番茄碳库强度对其与伴生分蘖洋葱和AMF的关系及钾吸收的影响

doi:10.16420/j.issn.0513-353x.2023-0413

摘要/Abstract

摘要：

以番茄（Solanum lycopersicum L.）和分蘖洋葱（Allium cepa L. var. aggregatum G. Don）为试验材料，通过分蘖洋葱伴生番茄（四叶一心时定植）盆栽试验，研究番茄遮荫（25%、50%）、去叶（25%、50%）和调整定植苗大小（在番茄子叶展平、番茄两叶一心时定植）3种方式对番茄碳库强度的影响。遮荫和去叶处理均降低了番茄的生长且促进了分蘖洋葱生长，而缩小番茄和分蘖洋葱间的个体差异后，均能促进两者植株生长；削减番茄碳库强度后，番茄与分蘖洋葱的丛枝菌根真菌的丛枝丰度和菌丝侵染率减少，但囊泡丰度增加；番茄去叶和较小定植苗处理后降低了番茄叶片和分蘖洋葱叶片、根系钾含量，提高了番茄根系钾含量，番茄遮荫25%处理提高了分蘖洋葱根系、叶片钾含量，遮荫50%处理提高了番茄叶片、根系钾含量；番茄遮荫和较小定植苗处理对分蘖洋葱土壤速效钾含量没有影响，而去叶和较小定植苗处理使番茄土壤速效钾含量降低并且削减番茄碳库强度，使两者植株可溶性糖含量降低。因此，削减番茄碳库强度减弱了分蘖洋葱对番茄促生作用，且抑制了丛枝菌根真菌在分蘖洋葱伴生番茄系统中的作用效果。

关键词: 番茄, 伴生, 分蘖洋葱, 碳库强度, 丛枝菌根真菌

Abstract:

A pot experiment was conducted using tomato（Solanum lycopersicum L.）and potato-onion （Allium cepa L. var. aggregatum G. Don）in itercropping system to study the effects of shading（25%，50%）defoliation（25%，50%）and adjusting the size of planted tomato seedlings（tomato seedlings with two cotyledons or with two leaves planted with potato-onion）on carbon pool strength of tomato. Shading and defoliation treatments were found to decrease tomato growth while enhancing the growth of potato-onions. Interestingly，reducing the individual differences between tomato and potato-onion plants promoted the growth of both two plants. Reduction of tomato carbon pool strength reduced abundance of the arbuscular and hypha infection rate of tomato and potato-onion，but increased the abundance of vesicular. The potassium concentration in leaves and roots of potato-onion and the potassium concentration in leaves tomato were decreased，but increasing in roots of tomato instead after defoliation and adjusting the size of plant seedlings. Moreover，the rising concentration of potassium in leaves and roots of the tomato and potato-onion were respectively observed in shading 50% and 25% treatment. Shading and transplanting smaller tomato seedlings did not affect the available potassium content in the soil for potato-onions. Conversely，defoliation and transplanting smaller tomato seedlings reduced the available potassium in the soil for tomato，and weakened the tomato carbon pool strength can lead to a decrease in soluble sugar content in both plants. Overall，the reduction of carbon pool strength in tomato decreased to promoting effects of potato-onion on tomato plant growth，and reduced the role of arbuscular mycorrhizal fungi in the intercropping benifits between potato-onion and tomato system.

Key words: tomato, companion, potato-onion, carbon pool strength, arbuscular mycorrhizal fungi

陆建增, 周丽颜, 黄馨怡, 吴凤芝, 高丹美. 番茄碳库强度对其与伴生分蘖洋葱和AMF的关系及钾吸收的影响[J]. 园艺学报, 2024, 51(11): 2620-2632.

LU Jianzeng, ZHOU Liyan, HUANG Xinyi, WU Fengzhi, GAO Danmei. Effects of Carbon Pool Strength on Plant Growth and Potassium Uptake of Tomato Intercropped with Potato-Onion[J]. Acta Horticulturae Sinica, 2024, 51(11): 2620-2632.

