水稻生长过程中籽粒水分状态和横向弛豫特性分析

doi:10.3864/j.issn.0578-1752.2017.02.004

摘要/Abstract

摘要： 【目的】基于低场核磁共振（LF-NMR）技术观察水稻抽穗到成熟过程中籽粒水分状态的变化，探讨淀粉、蛋白质的积累效应对前者的影响，为水稻品质形成规律提供数据参考。【方法】对抽穗后63 d内的“越光”有机稻间隔采样，测定籽粒的百粒重、硬度、水分含量、淀粉含量和蛋白质含量，比较水稻在抽穗后不同生长时期的整体品质变化，通过LF-NMR测定的横向弛豫参数定性和定量地分析籽粒中的水分动态，探讨水稻品质形成过程与水分状态的相关联系。【结果】在抽穗后7—14 d百粒重和淀粉含量增长速率最快，两者表现极显著正相关（P＜0.01）。籽粒硬度分别与淀粉含量和蛋白质含量都极显著正相关（P＜0.01），与水分含量极显著负相关（P＜0.01）。水分含量在抽穗后7—56 d从57.16%呈指数型下降到22.39%。淀粉含量在抽穗后42 d内呈“S”型曲线增长至50.47 g/100 g湿基。蛋白质含量在抽穗后7—49 d线性地增长至峰值6.56 g/100 g湿基。总体而言，水稻籽粒在抽穗后49 d左右整体品质已经形成。LF-NMR反演图谱表明在抽穗后7 d内籽粒部分水分往高自由度方向移动。抽穗后7—21 d，反演曲线整体向左迁移，表征流动性最弱“结合水”的T_2b峰出现，表征束缚水的T₂₂峰发生峰的分化现象。4种横向弛豫时间T_2b、T₂₁、T₂₂和T₂₃随生长时期而减小，水稻籽粒中的整体氢质子自由度在逐渐降低。抽穗后21 d左右，“结合水”的峰比例超过束缚水和自由水的峰比例总和。籽粒内的水分含量和横向弛豫参数（T_2b、T₂₁、T₂₂、T₂₃、A_2b、A₂₂和A₂₃）都随生长时期极显著地变化（P＜0.01），而且淀粉和蛋白质积累与前者变化具有极显著的相关性（P＜0.01）。尤其是淀粉充实胚乳细胞，水分子被淀粉颗粒包围或与其中的亲水基团氢键作用，使得整体水分状态向“结合水”的方向迁移。基于对横向弛豫信号的主成分分析，发现抽穗后42 d内不同生长时期的籽粒水分状态差异显著，42 d后整体趋于稳定。【结论】水稻灌浆期间，籽粒内部水分状态与淀粉和蛋白质的积累显著相关；“结合水”的比例逐渐上升，束缚水和自由水的比例显著下降；利用LF-NMR技术可以有效分析抽穗后不同生长时期籽粒的整体水分动态变化。

