干湿交替灌溉对籼粳杂交稻产量与水分利用效率的影响及其生理基础

doi:10.3864/j.issn.0578-1752.2021.07.014

摘要/Abstract

摘要： 【目的】探明干湿交替灌溉对籼粳杂交稻产量与水分利用效率的影响及其生理基础。【方法】试验于2017—2018年在中国水稻研究所富阳基地皇天畈试验农场进行。以2个新选育的超高产籼粳杂交稻品种甬优1540和春优927以及2个当地高产粳型杂交稻品种常优5号和嘉优5号为材料,开展大田试验。自移栽后7 d至成熟期设置2种灌溉模式,分别为常规灌溉（CI）和干湿交替灌溉（AWD）。研究AWD对籼粳杂交稻品种产量形成以及水分利用效率的影响及其生理基础。【结果】与CI模式相比,AWD模式显著降低了2个当地高产粳型杂交稻品种的产量,降幅为12.3%—12.8%,但2个籼粳杂交稻品种的产量在2种灌溉模式间没有显著差异;AWD模式显著提高了4个供试水稻品种的水分利用效率,其中当地高产粳型杂交稻品种的增幅为5.9%—8.3%,籼粳杂交稻品种的增幅为13.7%—16.8%。与当地高产粳型杂交稻相比,籼粳杂交稻品种在AWD模式下获得相对较高的产量以及水分利用效率主要得益于其强大的分蘖发生能力、较高的群体颖花量和结实率、齐穗至成熟期较高的光合势与作物生长速率、齐穗后2次土壤落干期与复水期较高的根系氧化力、剑叶净光合速率以及籽粒中较高的蔗糖-淀粉代谢途径关键酶的活性。【结论】与当地高产粳型杂交稻品种相比,籼粳杂交稻品种在AWD模式下可获得更高的产量与水分利用效率;较好的根系性能（齐穗后2次土壤落干期与复水期较高的根系氧化力）和地上部植株较强的生理活性（齐穗至成熟期较高的光合势、作物生长速率、齐穗后2次土壤落干期与复水期较高的剑叶净光合速率以及籽粒中蔗糖-淀粉代谢途径关键酶活性）是其在AWD模式下获得高产与水分高效利用的重要生理基础。

关键词: 水稻, 干湿交替灌溉, 籼粳杂交稻, 产量, 水分利用效率, 生理基础

Abstract:

【Objective】 This study aimed to understand the yield formation characteristics and water use efficiency (WUE) of indica/japonica hybrid rice (IJHR) cultivar and their physiological bases under alternate wetting and soil drying (AWD) regime.【Method】The field experiment was conducted at Huangtianfan Experimental Farm which belonged to China National Rice Research Institute, Hangzhou, Zhejiang province, Southeast China, in 2018 and 2019. Two newly-bred IJHR cultivars, Yongyou 1540 and Chunyou 927, and two local high-yielding japonica hybrid rice (JHR) cultivars, Changyou 5 and Jiayou 5, were field grown. Two irrigation regimes, conventional irrigation (CI) and AWD, were imposed from 7 days after transplanting to maturity. The goals of this study were to investigate the effects of AWD on the grain yield and WUE of IJHR and its physiological bases. 【Result】Compared with those under CI regime, the grain yields of JHR cultivars were significantly decreased by 12.3%-12.8% under AWD regime, whereas the difference in grain yields of IJHR cultivars was not significant between CI and AWD regimes. Compared with the CI regime, AWD significantly reduced the amount of irrigation water and significantly increased WUE by 5.9%-8.3% and 13.7%-16.8% in JHR and IJHR cultivars, respectively. In comparison with JHR cultivars, IJHR cultivars showed greater tillering capacity, larger sink size and higher grain filling rate, higher leaf area duration and crop growth rate from heading to maturity, higher root oxidation activity, leaf photosynthetic rates, and activities of sucrose synthase and adenosine diphosphate-glucose pyrophosphorylase in grains at the first and second soil drying periods as well as re-watering periods after heading.【Conclusion】IJHR cultivars could obtain higher grain yields and higher WUE than JHR cultivars under AWD regime. Stronger physiological activities of root, including higher root oxidation activity at the first and second soil drying periods as well as re-watering periods after heading, and above ground plants, including higher leaf area duration, crop growth rate from heading to maturity, greater leaf photosynthetic rates, and activities of sucrose synthase and adenosine diphosphate-glucose pyrophosphorylase in grains at the first and second soil drying periods as well as re-watering periods after heading, contributed to their better yield performance and higher WUE of IJHR cultivars under AWD regime.

