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1 Chinese Journal of Eco-Agriculture, Jan. 2017, 25(1): DOI: /j.cnki.cjea ,,,,,. [J]., 2017, 25(1): Chen L, Dong K, Yang Z X, Dong Y, Tang L, Zheng Y. Alleviation mechanism of intercropping with wheat for faba bean autotoxicity under benzoic acid stress[j]. Chinese Journal of Eco-Agriculture, 2017, 25(1): * 陈玲 1, 董坤 2, 杨智仙 3, 董艳 1**, 汤利 1 1,4, 郑毅 ( ; ; ; ) : 苯甲酸是引起蚕豆连作障碍的主要自毒物质之一本文采用水培试验, 研究了不同浓度苯甲酸 [C 0 (0 mg L 1 ) C 1 (50 mg L 1 ) C 2 (100 mg L 1 ) 和 C 3 (200 mg L 1 )] 处理对与小麦间作的蚕豆幼苗生长和枯萎病发生的影响, 从生理抗性角度探讨小麦与蚕豆间作对缓解苯甲酸自毒效应的机制, 为合理利用间作缓解连作障碍, 实现农业可持续发展提供科学依据结果表明 : 与 C 0 处理相比, 不同浓度苯甲酸处理均显著抑制了蚕豆幼苗的生长, 并且随处理浓度升高, 抑制效应增强 ; 同时显著提高了蚕豆枯萎病发病率和病情指数 ; 蚕豆根系和叶片的 MDA 含量显著提高, 但抗氧化酶 (POD 和 CAT) 活性和病程相关蛋白 (β-1,3- 葡聚糖酶和几丁质酶 ) 活性均随苯甲酸处理浓度升高而降低表明不同浓度苯甲酸处理均显著抑制了蚕豆的生长, 降低蚕豆的生理抗性而促进枯萎病发生与单作蚕豆相比, 蚕豆与小麦间作显著提高了苯甲酸胁迫下蚕豆的地上部干重 (17.0%~47.1%), 降低了发病率 (11.1%~25.0%) 和病情指数 (20.0%~42.1%); 蚕豆根系和叶片中 POD 活性分别提高 12.9%~16.9% 和 9.3%~24.9%, CAT 活性分别提高 10.3%~54.0% 和 6.6%~20.5%, 蚕豆根系的 β-1,3- 葡聚糖酶和几丁质酶活性分别提高 4.7%~13.1% 和 6.7%~15.8%, MDA 含量分别降低 19.5%~25.4% 和 20.5%~29.9% C 2 处理下间作提高抗氧化酶和病程相关蛋白活性的效果最好, 抗病效果最佳表明小麦与蚕豆间作通过提高蚕豆的生理抗性而减轻苯甲酸引起的枯萎病危害, 促进蚕豆生长, 是缓解苯甲酸自毒效应的有效措施 : 苯甲酸 ; 蚕豆 ; 连作障碍 ; 间作 ; 抗氧化酶 ; 病程相关蛋白 ; 枯萎病 : S : A : (2017) Alleviation mechanism of intercropping with wheat for faba bean autotoxicity under benzoic acid stress * CHEN Ling 1, DONG Kun 2, YANG Zhixian 3, DONG Yan 1**, TANG Li 1, ZHENG Yi 1,4 (1. College of Resources and Environment, Yunnan Agricultural University, Kunming , China; 2. College of Food Science and Technology, Yunnan Agricultural University, Kunming , China; 3. Yunnan University, Kunming , China; 4. Southwest Forestry University, Kunming , China) Abstract: Continuous cropping of faba bean causes serious growth obstacle, and benzoic acid is one of the main autotoxins in faba bean continuous cropping. Faba bean-wheat intercropping is a general planting pattern in Southwest China. To investigate the effect of benzoic acid on faba bean seedling growth and Fusarium wilt occurrence, and to explore the effect and alleviation * ( , , , )(2015FA022) ** :, dongyanyx@163.com, @qq.com : : * The study was supported by the National Natural Science Foundation of China ( , , , ) and the Key Science and Technology Project of Yunnan Province of China (2015FA022). ** Corresponding author, dongyanyx@163.com Received Jun. 15, 2016; accepted Sep. 6, 2016

