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1 Chinese Journal of Eco-Agriculture, Jun. 2018, 26(6): DOI: /j.cnki.cjea ,,,. [J]., 2018, 26(6): LIU Y C, XIAO J X, TANG L, ZHENG Y. Effect of nitrogen application rate on root soy isoflavone exudate of wheat/faba bean intercropping system[j]. Chinese Journal of Eco-Agriculture, 2018, 26(6): * 刘英超 1, 肖靖秀 1, 汤利 1 1,2**, 郑毅 ( ; ) : 通过盆栽和水培试验, 采用小麦蚕豆间作蚕豆单作小麦单作 3 种种植方式, 研究了不同生育期不同氮水平 ( 低氮常规氮和高氮 ) 处理下, 单间作小麦和蚕豆根系大豆异黄酮分泌量的变化, 为进一步探明间作增产和控病机制提供依据结果表明, 随作物生育期推移, 小麦根系分泌大豆异黄酮数量明显减少, 蚕豆根系大豆异黄酮的分泌量先增加后减少随施氮量增加, 小麦蚕豆根系大豆异黄酮分泌量均减少, 且多达显著水平与低氮处理相比, 常规氮和高氮处理下, 单间作小麦大豆异黄酮分泌量分别显著减少, 且多达显著水平间作可以显著提高作物大豆异黄酮的分泌量, 但间作优势仅在低氮和常规氮处理下明显, 高氮处理下, 单间作小麦和蚕豆根系分泌大豆异黄酮数量差异不显著, 并且这种间作效应会随生育期的推移逐渐消失总之, 间作种植和施氮量均影响作物根系大豆异黄酮分泌量, 且低施氮量的影响更明显 : 小麦蚕豆间作 ; 施氮量 ; 根系分泌物 ; 大豆异黄酮 : S311 : A : (2018) Effect of nitrogen application rate on root soy isoflavone exudate of wheat/faba bean intercropping system * LIU Yingchao 1, XIAO Jingxiu 1, TANG Li 1, ZHENG Yi 1,2** (1. College of Resources and Environmental Sciences, Yunnan Agricultural University, Kunming , China; 2. Yunnan Provincial Department of Education, Kunming , China) Abstract: Through pot and hydroponics experiments of wheat/faba-bean, mono-cropped faba-bean and mono-cropped wheat, soy isoflavone exudate of faba-bean and wheat were studied under different nitrogen (deficient, adequate and excessive) conditions at different growth stages. The study aimed at providing the basis for further understanding of yield increase and disease control mechanisms of intercropping systems. The results showed that with the growth of crops, soy isoflavone exudate of wheat roots decreased, while that of faba-bean first increased before decreased. With the increase of nitrogen application rate, soy isoflavone exudate of both wheat and faba-bean decreased, and most changes were significant. Compared with deficient nitrogen treatment, soy isoflavone exudates of intercropped wheat were increased respectively by 18.9% and 122.1% at stem elongation stage (60 d after sowing) under adequate and excessive nitrogen conditions. Accordingly, soy isoflavone exudate of mono-cropped wheat was increased by 28.9% and 72.7% under adequate and excessive nitrogen conditions. Crops intercropping increased soy isolavone secretion of crops, which was significant only at lower nitrogen level. Compared with mono-cropped wheat under deficient and * ( ) ** :, yzheng@ynau.edu.cn, liuyingchao_1988@163.com : : * This work was supported by the National Natural Science Foundation of China ( , ) and the Yunnan Province Graduate Student Academic Award. ** Corresponding author, yzheng@ynau.edu.cn Received Oct. 7, 2017; accepted Dec. 2, 2017

2 adequate nitrogen condition at stem elongation stage of wheat (60 d after sowing), soy isoflavone exudate of intercropped wheat was significantly increased by 26.9% and 12.0%, respectively, in pot experiment, and 15.3% and 59.2% in hydroponics. Compared with mono-cropped faba-bean under deficit and adequate nitrogen rates at branching stage (60 d after sowing), root soy isoflavone secretion of intercropped faba-bean was significantly increased by 64.6%, and 11.4% in pot experiment, and 23.8% and 14.1% in hydroponics experiment, respectively. At flowering and filling stages of wheat, and flowering and pod-bearing stages of faba-bean, the similar tendency was observed. In short, both intercropping and nitrogen fertilization changed root secretion of soy isoflavone by wheat and faba-bean, which was more obvious under lower nitrogen application rate. Keywords: Wheat/faba bean intercropping; Nitrogen fertilizer rate; Root exudate; Soy isoflavone, [1], [2-3],,,, [4] (Glycine max)(medicago sativa)(pisum sativum)(cajanus cajan), [5-8],,, [9],, [10], [11], (Tricum aestivum)(vicia faba), [12] [13], [14] [15-16] [17] [18] [19],,, (HPLC),,,, 1 材料与方法 ,, : 68 mg kg 1, 16 mg kg 1, 137 mg kg 1, ph 6.08, mg kg mm 320 mm, 10 kg (mol L 1 ): K 2 SO MgSO KCl Ca(NO 3 ) KH 2 PO H 3 BO MnSO CuSO ZnSO (NH 4 ) 6 Mo 7 O Fe-EDTA L PVC, 160 mm 240 mm 42,, 1.2 A, 3 ; B, () 3,, 1.5 9, 3, 4, , 8 16 ; 6 12, 12 24, 2 :, 150 mg kg 1, 75 mg kg 1, 225 mg kg 1 ;, 100 mg kg 1 ; 100 mg kg 1