关键词: 水稻, 籽粒淀粉积累, 水分状态, 核磁共振, 横向弛豫特性, 主成分分析

Abstract: 【Objective】This paper is mainly to observe the changes of moisture state of grains during the growth process of rice after heading by LF-NMR, and investigate the effects of accumulation of starch and protein on the moisture state of kernels, which provides reference for formation regularity of rice qualities.【Method】The spikes of “Yueguang” organic rice were sampled at intervals for 63 days after heading (DAH), The 100-grain weight, hardness, moisture content, starch content and protein content were measured, separately, which were used to compare the changes of whole qualities of grains at different growth periods of rice after heading. The transverse relaxation parameters determined by LF-NMR were adopted to qualitatively and quantitatively analyze the moisture dynamics of grains. What’s more, the internal relationships between the formation process of rice qualities and moisture state of grains were discussed.【Result】Both the 100-grain weight and starch content for “Yueguang” organic rice increased most quickly within 7-14 DAH, and they were highly significantly and positively correlated with each other (P＜0.01). Hardness was highly significantly and positively correlated with starch content and protein content, respectively (P＜0.01), and negatively correlated with moisture content (P＜0.01). Within 7-56 DAH, moisture content exponentially declined from 57.16% to 22.39%. With showing an S-shaped growth curve, starch content increased to 50.47 g/100g wet-basis within 42 DAH. Protein content linearly increased to the peak value of 6.56 g/100g wet-basis within 7-49 DAH. On the whole, the whole qualities of rice grains had been formed within 49 DAH. The inversion spectrum of LF-NMR data showed that part of water in the grains moved towards the direction of high mobility within 7 DAH. Within 7-21 DAH, when inversion curves of LF-NMR data gradually moved towards the left, T_2b peak representing for “bound water" with least mobility occurred, followed by the differentiation phenomenon of T₂₂ peak representing for capillary water. Four kinds of transverse relaxation times including T_2b, T₂₁, T₂₂ and T₂₃, gradually decreased with growth periods, which indicated that whole proton degree of freedom in the kernels dropped obviously. The peak ratio of “bound water” was more than peak ratio summation of capillary water and free water at 21 DAH. The moisture content and transverse relaxation parameters (T_2b, T₂₁, T₂₂, T₂₃, A_2b, A₂₂ and A₂₃) of grains varied with growth periods highly significantly (P＜0.01), which were also highly significantly (P＜0.01) related to the accumulation of starch and protein. With endosperm cells filled with starch granules, most of water molecules in the grains were mainly surrounded by starch granules and formed hydrogen bonds with hydrophilic groups of starch, which resulted in the migration of moisture state towards the direction of “bound water”. According to principal component analysis of transverse relaxation signals, there were significant differences among the moisture state of rice grains for different growth periods within 42 DAH, followed by keeping steady.【Conclusion】It was found that during the grain-filling process of rice, the moisture state of grains was significantly related to the accumulation of starch and protein. The ratio of “bound water” gradually increased, and the ratios of capillary water and free water significantly decreased in the grains. LF-NMR can be adopted to effectively analyze the whole moisture dynamic changes of grains at different growth periods of rice after heading.

Key words: rice, starch accumulation of grains, moisture state, nuclear magnetic resonance, transverse relaxation characteristics, principal component analysis

邵小龙，汪楠，时小转，沈飞，宋伟，张强. 水稻生长过程中籽粒水分状态和横向弛豫特性分析[J]. 中国农业科学, 2017, 50(2): 240-249.

SHAO XiaoLong, WANG Nan, SHI XiaoZhuan, SHEN Fei, SONG Wei, ZHANG Qiang. Analysis of Moisture State and Transverse Relaxation Characteristics of Grains During the Growth Process of Rice[J]. Scientia Agricultura Sinica, 2017, 50(2): 240-249.

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参考文献

[1] Zhang H, Li H W, Yuan L M, Wang Z Q, Yang J C, Zhang J H. Post-anthesis alternate wetting and moderate soil drying enhances activities of key enzymes in sucrose-to-starch conversion in inferior spikelets of rice. Journal of Experimental Botany, 2012, 63(1): 215-227.

[2] Shao G C, Deng S, Liu N, Yu S N, Wang M H, She D L. Effects of controlled irrigation and drainage on growth, grain yield and water use in paddy rice. European Journal of Agronomy, 2014, 53: 1-9.

[3] Borisjuk L, Rolletschek H, Neuberger T. Surveying the plant’s world by magnetic resonance imaging. The Plant Journal, 2012, 70(1): 129-146.

[4] 李栋梁, 李小刚, 顾蕴洁, 王忠. 不同类型水稻品种胚乳发育的研究. 中国农业科学, 2014, 47(19): 3757-3768.

Li D L, Li X G, Gu Y J, Wang Z. Investigation of endosperm cell development of different rice varieties. Scientia Agricultura Sinica, 2014, 47(19): 3757-3768. (in Chinese)

[5] 孙金才, 杨泽敏. 灌浆期淀粉合成酶动态与米质的相关性分析. 中国农学通报, 2008, 24(7): 32-38.

Sun J C, Yang Z M. Studies on correlation of rice quality and enzyme activity in grain filling period. Chinese Agricultural Science Bulletin, 2008, 24(7): 32-38. (in Chinese)

[6] 马启林, 李阳生, 田小海, 鄢圣之, 雷慰慈, 中田升. 高温胁迫对水稻贮藏蛋白质的组成和积累形态的影响. 中国农业科学, 2009, 42(2): 714-718.

Ma Q L, Li Y S, Tian X H, Yan S Z, Lei W C, Zhong T S. Influence of high temperature stress on composition and accumulation configuration of storage protein in rice. Scientia Agricultura Sinica, 2009, 42(2): 714-718. (in Chinese)

[7] 王贺正, 马均, 李旭毅, 张荣萍. 水分胁迫对水稻籽粒灌浆及淀粉合成有关酶活性的影响. 中国农业科学, 2009, 42(5): 1550-1558.