Key words: rice, alternate wetting and soil drying, indica-japonica hybrid rice, grain yield, water use efficiency, physiological bases

褚光,徐冉,陈松,徐春梅,王丹英,章秀福. 干湿交替灌溉对籼粳杂交稻产量与水分利用效率的影响及其生理基础[J]. 中国农业科学, 2021, 54(7): 1499-1511.

CHU Guang,XU Ran,CHEN Song,XU ChunMei,WANG DanYing,ZHANG XiuFu. Effects of Alternate Wetting and Soil Drying on the Grain Yield and Water Use Efficiency of Indica-Japonica Hybrid Rice and Its Physiological Bases[J]. Scientia Agricultura Sinica, 2021, 54(7): 1499-1511.

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链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2021.07.014

https://www.chinaagrisci.com/CN/Y2021/V54/I7/1499

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参考文献 48

[1]	邹应斌, 黄敏. 转型期作物生产发展的机遇与挑战. 作物学报, 2018,44(6):791-795.
	ZOU Y B, HUANG M. Opportunities and challenges for crop production in China during the transition period. Acta Agronomica Sinica, 2018,44(6):791-795. (in Chinese)
[2]	ZHANG G Q . Prospects of utilization of inter-subspecific heterosis between indica and japonica rice. Journal of Integrative Agriculture. 2020,19(1):1-10.
[3]	林建荣, 宋昕蔚, 吴明国, 程式华. 籼粳超级杂交稻育种技术创新与品种培育. 中国农业科学, 2016,49(2):207-218.
	LIN J R, SONG X W, WU M G, CHENG S H. Breeding technology innovation of indica-japonica super hybrid rice and varietal breeding. Scientia Agricultura Sinica, 2016,49(2):207-218. (in Chinese)
[4]	BELDER P, BOUMAN R. CABANGON G, LU G, QUILANG Y H, LI J, SPIERTZ J H, TUONG T P . Effect of water-saving irrigation on rice yield and water use in typical lowland conditions in Asia. Agricultural Water Management, 2004,65(3):193-210.
[5]	BOUMAN B . A conceptual framework for the improvement of crop water productivity at different spatial scales. Agricultural Systems, 2007,93(1/3):43-60.
[6]	WEI H H, MENG T Y, LI C, XU K, HUO Z Y, WEI H Y, GUO B W, ZHANG H C, DAI Q G . Comparisons of grain yield and nutrient accumulation and translocation in high-yielding japonica/indica hybrids, indica hybrids, and japonica conventional varieties. Field Crops Research, 2017,204:101-109.
[7]	WEI H Y, ZHANG H C, BLUMWALD E, LI H L, CHENG J Q, DAI Q G, HUO Z Y, XU M, GUO B W . Different characteristics of high yield formation between inbred japonica super rice and inter-sub- specific hybrid super rice. Field Crops Research, 2016,198:179-187.
[8]	MENG T Y, WEI H H, LI X Y, DAI Q G, HUO Z Y . A better root morpho-physiology after heading contributing to yield superiority of japonica/indica hybrid rice. Field Crops Research, 2018,228:135-146.
[9]	周磊, 刘秋员, 田晋钰, 朱梦华, 程爽, 车阳, 王志杰, 邢志鹏, 胡雅杰, 刘国栋, 魏海燕, 张洪程. 甬优系列籼粳杂交稻产量及氮素吸收利用的差异. 作物学报, 2020,46(5):772-786.
	ZHOU L, LIU Q Y, TIAN J Y, ZHU M H, CHENG S, CHE Y, WANG Z J, XING Z P, HU Y J, LIU G D, WEI H Y, ZHANG H C. Differences in yield and nitrogen absorption and utilization of indica-japonica hybrid rice varieties of Yongyou series. Acta Agronomica Sinica, 2020,46(5):772-786. (in Chinese)