3 6 : 801 ; 1/2, 1/2,, Ca(NO) 3, mol L 1 Ca(NO) 3 50 ml, 25 ml, 75 ml, :, 7 d, 70%; 3 d,, ph 6.80~ d 95 d 131 d(, ), :,, 3, 5% 3 min, 500 ml mol L 1 CaCl 2,, 10:00, 2 h,, 20 [19], 1.4, 200 ml 100 ml 50 ml 3, 3, 10 ml 10 ml 0.45 μm HPLC Synergi 4u Hydro-RP 80A (250 mm 4.6 mm ID), A, B, : A 30%~40%(5 min) 40%~60%(10 min) 60%~90%(25 min) 90% (29 min) 90%~30%(34 min) 30%(37 min) (37 min): 270 nm ml min 1,, (1 000 ng ml 1 ) [20] 10 ng ml 1,, 1.5 Microsoft Excel 2010, SPSS 19.0 (Duncan, α=0.05) 2 结果与分析 2.1 ( 1),,,, 29.3% 50.4%, 27.2% 37.2%;, 17.6% 29.0%, 15.4% 21.0% ( 1),,,, (N/2 N 3N/2), 20.9%~ 43.4%, 13.5%~32.3%, (N/2 N 3N/2), 14.3%~17.3%, 6.7%~13.6% 表 1 不同施氮量对小麦蚕豆间作系统籽粒产量的影响 Table 1 Grain yields of wheat and faba bean intercropping system under different nitrogen application rates g plant 1 Crop Treatment Pot experiment Hydroponic experiment N/2 N 3N/2 N/2 N 3N/2 Intercropping 4.26±0.15c 5.48±0.22b 5.98±0.24a 4.84±0.19c 5.62±0.11b 6.06±0.07a Faba bean Monocropping 3.22±0.13d 4.19±0.15c 5.27±0.11b 4.26±0.20d 5.08±0.12c 5.68±0.07b Intercropping 3.47±0.14bc 4.19±0.17a 4.42±0.11a 5.22±0.12c 5.95±0.06b 6.29±0.12a Wheat Monocropping 2.42±0.13d 3.30±0.17c 3.66±0.03c 4.45±0.09d 5.21±0.11c 5.41±0.12c N/2: ; N: ; 3N/2: 5%N/2: deficient nitrogen; N: adequate nitrogen; 3N/2: excessive nitrogen. Different lowercase letters mean significant differences at 5% level among different treatments for the same crop at the same experiment.

4 ,, ;,, ( 131 d) ;, ( 131 d) ( 95 d) 2,,,,, ( 60 d),, 36.6% 104.8%, 18.1% 49.6%; ( 95 d),, 43.4% 75.9%, 13.1% 15.5%, ( 60 d),,, 1.3% 139.4%, 39.8% 95.7% 2,,,, ( 60 d),, 29.6% 12.0%, 15.3% 59.2%;, ( 95 d), 71.4%, 表 2 不同施氮量对单间作小麦不同生育期根系分泌大豆异黄酮的影响 1 Table 2 Soy isoflavone secreted by mono- and inter-cropped wheat root under different N rates at different growth stages ng plant 1 h N/2 N 3N/2 Pot experiment Hydroponic experiment Treatment Jointing stage Flower stage Filling stage Jointing stage Flower stage Filling stage Intercropping ±3.26a 85.00±2.40a 45.00±1.84a 98.75±2.36a 87.50±2.55a 22.50±2.44a Monocropping 81.67±2.60b 49.58±2.76cd 47.08±1.82a 85.63±1.40b 84.38±1.30a 15.75±1.59a Intercropping 77.50±2.07b 59.17±2.71b 31.25±2.21b 97.50±1.81a 55.00±1.60b Monocropping 69.17±2.77c 57.08±1.27bc 30.42±1.21b 61.25±2.79c 48.75±2.38b Intercropping 51.67±1.48d 48.33±2.51d 41.25±2.43d Monocropping 54.58±2.22d 42.92±2.95d 43.75±2.09d N/2: ; N: ; 3N/2: P 0.05, N/2: deficient nitrogen; N: adequate nitrogen; 3N/2: excessive nitrogen. Different lowercase letters in the same line mean significant differences at 5% level. means that the target is not detected. 2.3 PLS-EDA,, PLS-EDA ( 1), PC1,,, PC2,, ;,,, PLS-EDA ( 1b), 在 PC1 上,,,,,,, PC2,, PC2, ( 2),,, (95 d), ;,, (60 d),,, (131 d), 2,,,,, (60 d),