2 mechanism of intercropping with wheat on faba bean autotoxicity, a hydroponic experiment of faba bean-wheat intercropping with benzoic acid application was conducted in the greenhouse of Yunnan Agricultural University. Four benzoic acid rates were set, which were C 0 (0 mg L 1 ), C 1 (50 mg L 1 ), C 2 (100 mg L 1 ) and C 3 (200 mg L 1 ), with two planting patterns, faba bean monocropping and faba bean and wheat intercropping. The growth, enzymes activities and occurrence of faba bean Fusarium wilt were investigated. The results showed that in comparison with C 0 treatment, seedling growth was inhibited with increasing benzoic acid concentration. Benzoic acid increased the incidence and disease index of monocropped faba bean Fusarium wilt, significantly increased root and leaf MDA contents, and decreased the activities of antioxidant enzyme (POD and CAT) and pathogenesis-related proteins (β-1, 3-glacanase and chitinase) with increasing application rates. Comparison with monocropped faba bean, intercropped faba bean increased shoot dry weight by 17.0% 47.1%, reduced disease incidence and disease index by 11.1% 25.0% and 20.0% 42.1%, respectively, under benzoic acid application. Furthermore, the peroxidase (POD) activity increased by 12.9% 16.9% in root and by 9.3% 24.9% in leaf; and the catalase (CAT) activity increased by 10.3% 54.0% in root and by 6.6% 20.5% in leaf under intercropped condition. Compared with monocropping of faba bean, intercropping with wheat increased faba bean β-1,3-glacanase and chitinase activities in root by 4.7% 13.1% and 6.7% 15.8%, while decreased malondialdehyde (MDA) contents in both root and leaf significantly by 19.5% 25.4% and 20.5% 29.9%, respectively, with benzoic acid application. Under C 2 (100 mg L 1 ) treatment of benzoic acid stress, faba bean-wheat intercropping system showed the best effect of improving pathogenesis-related proteins and antioxidative enzymes activity. Intercropping of faba bean-wheat significantly increased physiological resistance to Fusarium oxysporum f. fabae and significantly decreased the incidence of Fusarium wilt. Therefore, it alleviated the damage of Fusarium wilt caused by benzoic acid, and promoted growth of faba bean. Thus, intercropping with wheat was an effective method to alleviate faba bean autotoxicity caused by benzoic acid. The results also provided an experimental evidence for developing an ecologicaleconomic and effective approach to control soil-borne diseases caused by continuous crop cultivation. Keywords: Benzoic acid; Faba bean; Continuous cropping obstacle; Intercropping; Antioxidative enzyme; Pathogenesisrelated protein; Fusarium wilt, [1],,, [1],, [2-3] [3-4], (Cunninghamia lanceolata) (Cucumis sativus) (Solanum melongena) (Solanum lycopersicum), (Citrullus lanatus) (Colocasia esculenta) (Panax ginseng) [1,5], [6-7], (Vicia faba),, ; 7 ( ), [8],,,, [5], (Triticum aestivum),,, [9],,, [10-11],,,, ;,,

3 1 : 97 1 材料与方法 , , ( ) [Fusarium oxysporum f. fabae, (FOF)], PDA, 28 7 d, 1.2, A, 4, C 0 (0 mg L 1, ), C 1 (50 mg L 1 ), C 2 (100 mg L 1 ), C 3 (200 mg L 1 ) B, (MF)(IF), 8, 3, 24 3, 3 9 ( 1), 24 h Fig. 1 图 1 蚕豆单作和小麦与蚕豆间作种植示意图 Schematic diagrams of faba bean monocropping and intercropping with wheat h, 25 Hoagland, 4~6, 2 L, (3 ) 2 d, cfu ml d, 3 5 [11], (%)=(/ ) 100 (1) = ( )/( ) 100 (2) (%)= ( )/ 100 (3) d ( ), 3,, d ( ), 3, (POD) (CAT) (MDA), [12] d ( ), 3, ( ), 1 g 1 mg N- β-1,3- β-1,3- ( ), 1 β-1,3- g 1 mg 1.6 Microsoft Excel 2007, SPSS 20.0, (LSD) 2 结果与分析 ,,, ( ) C 0, C 1 C 2 C % 35.4% 41.5%;