[10]	LIANG K M, ZHONG X H, HUANG R R, LAMPAYAN R M, PAN J F, TIAN K, LIU Y . Grain yield, water productivity and CH₄ emission of irrigated rice in response to water management in south China. Agricultural Water Management, 2016,163:319-331.
[11]	ZHANG Y N, LIU M J, SAIZ G, DANNENMANN M, GUO L, TAO Y Y, SHI J C, ZUO Q, BUTTERBACH K, LI G Y, LIN S . Enhancement of root systems improves productivity and sustainability in water saving ground cover rice production system. Field Crops Research, 2017,213:186-193.
[12]	张自常, 李鸿伟, 陈婷婷, 王学明, 王志琴, 杨建昌. 畦沟灌溉和干湿交替灌溉对水稻产量与品质的影响. 中国农业科学, 2011,44(24):4988-4998.
	ZHANG Z C, LI H W, CHEN T T, WANG X M, WANG Z Q, YANG J C. Effect of furrow irrigation and alternate wetting and drying irrigation on grain yield and quality of rice. Scientia Agricultura Sinica, 2011,44(24):4988-4998. (in Chinese)
[13]	YANG J C, ZHOU Q, ZHANG J H . Moderate wetting and drying increases rice yield and reduces water use, grain arsenic level, and methane emission. The Crop Journal, 2017,5(2):151-158.
[14]	褚光, 展明飞, 朱宽宇, 王志琴, 杨建昌. 干湿交替灌溉对水稻产量与水分利用效率的影响. 作物学报, 2016,42(7):1026-1036.
	CHU G, ZHAN M F, ZHU K Y, WANG Z Q, YANG J C. Effects of alternate wetting and drying irrigation on yield and water use efficiency of rice. Acta Agronomica Sinica, 2016,42(7):1026-1036. (in Chinese)
[15]	ZHANG H, XUE Y G, WANG Z Q, YANG J C, ZHANG J H . An alternate wetting and moderate soil drying regime improves root and shoot growth in rice. Crop Science, 2009,49(6):2246-2260.
[16]	陆大克, 段骅, 王维维, 刘明爽, 魏艳秋, 徐国伟. 不同干湿交替灌溉与氮肥形态耦合下水稻根系生长及功能差异. 植物营养与肥料学报, 2019,25(8):1362-1372.
	LU D K, DUAN H, WANG W W, LIU M S, WEI Y Q, XU G W. Comparison of rice root development and function among different degrees of dry-wet alternative irrigation coupled with nitrogen forms. Journal of Plant Nutrition and Fertilizers, 2019,25(8):1362-1372. (in Chinese)
[17]	陈鸿飞, 庞晓敏, 张仁, 张志兴, 徐倩华, 方长旬, 李经勇, 林文雄. 不同水肥运筹对再生季稻根际土壤酶活性及微生物功能多样性的影响. 作物学报, 2017,43(10):1507-1517.
	CHEN H F, PANG X M, ZHANG R, ZHANG Z X, XU Q H, FANG C X, LI J Y, LIN W X. Effects of different irrigation and fertilizer application regimes on soil enzyme activities and microbial functional diversity in rhizosphere of ratooning rice. Acta Agronomica Sinica, 2017,43(10):1507-1517. (in Chinese)
[18]	CARRIJO D R, LUNDY M E, LINQUIST B A . Rice yields and water use under alternate wetting and drying irrigation: A meta-analysis. Field Crops Research, 2017,203:173-180.
[19]	SAH R N, MIKKELSEN D S . Availability and utilization of fertilizer nitrogen by rice under alternate flooding; I. Kinetics of available nitrogen under rice culture. Plant Soil, 1983,75(2):221-226.
[20]	ERIKSEN A B, KJELDBY M, NILSEN S . The effect of intermittent flooding on the growth and yield of wetland rice and nitrogen-loss mechanism with surface applied and deep placed urea. Plant Soil, 1985,84(3):387-401.
[21]	CHU G, CHEN S, XU C M, WANG D Y, ZHANG X F . Agronomic and physiological performance of indica/japonica hybrid rice cultivar under low nitrogen conditions. Field Crops Research, 2019,243:107625.