图/表 5

图1 盆栽中番茄遮荫、去叶和定植苗大小对番茄和伴生分蘖洋葱植株生长的影响 CK：正常对照。采用Student’s t检验进行处理间的平均比较，*表示处理与对照间差异显著（P < 0.05）。

Fig. 1 Effects of shading，defoliation and size adjustment on growth of tomato intercropped with potato-onion under pot experiment CK：Normal control. Student’s t-test was used for the mean comparison between treatments. * indicated significant differences between treatments and the control（P < 0.05）.

图2 盆栽中番茄遮荫、去叶和定植苗大小对番茄和伴生分蘖洋葱根系菌根侵染率的影响 CK：正常对照。采用Student’s t检验进行处理间的平均比较，*表示处理与对照间差异显著（P < 0.05）。

Fig. 2 Effects of shading，defoliation and size adjustment on mycorrhizal infection rate in tomato intercropped with potato-onion under pot experiment CK：Normal control. Student’s t-test was used for the mean comparison between treatments. * indicated significant differences between treatments and the control（P < 0.05）.

图3 盆栽中番茄遮荫、去叶和定植苗大小对番茄和伴生分蘖洋葱叶片及根系钾含量的影响 CK：正常对照。采用Student’s t检验进行处理间的平均比较，*表示处理与对照间差异显著（P < 0.05）。

Fig. 3 Effects of shading，defoliation and size adjustment on potassium content in leaves and roots of tomato intercropped with potato-onion under pot experiment CK：Normal control. Student’s t-test was used for the mean comparison between treatments. * indicated significant differences between treatments and the control（P < 0.05）.

图4 盆栽中番茄遮荫、去叶和定植苗大小对番茄和伴生分蘖洋葱土壤速效钾含量的影响 CK：正常对照。采用Student’s t检验进行处理间的平均比较，*表示处理与对照间差异显著（P < 0.05）。

Fig. 4 Effects of shading，defoliation and size adjustment on available potassium of soil in tomato intercropped with potato-onion under pot experiment CK：Normal control. Student’s t-test was used for the mean comparison between treatments. * different lowercase letters indicated significant differences between treatments and the control（P < 0.05）.

图5 盆栽中番茄遮荫、去叶和定植苗大小对番茄和伴生分蘖洋葱植株可溶性糖含量的影响 CK：正常对照。采用Student’s t检验进行处理间的平均比较，*表示处理与对照间差异显著（P < 0.05）。

Fig. 5 Effects of shading，defoliation and size adjustment on soluble sugar of plant in tomato intercropped with potato-onion under pot experiment CK：Normal control. Student’s t-test was used for the mean comparison between treatments. * indicated significant differences between treatments and the control（P < 0.05）.