Wang H Z, Ma J, Li X Y, Zhang R P. Effects of water stress on grain filling and activities of enzymes involved in starch synthesis in rice. Scientia Agricultura Sinica, 2009, 42(5): 1550-1558. (in Chinese)

[8] 要世瑾, 杜光源, 牟红梅, 栾翔宇, 马鸿雁, 刘嘉男, 刘梦达, 齐笑, 何建强. 基于核磁共振技术检测小麦植株水分分布和变化规律. 农业工程学报, 2014, 30(24): 177-186.

Yao S J, Du G Y, Mou H M, Luan X Y, Ma H Y, Liu J N, Liu M D, Qi X, He J Q. Detection of water distribution and dynamics in body of winter wheat based on nuclear magnetic resonance. Transactions of the Chinese Society of Agricultural Engineering, 2014, 30(24): 177-186. (in Chinese)

[9] 宋平, 徐静, 马贺男, 王成, 杨涛, 高鹤. 用低场核磁共振检测水稻浸种过程中种子水分的相态及分布特征. 农业工程学报, 2016, 32(6): 204-210.

Song P, Xu J, Ma H N, Wang C, Yang T, Gao H. Moisture phase state and distribution characteristics of seed during rice seed soaking process by low field nuclear magnetic resonance. Transactions of the Chinese Society of Agricultural Engineering, 2016, 32(6): 204-210. (in Chinese)

[10] Shao X L, Li Y F. Application of low-field NMR to analyze water characteristics and predict unfrozen water in blanched sweet corn. Food and Bioprocess Technology, 2013, 6: 1593-1599.

[11] Ruan R R, Chen P L, Almaer S. Nondestructive analysis of sweet corn maturity using NMR. HortScience, 1999, 34(2): 319-321.

[12] Musse M, Quellec S, Cambert M, Devaux M F, Lahaye M, Mariette F. Monitoring the postharvest ripening of tomato fruit using quantitative MRI and NMR relaxometry. Postharvest Biology and Technology, 2009, 53(1): 22-35.

[13] Horigane A K, Takahashib H, Maruyamac S, Ohtsuboa K, Yoshidaa M. Water penetration into rice grains during soaking observed by gradient echo magnetic resonance imaging. Journal of Cereal Science, 2006, 44(3): 307-316.

[14] Hwang S S, Cheng Y C, Chang C, Lur H S, Lin T T. Magnetic resonance imaging and analyses of tempering processes in rice kernels. Journal of Cereal Science, 2009, 50(1): 36-42.

[15] Seefeldt H F, van den Berg F, Köckenberger W, Engelsenb S B, Wollenwebera B. Water mobility in the endosperm of high beta-glucan barley mutants as studied by nuclear magnetic resonance imaging. Magnetic resonance imaging, 2007, 25(3): 425-432.

[16] 邵小龙, 宋伟, 李云飞. 粮油食品低场核磁共振检测技术研究进展. 中国粮油学报, 2013, 28(7): 114-118.

Shao X L, Song W, Li Y F. Research process of low-field nuclear magnetic resonance (LF-NMR) detection technology in grain and oil food. Journal of the Chinese Cereals and Oils Association, 2013, 28(7): 114-118. (in Chinese)

[17] 王海莲, 万向元, 胡培松, 翟虎渠, 万建民. 稻米脂肪含量近红外光谱分析技术研究. 中国农业科学, 2005, 38(8): 1540-1546.

Wang H L, Wan X Y, Hu P S, Zhai H Q, Wan J M. Quantitative analysis of fat content in brown rice by near infrared spectroscopy (NIRS) technique. Scientia Agricultura Sinica, 2005, 38(8): 1540-1546. (in Chinese)

[18] Marcone M F, Wang S N, Albabish W, Nie S P, Somnarain D, Hill A. Diverse food-based applications of nuclear magnetic resonance (NMR) technology. Food Research International, 2013, 51(2): 729-747.

[19] Yang P F, Li X J, Wang X Q, Chen H, Chen F, Shen S H. Proteomic analysis of rice (Oryza sativa) seeds during germination. Proteomics, 2007, 7(18): 3358-3368.

