

由表5可见，轮作模式(A)对胶稠度和蛋白质含量的影响差异极显著，而对直链淀粉含量的影响差异显著；氮肥用量(B)对所有3项指标均有显著或极显著的影响；氮肥比例(C)对胶稠度和蛋白质含量的影响极显著，但对直链淀粉含量的影响不显著。轮作模式和氮肥用量(A×B)的交互作用对直链淀粉含量和蛋白质含量的影响达到极显著水平，但对胶稠度的影响不显著；轮作模式和氮肥比例(A×C)、氮肥用量和氮肥比例(B×C)的交互作用均对蛋白质含量影响极显著；轮作模式、氮肥用量和氮肥比例的三者互作(A×B×C)对胶稠度和蛋白质含量的影响均达到显著水平，但对直链淀粉含量的影响不显著。
表5 不同因素及其互作对蒸煮食味与营养品质影响的方差分析
处理 直链淀粉含量 胶稠度 蛋白质含量
A * ** **
B ** * **
C ns ** **
A×B ** ns **
A×C ns ns **
B×C ns ns **
A×B×C ns ** **
通过对不同处理组合下的数据分析（表6），发现在轮作模式中，稻-紫云英模式(A2)在B1和C2处理组合下表现出较高的直链淀粉含量和胶稠度；而对于蛋白质含量，A1B2C3的组合值最高，达到6.91 g/100g。综合考虑所有处理组合，A2B1C2组合在直链淀粉含量、胶稠度和蛋白质含量上均展现出优越性，分别为13.68%、83.94 mm和6.65 g/100g。这表明稻-紫云英模式结合适度的氮肥用量和氮肥比例有助于提高稻米的蒸煮食味与营养品质。相比之下，当采用稻-羊肚菌模式(A3)与减氮10%(B1)和基蘖肥:穗肥比例为6:4(C2)的组合时，直链淀粉含量、胶稠度和蛋白质含量分别为12.41%、85.14 mm和6.23 g/100g，这亦是一个相对较好的组合。
表6 不同处理组合对蒸煮食味与营养品质的影响
处理组合 直链淀粉含量/% 胶稠度/mm 蛋白质含量/(g/100g)
A1B1C1 12.34 bcdef 73.04 jk 5.81 s
A1B1C2 12.43 bcde 75.56 ij 5.80 t
A1B1C3 12.56 bc 79.41 cdefghi 5.63 w
A1B1C4 12.52 bcd 76.14 hij 5.84 q
A1B2C1 11.89 cdef 70.56 k 6.16 k
A1B2C2 11.78 def 77.89 fghi 6.78 b
A1B2C3 11.72 ef 73.07 jk 6.91 a
A1B2C4 11.65 f 72.45 jk 6.77 c
A2B1C1 13.63 a 77.24 ghi 6.10 l
A2B1C2 13.68 a 83.94 ab 6.65 d
A2B1C3 13.10 ab 80.48 bcdefg 5.97 n
A2B1C4 13.54 a 80.14 cdefgh 5.78 v
A2B2C1 12.12 cdef 78.08 efghi 6.31 h
A2B2C2 12.08 cdef 78.12 defghi 6.40 e
A2B2C3 12.01 cdef 82.18 abcd 6.32 g
A2B2C4 12.02 cdef 78.28 defghi 6.29 i
A3B1C1 12.46 bcde 79.74 cdefgh 5.82 r
A3B1C2 12.41 bcdef 85.14 a 6.23 j
A3B1C3 12.53 bcd 81.98 abcde 5.85 p
A3B1C4 12.35 bcdef 83.85 ab 5.29 x
A3B2C1 11.97 cdef 78.98 cdefghi 6.03 m
A3B2C2 11.91 cdef 80.77 bcdefg 6.36 f
A3B2C3 11.85 cdef 81.89 abcdef 5.94 o
A3B2C4 11.81 cdef 82.78 abc 5.79 u
因此，为了达到较好的稻米蒸煮食味与营养品质，应考虑采用稻-紫云英、或稻-羊肚菌的轮作模式，并结合适量减氮的氮肥施用和合理的氮肥比例策略。