参考文献 48

[1]	Awaydul A, Zhu W, Yuan Y, Xiao J, Hu H, Chen X, Koide R T, Cheng L. 2019. Common mycorrhizal networks influence the distribution of mineral nutrients between an invasive plant,Solidago canadensis,and a native plant,Kummerowa striata. Mycorrhiza, 29 (1):29-38.
[2]	Bao Shidan. 2000. Soil and agricultural chemistryanalysis. 3rd ed. Beijing:China Agriculture Press:263-268. (in Chinese)
	鲍士旦. 2000. 土壤农化分析. 3版. 北京:中国农业出版社:263-268.
[3]	Baslam M, Esteban R, García-Plazaola J I, Goicoechea N. 2013. Effectiveness of arbuscular mycorrhizal fungi(AMF)for inducing the accumulation of major carotenoids,chlorophylls and tocopherol in green and red leaf lettuces. Applied Microbiology and Biotechnology, 97:3119-3128.
[4]	Bi Y, Zhou H. 2021. Changes in peanut canopy structure and photosynthetic characteristics induced by an arbuscular mycorrhizal fungus in a nutrient-poor environment. Scientific Reports, 11 (1):1-10.
[5]	Cao Benfu, Jiang Haixia, Liu Li, Lu Yingang, Wang Maosheng. 2021. Research progress on mechanism of arbuscular common mycorrhizal networks in plant-plant interactions. The Journal of Applied Ecology, 32 (9):3385-3396. (in Chinese)
	曹本福, 姜海霞, 刘丽, 陆引罡, 王茂胜. 2021. 丛枝菌根菌丝网络在植物互作中的作用机制研究进展. 应用生态学报, 32 (9):3385-3396. doi: 10.13287/j.1001-9332.202111.032
[6]	Cui M, Caldwell M M. 1997. Shading reduces exploitation of soil nitrate and phosphate by Agropyron desertorum and Artemisia tridentata from soils with patchy and uniform nutrient distributions. Oecologia, 109:177-183. doi: 10.1007/s004420050072 pmid: 28307168
[7]	Dietze M C, Sala A, Carbone M S, Czimczik C I, Mantooth J A, Richardson A D, Vargas R. 2014. Nonstructural carbon in woody plants. Annual Review of Plant Biology, 65 (1):667-687.
[8]	Dong Sangjie, Ge Shibei, Li Lan, He Liqun, Fan Feijun, Qi Zhenyu, Yu Jingquan, Zhou Yanhong. 2022. Effects of supplemental lighting on growth,root colonization by arbuscular mycorrhizal fungi and phosphorus uptake in pepper seedlings. Acta Horticulturae Sinica, 49 (8):1699-1712. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0604
	董桑婕, 葛诗蓓, 李岚, 贺丽群, 范飞军, 齐振宇, 喻景权, 周艳虹. 2022. 不同光质补光对辣椒幼苗生长、丛枝菌根共生和磷吸收的影响. 园艺学报, 49 (8):1699-1712. doi: 10.16420/j.issn.0513-353x.2021-0604
[9]	Drigo B, Pijl A S, Duyts H, Kielak A M, Gamper H A, Houtekamer M J, Boschker H T, Bodelier P L, Whiteley A S, van Veen J A, Kowalchuk G A. 2010. Shifting carbon flow from roots into associated microbial communities in response to elevated atmospheric CO₂. Proceedings of the National Academy of Sciences, 107 (24):10938-10942.
[10]	Fellbaum C R, Mensah J A, Cloos A J, Strahan G E, Pfeffer P E, Kiers E T, Bücking H. 2014. Fungal nutrient allocation in common mycorrhizal networks is regulated by the carbon source strength of individual host plants. New Phytologist, 203 (2):646-656. doi: 10.1111/nph.12827 pmid: 24787049
[11]	Fellbaum C R, Mensah J A, Pfeffer P E, Kiers E T, Bücking H. 2012. The role of carbon in fungal nutrient uptake and transport:implications for resource exchange in the arbuscular mycorrhizal symbiosis. Plant Signaling & Behavior, 7 (11):1509-1512.
[12]	Gao D, Pan X, Khashi U, Rahman M, Zhou X, Wu F. 2021. Common mycorrhizal networks benefit to the asymmetric interspecific facilitation via K exchange in an agricultural intercropping system. Biology and Fertility of Soils, 57:959-971.