[22]	CHU G, CHEN T T, CHEN S, XU C M, WANG D Y, ZHANG X F . Agronomic performance of drought-resistance rice cultivars grown under alternate wetting and drying irrigation management in southeast China. The Crop Journal, 2018,6(5):482-494.
[23]	CHU G, CHEN T T, WANG Z Q, YANG J C, ZHANG J H . Morphological and physiological traits of roots and their relationships with water productivity in water-saving and drought-resistant rice. Field Crops Research, 2014,162:108-119.
[24]	YANG J C, ZHANG J H, WANG Z Q, ZHU Q S, LIU L J . Activities of enzymes involved in sucrose-to-starch metabolism in rice grains subjected to water stress during filling. Field Crops Research, 2003,81(1):69-81.
[25]	YANG J C, ZHANG J H, WANG Z Q, XU G W, ZHU Q S . Activities of key enzymes in sucrose-to-starch conversion in wheat grains subjected to water deficit during grain filling. Plant Physiology, 2004,135(3):1621-1629.
[26]	杨建昌, 张建华. 促进稻麦同化物转运和籽粒灌浆的途径与机制. 科学通报, 2018,63(28/29):2932-2943.
	YANG J C, ZHANG J H. Approach and mechanism in enhancing the remobilization of assimilates and grain-filling in rice and wheat. Chinese Science Bulletin, 2018,63(28/29):2932-2943. (in Chinese)
[27]	ZHOU Q, JU C X, WANG Z Q, ZHANG H, LIU L J, YANG J C, ZHANG J H . Grain yield and water use efficiency of super rice under soil water deficit and alternate wetting and drying irrigation. Journal of Integrative Agriculture, 2017,16(5):1028-1043.
[28]	HUANG M, YANG C L, JI Q M, JIANG L G, TAN J L, LI Y Q . Tillering responses of rice to plant density and nitrogen rate in a subtropical environment of southern China. Field Crops Research, 2013,149:187-192.
[29]	AO H J, PENG S B, ZOU Y B, TANG Q Y, VISPERAS R M . Reduction of unproductive tillers did not increase the grain yield of irrigated rice. Field Crops Research, 2010,116:108-115.
[30]	李杰, 张洪程, 龚金龙, 常勇, 吴桂成, 郭振华, 戴其根, 霍中洋, 许轲, 魏海燕. 稻麦两熟地区不同栽培方式超级稻分蘖特性及其与群体生产力的关系. 作物学报, 2011,37(2):309-320.
	LI J, ZHANG H C, GONG J L, CHANG Y, WU G C, GUO Z H, DAI Q G, HUO Z Y, XU K, WEI H Y. Tillering characteristics and its relationships with population productivity of super rice under different cultivation methods in rice-wheat cropping areas. Acta Agronomica Sinica, 2011,37(2):309-320. (in Chinese)
[31]	CHU G, WANG Z Q, ZHANG H, YANG J C, ZHANG J H . Agronomic and physiological performance of rice under integrative crop management. Agronomy Journal, 2016,108(1):117-128.
[32]	CHU G, WANG Z Q, ZHANG H, LIU L J, YANG J C, ZHANG J H . Alternate wetting and moderate drying increases rice yield and reduces methane emission in paddy field with wheat straw residue incorporation. Food and Energy Security, 2015,4(3):238-254.
[33]	张洪程, 龚金龙. 中国水稻种植机械化高产农艺研究现状及发展探讨. 中国农业科学, 2014,47(7):1273-1289.
	ZHANG H C, GONG J L. Research status and development discussion on high-yielding agronomy of mechanized planting rice in China. Scientia Agricultura Sinica, 2014,47(7):1273-1289. (in Chinese)
[34]	朱德峰, 章秀福, 张玉屏. 水稻高产栽培技术的发展与展望. 中国农业科学, 2007,40(增刊 1):127-132.