5 6 : % 40.7%, ; (95 d), 39.5% 154.4%, 16.3% 37.4%; (131 d), 27.6% 188.4%, 34.9% 56.8%,, (60 d), 50.3% 66.3%, ; (95 d), 37.4% 94.4%, 69.7% 97.1%; (131 d), 52.8% 58.5%, 72.7% 94.5% 图 1 盆栽 (a) 和水培 (b) 试验不同氮水平下单间作蚕豆根系分泌大豆异黄酮 PLS-EDA 分析 Fig. 1 PLS-EDA analysis of soy isoflavone secreted by mono- and inter-cropped faba bean root under different N rates in pot experiment (a) and hydroponic experiment (b) IF:, MF: N/2: ; N: ; 3N/2: IF: intercropped faba bean; MF: monocropped faba bean; N/2: deficient nitrogen; N: adequate nitrogen; 3N/2: excessive nitrogen. 2,,,,, (60 d),, 64.6% 11.4%, (95 d),, 29.9% 9.9%; (131 d),, 18%,,, (60 d), 23.8% 14.1%, (95 d) 49.6% 82%, (131 d) 14% -0.3%,,,

6 图 2 盆栽 ( 上 ) 和水培 ( 下 ) 试验不同氮水平下单间作蚕豆根系分泌大豆异黄酮的差异 Fig. 2 Soy isoflavone secreted by mono- and inter-cropped faba bean root under different N rates in pot experiment (upper) and hydroponic experiment at different growth stages (lower) IF: ; MF: N/2: ; N: ; 3N/2: P 0.05 IF: intercropped faba bean; MF: monocropped faba bean; IW: intercropped wheat; MW: monocropped wheat; N/2: deficient nitrogen; N: adequate nitrogen; 3N/2: excessive nitrogen. Different lowercase letters mean significant differences at 5% level. 3 讨论, (50 kg hm 2 ), (Trifolium pratense) ;, [21-22],,, 95 d 131 d,,,,,, ;,,, [23],, -41,,,,,,, (60 d),, (131 d),, ;,,, (95 d), (60 d),,,,,, ; (95 d),, [24],, ; (131 d),,,,,,,