[13]	Ge Shibei, Jiang Xiaochun, Wang Lingyu, Yu Jingquan, Zhou Yanhong. 2020. Recent advances in the role and mechanism of arbuscular mycorrhiza-induced improvement of abiotic stress tolerance in horticultural plants. Acta Horticulturae Sinica, 47 (9):1752-1776. (in Chinese)
	葛诗蓓, 姜小春, 王羚羽, 喻景权, 周艳虹. 2020. 园艺植物丛枝菌根抗非生物胁迫的作用机制研究进展. 园艺学报, 47 (9):1752-1776.
[14]	Gong Xiaoya, Li Xian, Zhou Xingang, Wu Fengzhi. 2024. Effect of rhizosphere microorganisms induced by potato-onion on tomato root-knot nematode disease. Acta Horticulturae Sinica, 51 (8):1913-1926. (in Chinese) doi: 10.16420/j.issn.0513-353x.2023-0306
	龚小雅, 李贤, 周新刚, 吴凤芝. 2024. 分蘖洋葱伴生番茄诱导的根际微生物对根结线虫病的影响. 园艺学报, 51 (8):1913-1926.
[15]	Hammer E C, Nasr H, Wallander H. 2011. Effects of different organic materials and mineral nutrients on arbuscular mycorrhizal fungal growth in a Mediterranean saline dryland. Soil Biology and Biochemistry, 43 (11):2332-2337.
[16]	Hu Hailing, Ma Yuwen, Geng Heyang, Chen Xiaoxuan, Shi Congcong, Wang Yingnan, Wang Jinghong, Lin Jixiang. 2022. Progress in AMF enhancing plant stress tolerance using omics techniques. Journal of Plant Nutrition and Fertitizer, 28 (10):1928-1936. (in Chinese)
	胡海玲, 马钰雯, 耿赫阳, 陈晓璇, 石聪聪, 王英男, 王竞红, 蔺吉祥. 2022. 丛枝菌根真菌AMF提高植物抗逆性的组学技术研究进展. 植物营养与肥料学报, 28 (10):1928-1936.
[17]	Huang J, Liu W, Yang S, Yang L, Peng Z, Deng M, Liu L. 2021. Plant carbon inputs through shoot,root,and mycorrhizal pathways affect soil organic carbon turnover differently. Soil Biology and Biochemistry, 160:108322.
[18]	Kering M K, Butler T J, Biermacher J T, Mosali J, Guretzky J A. 2013. Effect of potassium and nitrogen fertilizer on switchgrass productivity and nutrient removal rates under two harvest systems on a low potassium soil. Bioenergy Research, 6:329-335.
[19]	Lang Linjie, Yang Chi. 1997. The effects of shading and defoliation on translocation of carbon resources between clonal ramets of Leymus chinensis. Acta Scientiarum Naturalium Universitatis NeiMongol, 28 (3):430-434. (in Chinese)
	郎林杰, 杨持. 1997. 遮光和去叶处理对羊草(Leymus chinensis)无性系分株间碳物质转移的影响. 内蒙古大学学报(自然科学版), 28 (3):430-434.
[20]	Lekberg Y, Hammer E C, Olsson P A. 2010. Plants as resource islands and storage units-adopting the mycocentric view of arbuscular mycorrhizal networks. FEMS Microbiology Ecology, 74 (2):336-345. doi: 10.1111/j.1574-6941.2010.00956.x pmid: 20722732
[21]	Li Min,Liu Runjin,Li Xiaolin. 2004. Influences of arbuscular mycorrhizal fungi on growth and Fusarium-wilt disease of watermelon in field. Acta Phytopathologica Sinica, 34 (5):472-473. (in Chinese)
	李敏, 刘润进, 李晓林. 2004. 大田条件下丛枝菌根真菌对西瓜生长和枯萎病的影响. 植物病理学报, 34 (5):472-473.
[22]	Li Wei,Cheng Zhihui,Meng Huanwen,Zhou Jing,Liang Jing,Liu Xuejiao. 2012. Effect of rotating different vegetables on micro-biomass and enzyme in tomato continuous cropped substrate and afterculture tomato under plastic tunnel cultivation. Acta Horticulturae Sinica, 39 (1):73-80. (in Chinese)
	李威, 程智慧, 孟焕文, 周静, 梁静, 刘雪娇. 2012. 轮作不同蔬菜对大棚番茄连作基质中微生物与酶及后茬番茄的影响. 园艺学报, 39 (1):73-80.
[23]	Liang Shengmin, Zhang Fei, Wu Qiangsheng. 2023. Arbuscular mycorrhizal fungi improve drought tolerance of trifoliate orange seedlings by regulating root polyamines. Acta Horticulturae Sinica, 50 (12):2680-2688. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-1036
	梁圣敏, 张菲, 吴强盛. 2023. 丛枝菌根真菌通过调节枳根系多胺提高抗旱性. 园艺学报, 50 (12):2680-2688. doi: 10.16420/j.issn.0513-353x.2022-1036