2.2 RVA谱特征值分析

根据表7数据，可以看出，轮作模式对崩解值、冷胶黏度、消减值及糊化温度有显著影响；氮肥用量主要影响糊化温度；而氮肥比例则对峰值黏度、热浆黏度和崩解值有显著或极显著效应。当考虑轮作模式与氮肥用量的互作时，峰值黏度、崩解值和糊化温度均受到了显著或极显著影响。轮作模式与氮肥比例的互作对峰值黏度和糊化温度有极显著影响。但在考虑所有3个因子的互作时，仅峰值黏度受到极显著影响。
表7 不同因素及其互作对RVA谱特征值影响的方差分析
因素 峰值黏度 热浆黏度 崩解值 冷胶黏度 消减值 峰值时间 糊化温度
A ns ns * * * ns *
B ns ns ns ns ns ns **
C * ** * ns ns ns ns
A×B ** ns ** ns ns ns *
A×C ** ns ns ns ns ns **
B×C ns ns ns ns ns ns ns
A×B×C ** ns ns ns ns ns ns
对于不同处理组合下的RVA谱特征值，从表8中可以发现明显的变化规律和趋势。从峰值黏度来看，A2B2C4处理组合达到了最高的2239 cP，明显高于其他组合，而A1B1C2、A1B1C3和A1B1C4的处理组合在2065~2076 cP的范围内，数值相对较低，表明氮肥运筹和轮作模式在这些组合下对稻米的峰值黏度产生了显著的影响。热浆黏度方面，A2B2C4达到最高的1212 cP，而A3B1C3处理组合则降至最低的933 cP，显示出氮肥比例与轮作模式可能与热浆黏度的差异有关。崩解值中，A3B1C3处理组合达到最高的1087 cP，而A3B1C2和A3B2C3的处理组合却表现出较低的804 cP和985 cP，这可能意味着氮肥比例在这一特性上起到了关键作用。观察冷胶黏度，A1B2C3处理组合有最高值1914 cP，与A3B1C1的最低值1615 cP相比，显示出轮作模式与氮肥用量可能对冷胶黏度产生了较大影响。消减值方面，A1B2C1和A3B2C3的处理组合达到了相对较低的712 cP和615 cP，而A1B2C2则达到最高的849 cP，表明氮肥比例可能与此特征有显著关联。糊化温度在A2B1C2和A3B1C1的处理组合中达到最低的73.1℃和72.3℃，与多数其他处理在84~86℃范围内的数值相比，显示出显著差异。这可能表示轮作模式与氮肥比例共同影响了糊化温度。
表8 不同处理组合对RVA谱特征值的影响
处理组合 峰值黏度/cP 热浆黏度/cP 崩解值/cP 冷胶黏度/cP 消减值/cP 峰值时间/min 糊化温度/℃
A1B1C1 2182 ab 1147 ab 1035 abc 1872 abc 725 bc 6.0 a 84.5 a
A1B1C2 2065 bcdef 1113 abc 952 cdefg 1870 abc 757 abc 6.1 a 85.2 a
A1B1C3 2067 bcdef 1095 abc 972 cdef 1819 abcd 724 abc 6.0 a 84.5 a
A1B1C4 2076 abcdef 1144 ab 932 defgh 1893 a 749 abc 6.0 a 84.3 a
A1B2C1 2082 abcdef 1143 ab 936 defg 1855 abc 712 abc 6.0 a 85.2 a
A1B2C2 1878 hi 987 cd 923 efgh 1836 abcd 849 a 5.9 a 86.0 a
A1B2C3 1973 efghi 1110 abc 863 ghi 1914 a 804 ab 6.1 a 86.0 a
A1B2C4 1984 defghi 1147 ab 836 hi 1887 ab 739 abc 6.1 a 86.0 a
A2B1C1 2094 abcdef 1102 abc 992 abcdef 1738 abcd 636 c 6.0 a 84.4 a
A2B1C2 2146 abcd 1127 abc 1019 abcde 1696 abcd 669 bc 5.9 a 73.1 b
A2B1C3 2112 abcde 1116 abc 996 abcdef 1772 abcd 656 c 5.9 a 84.4 a
A2B1C4 1960 efghi 1103 abc 857 ghi 1758 abcd 655 c 6.1 a 74.8 b
A2B2C1 2034 bcdefgh 1152 ab 915 fgh 1779 abcd 693 bc 6.0 a 85.2 a
A2B2C2 2161 abc 1081 abcd 1080 ab 1723 abcd 642 c 5.9 a 82.8 a
A2B2C3 2113 abcde 1068 abcd 1045 abc 1716 abcd 648 c 5.9 a 84.5 a
A2B2C4 2239 a 1212 a 1027 abcd 1885 ab 673 bc 6.0 a 84.5 a
A3B1C1 1894 ghi 1002 bcd 925 efgh 1615 d 646 c 5.9 a 72.3 b
A3B1C2 1854 i 1050 bcd 1087 a 1687 abcd 637 c 6.0 a 85.2 a
A3B1C3 2020 bcdefgh 933 d 804 i 1615 d 682 bc 5.8 a 73.3 b
A3B1C4 2144 abcd 1143 ab 1000 abcdef 1767 abcd 623 c 6.0 a 82.8 a
A3B2C1 2082 abcdef 1116 abc 966 cdef 1769 abcd 653 c 6.0 a 73.2 b
A3B2C2 1945 fghi 1021 bcd 924 efgh 1652 bcd 631 c 5.9 a 84.4 a
A3B2C3 2012 cdefghi 1027 bcd 985 bcdef 1642 cd 615 c 5.9 a 72.3 b
A3B2C4 2053 bcdefg 1101 abc 952 cdefg 1757 abcd 656 c 6.0 a 83.6 a
在主效应中，稻-羊肚菌模式(A3)和稻-紫云英模式(A2)在峰值黏度、热浆黏度、冷胶黏度等方面表现出优于稻-麦模式(A1)的效果，这可能是由于紫云英和羊肚菌与水稻的生物互动改善了土壤的肥力状况，进而影响了稻米的品质。在氮肥用量对RVA谱特征值的影响中，减氮10%的处理(B1)在热浆黏度、崩解值等方面表现出较好的效果，这可能是由于适量的减氮能够促进稻米的营养积累，进一步影响稻米的口感。而在氮肥运筹对RVA谱特征值的影响中，基蘖肥:穗肥比例为6:4(C2)的处理在峰值黏度和冷胶黏度等方面表现最佳。
在不同的处理组合中，A2B1C2（稻-紫云英模式，减氮10%，基蘖肥:穗肥比例6:4）处理的稻米表现出较高的峰值黏度和热浆黏度，这对于提高稻米的加工品质和口感具有积极的作用。而A3B1C2（稻-羊肚菌模式，减氮10%，基蘖肥:穗肥比例6:4）处理则表现出最高的崩解值，说明这种处理的稻米在糊化过程中，淀粉颗粒的崩解程度更高，进而可能会提高稻米的稳定性和抗老化能力。