4 Table 1 表 1 苯甲酸胁迫下与小麦间作对蚕豆幼苗生长的影响 Effect of intercropping with wheat on faba bean growth parameters under benzoic acid stress C 0 C 1 C 2 C 3 Growth parameter MF IF MF IF MF IF MF IF Leaf number per plant 21.67±0.57b 24.00±1.00a 14.67±1.15de 16.23±0.58c 14.00±1.00def 15.33±0.58cd 12.67±1.15f 13.67±0.58ef Max leaf length (cm) 8.53±0.38b 9.37±0.12a 7.20±0.20cd 7.80±0.36c 5.77±0.47e 6.12±0.30d 4.73±0.49f 4.90±0.35e Max leaf width (cm) 5.43±0.51a 5.53±0.12a 4.07±0.25b 4.47±0.35b 3.71±0.38c 3.97±0.21b 2.70±0.26d 2.80±0.38cd Plant height (cm) 39.23±0.49a 37.87±1.44a 32.17±1.06b 30.47±0.61c 25.80±1.04d 23.90±0.56e 21.10±0.11f 19.37±0.76f Main root length (cm) 18.33±1.45bc 19.83±0.97a 18.13±0.42bcd 19.23±0.38ab 16.83±0.76d 17.40±0.17cd 13.13±0.45e 13.43±0.21e Shoot dry weight (g) 2.23±0.05b 3.28±0.07a 1.52±0.12c 2.14±0.07b 0.87±0.35e 1.14±0.07d 0.53±0.01f 0.62±0.15e Root dry weight (g) 0.54±0.03b 0.86±0.06d 0.36±0.01a 0.56±0.05c 0.19±0.05c 0.32±0.01e 0.09±0.01b 0.16±0.02d Root-shoot ratio 0.24±0.02a 0.26±0.01ab 0.24±0.01a 0.26±0.03a 0.22±0.04ab 0.28±0.01c 0.17±0.01a 0.26±0.01c MF: ; IF: C 0 C 1 C 2 C 3, 0 mg L 1 50 mg L mg L mg L 1 (P<0.05) MF: monocropped faba bean; IF: intercropped faba bean. C 0, C 1, C 2 and C 3 are treatments of benzoic acid stress of 0 mg L 1, 50 mg L 1, 100 mg L 1 and 200 mg L 1. Different letters in the same row mean significant differences at 0.05 level. 15.6% 32.4% 44.5%; 25.0% 31.7% 50.3%; 18.0% 34.2% 46.2%; 31.8% 61.0% 76.2%; 33.3% 64.8% 83.3% C 0, C 2 C % 28.4%, (C 0 ), 10.8% 12.2% 8.2% 47.1% 59.3%; C 1, 10.6% 40.8% 55.6%; C 2, 6.1% 7.0% 31.0% 68.4%; C 3, 3.6% 17.0% 77.8%, ( 1) 2.2 2,, C 0, C 1 C 2 C % 50.0% 50.0%, 25.0% 137.5% 362.4% C 1 C 2,, 25.0% 11.1%; C 0 C 3 C 0 C 1 C 2 C 3, 25.0% 20.0% 42.1% 21.6%, C 2, 20.0%~42.1%, C 2, 42.1% Table 2 表 2 苯甲酸胁迫下间作对蚕豆枯萎病发病率和病情指数的影响 Effect of intercropping with wheat on occurrence and index of faba bean Fusarium wilt under benzoic acid stress Incidence (%) Disease index (%) MF IF MF IF Relative control effect (%) C ±3.61c 66.67±3.33c 17.78±2.04ef 13.33±1.67f 25.03±2.00b C ±1.92b 66.67±1.67c 22.22±3.85de 17.78±0.96ef 19.98±2.32c C ±0.00a 88.89±8.38b 42.22±1.92c 24.44±3.85d 42.11±2.31a C ±0.00a ±0.00a 82.22±3.85a 64.44±1.95b 21.62±4.94bc MF: ; IF: C 0 C 1 C 2 C 3, 0 mg L 1 50 mg L mg L mg L 1 (P<0.05) MF: monocropped faba bean; IF: intercropped faba bean. C 0, C 1, C 2 and C 3 are treatments of benzoic acid stress of 0 mg L 1, 50 mg L 1, 100 mg L 1 and 200 mg L 1. Different letters in the same row mean significant differences at 0.05 level. 2.3 POD CAT POD 2, POD, C 0, C 1 POD C 0, ; C 2 C 3 POD