[24]	Loch-Temzelides T. 2021. Walrasian equilibrium behavior in nature. Proceedings of the National Academy of Sciences, 118 (27):1-6.
[25]	Ma Junqing, Hou Ning, Sun Chenyu, Yang Yisen, Qin Shengfeng, Wang Yong, Liu Lu, Liao Honglin, Huang Jinghua. 2022. Effects of different hosts on propagation of arbuscular mycorrhizal fungi. Chinese Agricultural Science Bulletin,(1):7-14. (in Chinese) doi: 10.11924/j.issn.1000-6850.casb2021-0050
	马俊卿, 侯宁, 孙晨瑜, 杨怡森, 覃圣峰, 王勇, 刘璐, 廖虹霖, 黄京华. 2022. 宿主不同对丛枝菌根真菌扩繁效应的影响. 中国农学通报,(1):7-14. doi: 10.11924/j.issn.1000-6850.casb2021-0050
[26]	McGonigle T P, Miller M H, Evans D G, Fairchild G L, Swan J A. 1990. A new method which gives an objective measure of colonization of roots by vesicular-arbuscular mycorrhizal fungi. New Phytologist, 115 (3):495-501. doi: 10.1111/j.1469-8137.1990.tb00476.x pmid: 33874272
[27]	Merrild M P, Ambus P, Rosendahl S, Jakobsen I. 2013. Common arbuscular mycorrhizal networks amplify competition for phosphorus between seedlings and established plants. New Phytologist, 200 (1):229-240. doi: 10.1111/nph.12351 pmid: 23738787
[28]	Phillips J M, Hyman D S. 1970. Improved procedures for clearing roots and staining parasitic and vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection. Transactions of the British Mycological Society, 55 (1):158-161.
[29]	Pietikäinen A, Kytöviita M M. 2007. Defoliation changes mycorrhizal benefit and competitive interactions between seedlings and adult plants. Journal of Ecology, 95 (4):639-647.
[30]	Pietikäinen A, Kytöviita M M, Husband R, Young J P W. 2007. Diversity and persistence of arbuscular mycorrhizas in a low‐arctic meadow habitat. New Phytologist, 176 (3):691-698. doi: 10.1111/j.1469-8137.2007.02209.x pmid: 17725554
[31]	Qiao X, Bei S, Li C, Dong Y, Li H, Christie P, Zhang J. 2015. Enhancement of faba bean competitive ability by arbuscular mycorrhizal fungi is highly correlated with dynamic nutrient acquisition by competing wheat. Scientific Reports, 5 (1):8122.
[32]	Qu Minghua, Yu Yuanchun, Li Sheng, Zhang Jinchi. 2019. Advances in research on activation of mineral nutrients by arbuscular mycorrhizal fungi. Journal of Zhejiang A & F University, 36 (2):394-405. (in Chinese)
	屈明华, 俞元春, 李生, 张金池. 2019. 丛枝菌根真菌对矿质养分活化作用研究进展. 浙江农林大学学报, 36 (2):394-405.
[33]	Rosa M, Prado C, Podazza G, Interdonato R, González J A, Hilal M, Prado F E. 2009. Soluble sugars:metabolism,sensing and abiotic stress:a complex network in the life of plants. Plant Signaling & Behavior, 4 (5):388-393.
[34]	Scheloske S, Maetz M, Schneider T, Hildebrandt U, Bothe H, Povh B. 2004. Element distribution in mycorrhizal and nonmycorrhizal roots of the halophyte Aster tripolium determined by proton induced X-ray emission. Protoplasma, 223:183-189. pmid: 15221523
[35]	Sun M F, Yuan D, Hu X C, Zhang D, Li Y Y. 2020. Effects of mycorrhizal fungi on plant growth,nutrient absorption and phytohormones levels in tea under shading condition. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 48 (4):2006-2020.
[36]	Tan Deshui, Jin Jiyun, Huang Shaowen. 2008. Effect of long-term K fertilizer application and returning wheat straw to soil on crop yield and soil K under different planting systems in northwestern China. Plant Nutrition and Fertilizer Science, 14 (5):886-893. (in Chinese)