7 6 : 805,,,,,,,, 4 结论,,, ;,, (131 d), ( 60 d) ( 95 d), 22.5% 35.6% 28.8% 7.9%; ( 60 d) ( 95 d) ( 131 d), 44.2% 12.8%, 39.8% 46.0% 16.0% 27.0%;, 参考文献 References [1],,,. [J]., 2015, 52(4): DONG Y, DONG K, TANG L, et al. Relationship of free amino acids in root exudates with wilt disease (Fusarium oxysporum) of faba bean[j]. Acta Pedologica Sinica, 2015, 52(4): [2] FAN F L, ZHANG F S, SONG Y N, et al. Nitrogen fixation of faba bean (Vicia faba L.) interacting with a non-legume in two contrasting intercropping systems[j]. Plant and Soil, 2006, 283(1/2): [3],,. [J]., 2008, 14(5): LIU X Y, HE P, JIN J Y. Effect of potassium chloride on the exudation of sugars and phenolic acids by maize root and its relation to growth of stalk rot pathogen[j]. Plant Nutrition and Fertilizer Science, 2008, 14(5): [4] MAJ D, WIELBO J, MAREK-KOZACZUK M, et al. Response to flavonoids as a factor influencing competitiveness and symbiotic activity of Rhizobium leguminosarum[j]. Microbiological Research, 2010, 165(1): [5] LI L, LI S M, SUN J H, et al. Diversity enhances agricultural productivity via rhizosphere phosphorus facilitation on phosphorus-deficient soils[j]. Proceedings of the National Academy of Sciences of the United States of America, 2007, 104(27): [6] NOVÁK K, CHOVANEC P, ŠKRDLETA V, et al. Effect of exogenous flavonoids on nodulation of pea (Pisum sativum L.)[J]. Journal of Experimental Botany, 2002, 53(375): [7] LI B, KRUMBEIN A, NEUGART S, et al. Mixed cropping with maize combined with moderate UV-B radiations lead to enhanced flavonoid production and root growth in faba bean[j]. Journal of Plant Interactions, 2012, 7(4): [8] NEUMANN G, RÖMHELD V. The release of root exudates as affected by the plant physiological status[m]//pinton R, VARANINI Z, NANNIPIERI P. The Rhizosphere: Biochemistry and Organic Substances at the Soil-Plant Interface. 2nd ed. Boca Raton, FL, USA: CRC Press, 2007 [9] LANDINI S, GRAHAM M Y, GRAHAM T L. Lactofen induces isoflavone accumulation and glyceollin elicitation competency in soybean[j]. Phytochemistry, 2003, 62(6): [10] DELMONTE P, PERRY J, RADER J I. Determination of isoflavones in dietary supplements containing soy, red clover and kudzu: Extraction followed by basic or acid hydrolysis[j]. Journal of Chromatography A, 2006, 1107(1/2): [11],,,. [J]., 2009, (5): 1 4 GAO R H, ZHANG C H, ZHAO X H, et al. Research progress on soybean isoflavone[j]. Cereals & Oils, 2009, (5): 1 4 [12],,,. - [J]., 2015, 41(9): FENG X M, YANG Y, REN C Z, et al. Effects of legumes intercropping with oat on photosynthesis characteristics of and grain yield[j]. Acta Agronomica Sinica, 2015, 41(9): [13],,,. / [J]., 2015, 24(8): WU N, LIU X X, LIU J L, et al. Effect of intercropping potatoes with oats on the photosynthetic characteristics and yield of potato[j]. Acta Prataculturae Sinica, 2015, 24(8): [14],,. [J]., 2015, 21(5): ZHANG L C, TANG L, ZHENG Y. Phosphorus absorption of crops affected by root interaction in maize and soybean intercropping system[j]. Journal of Plant Nutrition and Fertilizer, 2015, 21(5): [15] HAUGGAARD-NIELSEN H, GOODING M, AMBUS P, et al. Pea-barley intercropping for efficient symbiotic N 2 -fixation, soil N acquisition and use of other nutrients in European organic cropping systems[j]. Field Crops Research, 2009, 113(1): [16] LI X P, MU Y H, CHENG Y B, et al. Effects of intercropping sugarcane and soybean on growth, rhizosphere soil microbes, nitrogen and phosphorus availability[j]. Acta Physiologiae Plantarum, 2013, 35(4): [17] WADDINGTON S R, MEKURIA M, SIZIBA S, et al.

8 Long-term yield sustainability and financial returns from grain legume-maize intercrops on a sandy soil in subhumid North Central Zimbabwe[J]. Experimental Agriculture, 2007, 43(4): [18] WANG D, YANG S M, TANG F, et al. Symbiosis specificity in the legume-rhizobial mutualism[j]. Cellular Microbiology, 2012, 14(3): [19],,. [J]., 2014, 25(6): XIAO J X, ZHENG Y, TANG L. Effect of wheat and faba bean intercropping on root exudation of low molecular weight organic acids[j]. Chinese Journal of Applied Ecology, 2014, 25(6): [20],,,. [J]., 2017, 53(6): LIU Y C, XIAO J X, TANG L, et al. Effects of nitrogen application rate on the naringenin exudation from intercropped faba bean s roots in different separation patterns[j]. Plant Physiology Journal, 2017, 53(6): [21],,. [J]., 2010, 19(7): LIU X L, DU W H, SONG C. Effects of nitrogen and phosphorus fertilization on isoflavone content in red clover[j]. Acta Agriculturae Boreali-Occidentalis Sinica, 2010, 19(7): [22],,,. [J]., 2006, (3): GU W Y, YU F, CHEN Y, et al. Study on the distribution of isoflavone in Trifolium pratense at different growth stages[j]. Grassland and Turf, 2006, (3): [23],,,. / [J]., 2008, 14(6): LU G L, TANG L, CHU Y O, et al. Effect of nitrogen levels on the changes of phenol and flavonoid contents under rice monocropping and intercropping system[j]. Plant Nutrition and Fertilizer Science, 2008, 14(6): [24] LI B, LI Y Y, WU H M, et al. Root exudates drive interspecific facilitation by enhancing nodulation and N 2 fixation[j]. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(23):

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