5 1 : 99 图 2 苯甲酸胁迫下与小麦间作对蚕豆根系和叶片中 POD 和 CAT 活性的影响 Fig. 2 Effect of intercropping with wheat on POD and CAT activities in roots and leaves of faba bean under benzoic acid stress MF: ; IF: C 0 C 1 C 2 C 3, 0 mg L 1 50 mg L mg L mg L 1 (P<0.05) MF: monocropped faba bean; IF: intercropped faba bean. C 0, C 1, C 2 and C 3 are treatments of benzoic acid stress of 0 mg L 1, 50 mg L 1, 100 mg L 1 and 200 mg L 1. Different letters for the same organ of faba bean mean significant differences at 0.05 level. 15.7% 31.4%; POD 21.3% 38.7%,, POD C 0 C 1 C 2 C 3, POD 12.9% 15.1% 16.9% 14.6%, POD, C 2 POD 24.9%( 2) C 2 POD CAT 2, C 0, C 1 CAT C 0, ; C 2 C 3 CAT 37.7% 42.8%; CAT 28.4% 44.8%, CAT C 0 C 1 C 2 C 3, CAT 10.3% 22.6% 54.0% 21.7%, CAT 6.6% 9.2% 11.5% 20.5%( 2) 2.4 MDA MDA 3, C 0, C 1 MDA C 0, ; C 2 C 3 MDA 28.9% 42.6%; MDA 16.4% 45.0%, MDA, C 0 C 1 C 2 C 3 图 3 苯甲酸胁迫下与小麦间作对蚕豆根系和叶片中 MDA 含量的影响 Fig. 3 Effect of intercropping with wheat on MDA content in roots and leaves of faba bean under benzoic acid stress MF: ; IF: C 0 C 1 C 2 C 3, 0 mg L 1 50 mg L mg L mg L 1 (P<0.05) MF: monocropped faba bean; IF: intercropped faba bean. C 0, C 1, C 2 and C 3 are treatments of benzoic acid stress of 0 mg L 1, 50 mg L 1, 100 mg L 1 and 200 mg L 1. Different letters for the same organ of faba bean mean significant differences at 0.05 level., MDA 25.4% 22.6% 20.0% 19.5%, MDA 20.5% 29.9% 24.5% 26.5% 2.5 β-1,3 4, C 0, C 1 β-1,3

100 2017 25 C 0, ; C 2 C 3 β-1,3 17.4% 38.7% C 0 C 1 C 2 C 3, β-1,3 4.7% 7.9% 13.1% 10.3%, C 2 β-1,3 ( 4) 图 4 苯甲酸胁迫下与小麦间作对蚕豆根系 β-1,3 葡聚糖酶和几丁质酶活性的影响 Fig.

6 C 0, ; C 2 C 3 β-1,3 17.4% 38.7% C 0 C 1 C 2 C 3, β-1,3 4.7% 7.9% 13.1% 10.3%, C 2 β-1,3 ( 4) 图 4 苯甲酸胁迫下与小麦间作对蚕豆根系 β-1,3 葡聚糖酶和几丁质酶活性的影响 Fig. 4 Effect of intercropping with wheat on β-1,3-glacanase and chitinase activities in roots of faba bean under benzoic acid stress MF: ; IF: C 0 C 1 C 2 C 3, 0 mg L 1 50 mg L mg L mg L 1 (P<0.05) MF: monocropped faba bean; IF: intercropped faba bean. C 0, C 1, C 2 and C 3 are treatments of benzoic acid stress of 0 mg L 1, 50 mg L 1, 100 mg L 1 and 200 mg L 1. Different letters mean significant differences at 0.05 level., C 0, C 1 C 0, ; C 2 C % 39.4%(P<0.05),,, 6.7% 7.8% 15.8% 10.5%, C 2 3 讨论与结论 3.1,,, (Cucumis sativus) (Citrullus lanatus) (Solanum melongena), ( ) [13] ( ) [14],,, ( ),,,,, [15] (Fragaria ananassa), [16], [17] ; [18],, (Cucumis melo) [13], [19-20],, [21] (Oryza sativa), [22],,, (Artractylodes lancea), [23] ; [5],, 20.0%~42.1%,

7 1 : 101 C 2, 3.2,, CAT,, [22], [24-25], POD CAT [14,26] CAT, [15],, CAT,, (CAT),,, POD,,,, POD,, MDA,, MDA,,,,, [27] MDA,, [15],, MDA,,,,, MDA, MDA,,,, 5 d 10 d 15 d MDA, MDA [28], POD CAT,, MDA,,,, 3.3,,,, [28],,,,, ; β-1,3-, [28],, β-1,3- [29],, C 1 β-1,3-, ; C 1, β-1,3-,,, β-1,3- [28] β-1,3-, [29],, β-1,3-, β-1,3-,,,

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