	ZHU D F, ZHANG X F, ZHANG Y P. Development and prospect of high-yielding cultivation technology in rice. Scientia Agricultura Sinica, 2007,40(Suppl.1):127-132. (in Chinese)
[35]	BOYER J, WESTGATE M . Grain yields with limited water. Journal of Experimental Botany, 2004,55(407):2385-2394.
[36]	SAINI H, WESTGATE M . Reproductive development in grain crops during drought. Advances in Agronomy, 2000,68:59-96.
[37]	YANG J C, ZHANG J H, LIU K, WANG Z Q, LIU L J . Abscisic acid and ethylene interact in rice spikelets in response to water stress during meiosis. Journal of Plant Growth Regulation, 2007,26(4):318-328.
[38]	ZHANG W Y, SHENG J Y, XU Y J, XIONG F, WU Y F, WANG W L, WANG Z Q, YANG J C, ZHANG J H . Role of brassinosteroids in rice spikelet differentiation and degeneration under soil-drying during panicle development. BMC Plant Biology, 2019,19(1):409.
[39]	ZHANG W Y, SHENG J Y, FU L D, XU Y J, XIONG F, WU Y F, WANG W L, WANG Z Q, ZHANG J H, YANG J C . Brassinosteroids mediate the effect of soil-drying during meiosis on spikelet degeneration in rice. Environmental and Experimental Botany, 2020,169:103887.
[40]	KATO T, TAKEDA K . Associations among characters related to yield sink capacity in space-planted rice. Crop Science, 1996,36(5):1135-1139.
[41]	ZHANG H, CHEN T T, WANG Z Q, YANG J C, ZHANG J H . Involvement of cytokinins in the grain filling of rice under alternate wetting and drying irrigation. Journal of Experimental Botany, 2010,61(13):3719-3733.
[42]	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.
[43]	WANG G Q, LI H X, FENG L, CHEN M X, MENG S, YE N H, ZHANG J H . Transcriptomic analysis of grain filling in rice inferior grains under moderate soil drying. Journal of Experimental Botany, 2019,70(5):1597-1611.
[44]	LIANG J S, ZHANG J H, CAO X Z . Grain sink strength may be related to the poor grain filling of indica-japonica rice (Oryza Sativa) hybrids. Physiologia Plantarum, 2001,112(4):470-477.
[45]	AHMADI A, BAKER D A . The effect of water stress on the activities of key regulatory enzymes of the sucrose to starch pathway in wheat. Plant Growth Regulation, 2001,35(1):81-91.
[46]	杨建昌. 水稻根系形态生理与产量、品质形成及养分吸收利用的关系. 中国农业科学, 2011,44(1):36-46.
	YANG J C. Relationships of rice root morphology and physiology with the formation of grain yield and quality and the nutrient absorption and utilization. Scientia Agricultura Sinica, 2011,44(1):36-46. (in Chinese)
[47]	RAMASAMY S, BERGE H, PURUSHOTHAMAN S . Yield formation in rice in response to drainage and nitrogen application. Field Crops Research, 1997,51(1/2):65-82.
[48]	姜元华, 许俊伟, 赵可, 韦还和, 孙建军, 张洪程, 戴其根, 霍中洋, 许轲, 魏海燕, 郭保卫. 甬优系列籼粳杂交稻根系形态与生理特征. 作物学报, 2015,41(1):89-99.
	JIANG Y H, XU J W, ZHAO K, WEI H H, SUN J J, ZHANG H C, DAI Q G, HUO Z Y, XU K, WEI H Y, GUO B W. Root system morphological and physiological characteristics of indica-japonica hybrid rice of Yongyou series. Acta Agronomica Sinica, 2015,41(1):89-99. (in Chinese)