	谭德水, 金继运, 黄绍文. 2008. 长期施钾与秸秆还田对西北区不同种植制度下作物产量及土壤钾素的影响. 植物营养与肥料学报, 14 (5):886-893.
[37]	van der Heijden M G, Horton T R. 2009. Socialism in soil? The importance of mycorrhizal fungal networks for facilitation in natural ecosystems. Journal of Ecology, 97 (6):1139-1150.
[38]	van der Heijden M G A,Martin F M,Selosse M A,Sanders I R. 2015. Mycorrhizal ecology and evolution:the past,the present,and the future. New Phytologist, 205 (4):1406-1423. doi: 10.1111/nph.13288 pmid: 25639293
[39]	Walder F, Niemann H, Natarajan M, Lehmann M F, Boller T, Wiemken A. 2012. Mycorrhizal networks: common goods of plants shared under unequal terms of trade. Plant Physiology, 159 (2):789-797. doi: 10.1104/pp.112.195727 pmid: 22517410
[40]	Wang Haiyan, Sheng Yuefan, Li Qianjin, Wang Mei, Pan Fengbing, Chen Xue Sen, Shen Xiang, Yin Chengmiao, Mao Zhiquan. 2019. Effects of Allium fistulosum,Brassica juncea and Triticum aestivum rotation on soil environment of old apple orchard. Acta Horticulturae Sinica, 46 (11):2224-2238. (in Chinese)
	王海燕, 盛月凡, 李前进, 王玫, 潘凤兵, 陈学森, 沈向, 尹承苗, 毛志泉. 2019. 葱、芥菜和小麦轮作对老龄苹果园土壤环境的影响. 园艺学报, 46 (11):2224-2238. doi: 10.16420/j.issn.0513-353x.2019-0048
[41]	Wang Mingquan, Su Jun, Li Chunxia, Gong Shichen, Yan Shuqin, Li Guoliang, Hu Guanghui, Ren Honglei. 2014. Effects of different density and fertilizer amount on yield of maize variety Longdan 42. Heilongjiang Agricultural Sciences,(5):49-51. (in Chinese)
	王明泉, 苏俊, 李春霞, 龚士琛, 闫淑琴, 李国良, 扈光辉, 任洪雷. 2014. 不同密度及施肥量对玉米品种龙单42产量的影响. 黑龙江农业科学,(5):49-51.
[42]	Weremijewicz J, Janos D P. 2013. Common mycorrhizal networks amplify size inequality in Andropogon gerardii monocultures. New Phytologist, 198 (1):203-213. doi: 10.1111/nph.12125 pmid: 23356215
[43]	Weremijewicz J,Sternberg L D S L O R,Janos D P. 2016. Common mycorrhizal networks amplify competition by preferential mineral nutrient allocation to large host plants. New Phytologist, 212 (2):461-471. doi: 10.1111/nph.14041 pmid: 27265515
[44]	Zhan Lijie, Zhang Hongbao, Li Zongtai, Meng Wei, Xu Wei, Li Linghao. 2020. Impacts of shading on seedling growth and mineral accumulation in Paeonia lactiflora. Chinese Journal of Applied Ecology, 31 (10):3473-3479. (in Chinese) doi: 10.13287/j.1001-9332.202010.029
	战丽杰, 张宏宝, 李宗泰, 孟伟, 徐伟, 李凌浩. 2020. 遮荫处理对芍药幼苗生长和矿质营养积累的影响. 应用生态学报, 31 (10):3473-3479.. doi: 10.13287/j.1001-9332.202010.029
[45]	Zhang H, Ziegler W, Han X, Trumbore S, Hartmann H. 2015. Plant carbon limitation does not reduce nitrogen transfer from arbuscular mycorrhizal fungi to Plantago lanceolata. Plant and Soil, 396 (1):369-380.
[46]	Zhang Jing, Wang Jiao’ai, Dang Jianyou, Zhang Dingyi. 2013. Effect of sowing dates on agronomic character,yield and quality of the main stalks and tillers in wheat. Journal of Agriculture, 3 (10):1. (in Chinese)
	张晶, 王姣爱, 党建友, 张定一. 2013. 播期对小麦主茎及分蘖农艺性状,产量和品质的影响. 农学学报, 3 (10):1.
[47]	Zhang Shanquan, Chen Chuan, Xu Mu, Yin Shixue. 2003. Improvement in sulfuric-acid-hydrogen-peroxide assimilating method for determination of NPK in plant. Soil,(2):174-175. (in Chinese)
	张山泉, 陈川, 徐沭, 殷士学. 2003. 硫酸-过氧化氢消化法测定植株氮磷钾方法的改进. 土壤,(2):174-175.
[48]	Zheng C, Ji B, Zhang J, Zhang F, Bever J D. 2015. Shading decreases plant carbon preferential allocation towards the most beneficial mycorrhizal mutualist. New Phytologist, 205 (1):361-368. doi: 10.1111/nph.13025 pmid: 25243653