LED红蓝光强对水培生菜生长以及有机碳和自毒物质分泌的影响
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摘要
为了探明发光二极管(LED)红(R)蓝(B)光强对生菜(Lactuca sativa)生长的调控机理,在密闭植物工厂内,以红蓝LED (红光:蓝光=4:1, 4R:1B)为光源,研究了100μmol·m~(-2)·s~(-1)(RBL1)、200μmol·m~(-2)·s~(-1)(RBL2)和300μmol·m~(-2)·s~(-1) (RBL3)光强对水培生菜生长和根系分泌物的影响。结果表明:RBL1处理下生菜生物量较低, RBL2和RBL3地上部生物量均较高,地下部生物量以RBL3最高;三个处理下生菜叶片净光合速率随处理光强增加而逐渐升高;在根系的生长发育和形态结构方面以RBL3处理最佳, RBL1最低; RBL3处理下营养液中总有机碳(TOC)浓度较高,而积累单位生物量(地上部和地下部)释放的有机碳量和单位根系表面积释放的有机碳量各处理之间无显著差异;此外, 5种自毒物质分泌量在RBL1和RBL2处理下较低,与这两个处理相比, RBL3下5种自毒物质均显著提高超过150%,且生菜根系自毒物质分泌量依次为水杨酸>阿魏酸>苯甲酸>单宁酸>没食子酸。总之,各处理水培生菜通过根系释放的有机碳量差异不大,自毒物质以水杨酸含量最高。LED红蓝光强为200μmol·m~(-2)·s~(-1)时能够降低环境中自毒物质的含量,同时促进生菜的生长,降低光合产物在地下部的分配比例,以保证生菜产量。因此, 200μmol·m~(-2)·s~(-1)的光强更适于植物工厂水培生菜的生长。
In order to investigate the regulation mechanism of red and blue LED light intensities on growth and root exudates of hydroponic lettuce(Lactuca sativa), a study was conducted with three different light intensities [100 μmol·m~(-2)·s~(-1)(RBL1), 200 μmol·m~(-2)·s~(-1)(RBL2), and 300 μmol·m~(-2)·s~(-1)(RBL3)] delivered by red and blue LED light(4:1) in a closed plant factory. The results show that the aboveground biomass of lettuce under RBL1 treatment was lower than those under RBL2 and RBL3 treatments, and the underground biomass was the highest under RBL3 treatment. The net photosynthetic rates of lettuce leaves improved with the increases of light intensities under the three treatments, as well as the root growth and morphological development. The total organic carbon(TOC) concentration in the nutrient solution was higher under RBL3 treatment, but the amount of organic carbon released by unit biomass(aboveground and underground) and the amount of organic carbon released by unit root surface area showed no significant difference in all treatments. In addition, the secretions of five autotoxins were lower under the RBL1 and RBL2 treatments. Compared to these two treatments, the secretions of five autotoxins were significantly increased by over 150% under RBL3 treatment. The amounts of autotoxins secreted by lettuce roots were salicylic acid, ferulic acid, benzoic acid, tannic acid, and gallic acid in descending order. In summary, the amount of organic carbon released from the root was not much different under all treatments, and the salicylic acid content was highest in all autotoxins. 200 μmol·m~(-2)·s~(-1) red and blue LED light could reduce the contents of autotoxins, improve the growth of lettuce, and reduce the distribution ratios of photosynthetic products in the underground to ensure the yield of lettuce. Therefore, 200 μmol·m~(-2)·s~(-1) red and blue LED light is more suitable for the growth of hydroponic lettuce in plant factory.