年份/气象条件 Year/ Meteorological condition	六月 June	七月 July	八月 August	九月 September	十月 October
2017
降雨量 Precipitation (mm)	112	218	164	72	62
日照时长 Sunshine (h)	98	220	208	185	157
平均气温 Temperature (℃)	26.5	28.6	28.5	26.8	22.5
2018
降雨量 Precipitation (mm)	71	182	151	84	77
日照时长 Sunshine (h)	118	241	185	178	164
平均气温 Temperature (℃)	26.8	28.9	28.1	26.5	22.8

方差分析 Analysis of variance	产量 Grian yield	水分利用效率 Water use efficiency	茎蘖成穗率 Percentage of productive tillers	作物生长速率 Crop growth rate	光合势 Leaf area duration	根系氧化力 Root oxidation activity	剑叶净光合速率 Flag leaf photosynthetic rate	蔗糖合酶 SuSase	腺苷二磷酸葡萄糖焦磷酸化酶 AGPase
年份 Year	NS	NS	NS	NS	NS	NS	NS	NS	NS
年份×品种 Year ×Cultivar	NS	NS	NS	NS	NS	NS	NS	NS	NS
年份×处理 Year ×Treatment	NS	NS	NS	NS	NS	NS	NS	NS	NS

品种 Cultivar	处理 Treatment	稻谷产量 Grain yield (t·hm^-2)	穗数 Number of panicles (×10⁴ hm^-2)	每穗粒数 Spikelets per panicle	总颖花量 Total spikelets (×10³ m^-2)	结实率 Filled grains (%)	千粒重 1000-grain weight (g)
甬优1540 YY-1540	CI	11.5a	202c	326a	65.7a	79.2c	23.0c
甬优1540 YY-1540	AWD	11.4a	198c	304b	59.9b	84.5b	23.3c
春优927 CY-927	CI	11.6a	189d	311b	58.8b	84.2b	24.7b
春优927 CY-927	AWD	11.5a	186d	287c	53.2c	89.6a	25.0b
常优5号 CY-5	CI	10.1b	251a	186e	46.7d	84.4b	26.8a
常优5号 CY-5	AWD	8.86c	240b	169f	40.4e	85.3b	26.8a
嘉优5号 JY-5	CI	9.82b	248a	198d	48.9d	80.7c	25.8b
嘉优5号 JY-5	AWD	8.56c	235b	181e	42.3e	81.9c	25.9b
方差分析 ANOVA
品种 Cultivar (C)		**	**	**	**	**	**
处理 Treatment (T)		**	**	**	**	**	NS
品种×处理 C×T		**	*	*	*	**	*

品种 Cultivar	处理 Treatment	茎蘖数 Number of tillers and mail stems (m^-2)			茎蘖成穗率 Percentage of productive tillers (%)
品种 Cultivar	处理 Treatment	拔节期 Jointing	齐穗期 Heading	成熟期 Maturity	茎蘖成穗率 Percentage of productive tillers (%)
甬优1540 YY-1540	CI	281c	208c	202c	71.7b
甬优1540 YY-1540	AWD	258d	207c	198c	76.2a
春优927 CY-927	CI	260d	198d	189d	72.7b
春优927 CY-927	AWD	239e	197d	186d	77.5a
常优5号 CY-5	CI	360a	259a	251a	70.4b
常优5号 CY-5	AWD	315b	245b	240b	76.1a
嘉优5号 JY-5	CI	357a	255a	248a	69.4b
嘉优5号 JY-5	AWD	309b	241b	235b	75.9a
方差分析 ANOVA
品种 Cultivar (C)		**	**	**	NS
处理 Treatment (T)		**	**	**	**
品种×处理 C×T		**	*	*	NS

品种 Cultivar	处理 Treatment	叶面积指数 LAI (m²·m^-2)			光合势 LAD (m²·m^-2·d)
品种 Cultivar	处理 Treatment	拔节期 Jointing	齐穗期 Heading	成熟期 Maturity	拔节前 Before Jointing	拔节-齐穗 Jointing-Heading	齐穗-成熟 Heading-Maturity
甬优1540 YY-1540	CI	4.67a	7.87a	1.85a	103a	251a	243a
甬优1540 YY-1540	AWD	3.71b	7.58a	1.80a	84.2b	226b	235a
春优927 CY-927	CI	4.88a	7.92a	1.91a	108a	256a	246a
春优927 CY-927	AWD	3.87b	7.63a	1.88a	87.4b	220bc	238a
常优5号 CY-5	CI	3.52b	7.01b	1.14b	80.4b	211c	204b
常优5号 CY-5	AWD	2.30c	5.56c	0.77c	56.0c	157d	158c
嘉优5号 JY-5	CI	3.68b	6.97b	1.12b	83.6b	213c	202b
嘉优5号 JY-5	AWD	2.12c	5.48c	0.71c	52.4c	152d	155c
方差分析 ANOVA
品种 Cultivar (C)		**	**	**	**	**	**
处理 Treatment (T)		**	**	**	**	*	**
品种×处理 C×T		*	**	*	**	*	*