3 讨论

3.1 轮作模式对稻米品质的影响

本研究对不同轮作模式对优质食味粳稻稻米品质的影响进行了深入探讨，结果表明轮作模式对稻米的加工品质、外观品质和蒸煮食味与营养品质均产生了显著影响。研究表明，稻-羊肚菌轮作模式在提高稻米加工品质和外观品质上均展现出了优越性，这可能是由于羊肚菌有利于稻田土壤中的有机质和微生物的积累和改良。刘天海等[17]发现羊肚菌栽培能够显著改善土壤理化性质，对土壤容重、有机质、碱解氮、有效磷、pH、腐殖酸等影响较大。张津京等[18]进一步证实，羊肚菌菌丝能够提升土壤有机质含量，促进有益微生物的增长，提高蔗糖酶和淀粉酶酶活性。王杰鹏等[19]指出，这种微生物的增长可以调节水稻生长，提高植株抗逆性和抗病能力，进而改善稻米品质。研究也表明，稻-紫云英模式在提高稻米蒸煮食味与营养品质上表现出较好的效果。这与汤文光等[20]研究结果一致，稻-紫云英轮作有利于稻田植株地上部生长发育和改善稻米品质，表现为胶稠度和直链淀粉含量增加，同时提高了稻米出糙率、整精米率，降低了垩白米率和垩白度。可能是因为紫云英作为一种绿肥作物，能够提高稻田土壤肥力和促进土壤养分循环。周玲红等[21]证实，冬季种植紫云英可有效增加土壤可溶性有机碳氮和微生物生物量碳氮含量，促进土壤有机碳氮的矿化速率，从而有利于稻米的营养吸收。本研究还表明稻-麦轮作模式在某些品质指标上表现相对较为一般，这可能与小麦与水稻之间的生长周期和土壤需求存在差异有关。田伟等[22]发现小麦与水稻的连续轮作可能导致土壤中某些病原微生物的累积，张国发等[23]进一步指出，小麦的生长可能会消耗土壤中的某些关键营养元素，对后续水稻生长和稻米品质可能存在潜在影响。