In order to investigate the regulation mechanism of red and blue LED light intensities on growth and root exudates of hydroponic lettuce(Lactuca sativa), a study was conducted with three different light intensities [100 μmol·m~(-2)·s~(-1)(RBL1), 200 μmol·m~(-2)·s~(-1)(RBL2), and 300 μmol·m~(-2)·s~(-1)(RBL3)] delivered by red and blue LED light(4:1) in a closed plant factory. The results show that the aboveground biomass of lettuce under RBL1 treatment was lower than those under RBL2 and RBL3 treatments, and the underground biomass was the highest under RBL3 treatment. The net photosynthetic rates of lettuce leaves improved with the increases of light intensities under the three treatments, as well as the root growth and morphological development. The total organic carbon(TOC) concentration in the nutrient solution was higher under RBL3 treatment, but the amount of organic carbon released by unit biomass(aboveground and underground) and the amount of organic carbon released by unit root surface area showed no significant difference in all treatments. In addition, the secretions of five autotoxins were lower under the RBL1 and RBL2 treatments. Compared to these two treatments, the secretions of five autotoxins were significantly increased by over 150% under RBL3 treatment. The amounts of autotoxins secreted by lettuce roots were salicylic acid, ferulic acid, benzoic acid, tannic acid, and gallic acid in descending order. In summary, the amount of organic carbon released from the root was not much different under all treatments, and the salicylic acid content was highest in all autotoxins. 200 μmol·m~(-2)·s~(-1) red and blue LED light could reduce the contents of autotoxins, improve the growth of lettuce, and reduce the distribution ratios of photosynthetic products in the underground to ensure the yield of lettuce. Therefore, 200 μmol·m~(-2)·s~(-1) red and blue LED light is more suitable for the growth of hydroponic lettuce in plant factory.
引文
Cao G,Zhang GB,Yu JH,et al(2013).Effects of different LED light qualities on cucumber seedling growth and chlorophyll fluorescence parameters.Sci Agr Sin,46(6):1297-1304(in Chinese with English abstract)[曹刚,张国斌,郁继华等(2013).不同光质LED光源对黄瓜苗期生长及叶绿素荧光参数的影响.中国农业科学,46(6):1297-1304]
Cluis CP,Mouchel CF,Hardtke CS(2004).The Arabidopsis transcription factor HY5 integrates light and hormone signaling pathways.Plant J,38(2):332-347
Duan K(2009).Experiments of Instrument Analysis.Beijing:Chemical Industry Press(in Chinese)[段科欣(2009).仪器分析实验.北京:化学工业出版社]
Fu YR,Zhang WM,Chen GM,et al(2006).Determination of gallic acid and total phenolic acid in Sedum aizoon L.Chin Tradit Pat Med,28(7):1016-1018(in Chinese with English abstract)[付煜荣,张万明,陈桂敏等(2006),景天三七中没食子酸和总酚酸含量测定.中成药,28(7):1016-1018]
Gao J,Shi JG,Dong ST,et al(2017).Effect of different light intensities on root characteristics and grain yield of summer maize(Zea mays L.).Sci Agr Sin,50(11):2104-2113(in Chinese with English abstract)[高佳,史建国,董树亭等(2017).花粒期光照强度对夏玉米根系生长和产量的影响.中国农业科学,50(11):2104-2113]
Guo YS,Gu AS,Cui J(2011).Effects of light quality on rice seedlings growth and physiological characteristics.Chin J Appl Ecol,22(6):1485-1492(in Chinese with English abstract)[郭银生,谷艾素,崔瑾(2011).光质对水稻幼苗生长及生理特性的影响.应用生态学报,22(6):1485-1492]
Hierro JL,Callaway RW(2003).Allelopathy and exotic plant invasion.Plant Soil,256:29-39
Hogewoning SW,Trouwborst G,Maljaars H,et al(2010).Blue light dose-esponses of leaf photosynthesis,morphology,and chemical composition of Cucumis sativus grown under different combinations of red and blue light.J Exp Bot,61(11):3107-3117
Jiang N,Liu Z,Ren H,et al(2010).Changes of phenolic acids and total phenolic acids contents in cell walls of Moso bamboos at different ages.J Bamboo Res,29(1):24-28(in Chinese with English abstract)[蒋乃翔,刘志明,任海青等(2010),不同竹龄毛竹细胞壁总酚酸和酚酸类物质的含量变化.竹子研究汇刊,29(1):24-28]