[20] Rathjen J R, Strounina E V, Mares D J. Water movement into dormant and non-dormant wheat (Triticum aestivum L.) grains. Journal of Experimental Botany, 2009, 60(6): 1619-1631.

[21] 金正勋, 杨静, 钱春荣, 刘海英, 金学泳, 秋太权. 灌浆成熟期温度对水稻籽粒淀粉合成关键酶活性及品质的影响. 中国水稻科学, 2005, 19(4): 377-380.

Jin Z X, Yang J, Qian C R, Liu H Y, Jin X Y, Qiu T Q. Effects of temperature during grain filling period on activities of key enzymes for starch synthesis and rice grain quality. Chinese Journal of Rice Science, 2005, 19(4): 377-380. (in Chinese)

[22] Ishimaru T, Horigane A K, Ida M, Iwasawa N, San-oh Y A, Nakazono M, Nishizawa N K, Masumura T, Kondo M, Yoshida M. Formation of grain chalkiness and changes in water distribution in developing rice caryopses grown under high-temperature stress. Journal of Cereal Science, 2009, 50(2): 166-174.

[23] Tanaka K, Onishi R, Miyazaki M, Ishibashi Y, Yuasa T, Inoue M. Changes in NMR relaxation of rice grains, kernel quality and physicochemical properties in response to a high temperature after flowering in heat-tolerant and heat-sensitive rice cultivars. Plant Production Science, 2009, 12(2): 185-192.

[24] Castro C, Gazza L, Ciccoritti R, Pogna N, Rossi C O, Manetti C. Development of wheat kernels with contrasting endosperm

texture characteristics as determined by magnetic resonance imaging and time domain-nuclear magnetic resonance. Journal of Cereal Science, 2010, 52(2): 303-309.

[25] Krishnan P, Chopra U K, Verma A P S, Joshi D K, Chand I. Nuclear magnetic resonance relaxation characterisation of water status of developing grains of maize (Zea mays L.) grown at different nitrogen levels. Journal of Bioscience and Bioengineering, 2014, 117(4): 512-518.

[26] Kasai M, Lewis A R, Ayabe S, Hatae K, Fyfe C A. Quantitative NMR imaging study of the cooking of japonica and indica rice. Food Research International, 2007, 40(8): 1020-1029.

[27] 陈银基, 蒋伟鑫, 曹俊, 戴炳业, 董文. 温湿度动态变化过程中不同含水量稻谷的储运特性. 中国农业科学, 2016, 49(1): 163-175.

Chen Y J, Jiang W X, Cao J, Dai B Y, Dong W. Storage and transportation characteristic of different moisture paddy rice dealt with dynamic temperature and humidity. Scientia Agricultura Sinica, 2016, 49(1): 163-175. (in Chinese)

[28] Thompson J F, Mutters R G. Effect of weather and rice moisture at harvest on milling quality of California medium-grain rice. Transactions of the ASABE, 2006, 49(2): 435-440.

[29] 徐兴凤, 钟业俊, 夏文, 刘成梅, 刘伟, 左艳娜, 艾亦旻. 不同采收期籼米外观与米饭食味品质的相关性分析. 食品科学, 2013, 34(23): 147-150.

Xu X F, Zhong Y J, Xia W, Liu C M, Liu W, Zuo Y N, Ai Y M. Correlational analysis between appearance and eating quality of indica rice harvested at different times. Food Science, 2013, 34(23): 147-150. (in Chinese)

[30] Fang G H, Goh J Y, Tay M, Lau H F, Li S F Y. Characterization of oils and fats by ¹H NMR and GC/MS fingerprinting: classification, prediction and detection of adulteration. Food chemistry, 2013, 138(2): 1461-1469.

[31] Monakhova Y B, Rutledge D N, Roßmann A, Waiblinger H U, Mahler M, Ilse M, Kuballa T, Lachenmeier D W. Determination of rice type by ¹H NMR spectroscopy in combination with different chemometric tools. Journal of Chemometrics, 2014, 28(2): 83-92.

[32] Li T, Tu C H, Rui X, Gao Y W, Li W, Wang K, Xiao Y. Study of water dynamics in the soaking, steaming, and solid-state fermentation of glutinous rice by LF-NMR: A novel monitoring approach. Journal of Agricultural and Food Chemistry, 2015, 63(12): 3261-3270.