3.2 氮肥运筹对稻米品质的调控

在稻米品质的调控中，氮肥运筹起到了至关重要的作用。本研究的结果明确显示，不同的氮肥运筹策略对稻米的加工品质、外观品质以及蒸煮食味与营养品质都产生了显著的影响，在稻-羊肚菌、稻-紫云英轮作模式和减氮10%的施肥策略下，6:4的基蘖肥:穗肥在多数情况下都能表现出较好的稻米品质。这与胡群等[24]和刘代银等[25]对钵苗机插和免耕抛秧优质食味水稻的研究结果类似，均表明6:4的基蘖肥:穗肥较为合理，对于提高稻米的糙米率和精米率具有积极作用。然而，陈梦云等[26]认为秸秆全量还田下水稻7:3的氮肥运筹策略更为有利，提高穗肥比例显著改善稻米的加工和营养品质，但同时降低了稻米外观品质，且稻米蒸煮食味品质也有所下降。这种不一致性可能是由于所选群体、研究条件、土壤类型等因素的差异所致。不同轮作模式下氮肥运筹对优质食味粳稻稻米品质的综合调控效应尚未见报道。轮作模式与氮肥运筹的互作效应可能涉及到土壤微生物环境[27]、土壤养分动态[28]、植物生理响应[29]和环境应激[30]等多个方面。本研究表明，氮肥运筹对稻米品质的影响也与轮作模式紧密相关，为了获得最佳的稻米品质，除了选择合适的轮作模式外，还需要根据具体的土壤条件和轮作模式来调整氮肥运筹策略。

3.3 RVA谱特征值与稻米品质

淀粉RVA谱特征是评估稻米食味品质的核心参数。当其峰值黏度较高、崩解值较大以及消减值的绝对值较小时，代表稻米的食味品质较好[31]。本研究中不同的轮作模式和氮肥运筹对RVA谱特征值产生了显著的影响，稻-紫云英模式和稻-羊肚菌模式在峰值黏度、热浆黏度、崩解值等方面均表现出优于稻-麦模式的效果。这可能是由于紫云英和羊肚菌与水稻的生物互动改善了土壤的肥力状况，进而影响了稻米的淀粉特性。与刘建等[32]的研究结果相一致，本研究也发现，提高穗肥用量可能会降低最高黏度、热浆黏度和崩解值，但会使消解值与糊化温度升高。而胡雅杰等[33]指出，施氮量增加会导致稻米的峰值黏度与崩解值下降，而热浆黏度和消减值则上升，可能会对稻米蒸煮食味品质产生负面效果。此外，由于优良食味粳稻中的直链淀粉与支链淀粉的比例的特殊性，也会对稻米淀粉的RVA谱特征产生影响，进而对其蒸煮食味品质产生影响[34]。在本研究中，不同的轮作模式和氮肥运筹组合对稻米的RVA谱特征值产生了不同的影响，这为进一步优化稻米的食味品质提供了重要的参考。
本研究的结果对于农业生产具有重要的实用价值。通过选择合适的轮作模式和氮肥运筹，可以有效地提高稻米的品质，从而增加其市场价值。然而，本研究也存在局限性，如研究期限、轮作模式和土壤类型的选择等，可能无法在不同的地理和气候条件下全面反映长期轮作对稻米品质的影响。在未来的研究中，将进一步探讨不同轮作模式对土壤微生物群落的影响，以及这些变化与稻米品质的联系。

4 结论

本研究初步明确了不同轮作模式与氮肥运筹对优质食味粳稻稻米品质的显著调控效应。与传统的稻-麦轮作模式相比，稻-紫云英和稻-羊肚菌轮作模式在品质调控上表现出更为显著的优势，结合减氮10%与6:4的氮肥运筹能够显著提高稻米的加工品质、外观品质、蒸煮食味与营养品质及RVA谱特征值。轮作模式、氮肥用量和氮肥比例之间存在的互作效应进一步强化了这些调控效果。为实现优质食味粳稻的高品质稻米生产，可采用稻-羊肚菌或稻-紫云英轮作模式，并结合适宜的氮肥管理策略。

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