Lee JG,Lee BY,Lee,HJ(2006).Accumulation of phytotoxic organic acids in reused nutrient solution during hydroponic cultivation of lettuce(Lactuca sativa L.).Sci Hortic,110(2):119-128
Li S,Pan R(1994).Effects of blue light on the growth of rice seedlings.Chin J Rice Sci,8(2):115-118(in Chinese with English abstract)[李韶山,潘瑞炽(1994).蓝光对水稻幼苗生长效应的研究.中国水稻科学,8(2):115-118]
Liu W,Yang Q,Wei L(2012).Light-Emitting Diodes(LEDs)and Their Applications in Protected Horticulture as Light Sources.Beijing:China Agricultural Science and Technology Press(in Chinese)[刘文科,杨其长,魏灵玲(2012).LED光源及其设施园艺应用.北京:中国农业科学技术出版社]
Ma G,Lian H,Yan M(2002).Effects of different mulching materials on the growth and development of tomato in greenhouse.J Jilin Agr Sci,27(4):41-43(in Chinese)[马光恕,廉华,闫明伟(2002).不同覆盖材料对大棚内番茄生长发育的影响.吉林农业科学,27(4):41-43]
Molas ML,Kiss JZ,Correll MJ(2006).Gene profiling of the red light signalling pathways in roots.J Exp Bot,57(12):3217-3229
Neumann G,Bott S,Ohler MA,et al(2014).Root exudation and root development of lettuce(Lactuca sativa L.cv.Tizian)as affected by different soils.Front Microbiol,5:2
Qian H,Xu X,Chen W,et al(2009).Allelochemical stress causes oxidative damage and inhibition of photosynthesis in Chlorella vulgaris.Chemosphere,75(3):368-375
Qiu ZP,Yang QC,Liu WK(2013).Effect of nano-TiO2 photocatalysis on yield and quality of hydroponic lettuce.Acta Agr Boreali Sin,28(2):103-107(in Chinese with English abstract)[邱志平,杨其长,刘文科(2013).纳米TiO2光催化处理对水培生菜产量及品质的影响.华北农学报,28(2):103-107]
S?b?A,Krekling T,Appelgren M(1995).Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell Tissue Organ Cult,41(2):177-185
Savvides A,Fanourakis D,van Ieperen W(2012).Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves.J Exp Bot,63(3):1135-1143
Talukder MR,Asaduzzaman M,Tanaka H,et al(2019).Electro-degradation of culture solution improves growth,yield and quality of strawberry plants grown in closed hydroponics.Sci Hortic,243:243-251
Ullah N,Haq IU,Safdar N,et al(2015).Physiological and biochemical mechanisms of allelopathy mediated by the allelochemical extracts of Phytolacca latbenia(Moq.)H.Walter.Toxicol Ind Health,31(10):931-937
Weir TL,Park SW,Vivanco JM(2004).Biochemical and physiological mechanisms mediated by allelochemicals.Curr Opin Plant Biol,7:472-479
Wu FZ,Wang W(1999).Research on soil microflora of greenhouse tomato.North Hortic,(3):1-2(in Chinese)[吴凤芝,王伟(1999).大棚番茄土壤微生物区系研究.北方园艺,(3):1-2]
Xu DQ,Gao W,Ruan J(2015).Effects of light quality on plant growth and development.Plant Physiol J,51(8):1217-1234(in Chinese with English abstract)[许大全,高伟,阮军(2015).光质对植物生长发育的影响.植物生理学报,51(8):1217-1234]
Xu J,Ren S,Zhang L,et al(2015).Effects of additional light irradiation on growth of tobacco seedling root system in a multilayer seedling culturing system.Tob Sci Tech,48(5):9-13(in Chinese with English abstract)[徐经年,任四海,张林等(2015).立体育苗模式下补光光源对烟苗根系生长的影响.烟草科技,48(5):9-13]
Yan X,Yu J,Xie J(2016).Effects of supplemental light quality and durations of illumination on growth and root activity of cucumber seedling.J Nucl Agr Sci,30(6):1211-1217(in Chinese with English abstract)[闫晓花,郁继华,颉建明(2016).补光时间及光质对黄瓜幼苗生长及根系活力的影响.核农学报,30(6):1211-1217]
Yang J,Pan T,Wang J,et al(2018).Effects of different light quality on tomato seedlings growth and roots.North Hortic,(12):77-82(in Chinese with English abstract)[杨俊伟,潘铜华,王嘉维等(2018).不同光质对番茄幼苗生长和根系的影响.北方园艺,(12):77-82]
Yu JQ,Matsui Y(1994).Phytotoxic substances in root exudates of cucumber(Cucumis sativus L.).J Chem Ecol,20(1):21-31
Yu JQ,Ye SF,Zhang MF,et al(2003).Effects of root exudates and aqueous root extracts of cucumber(Cucumis sativus)and allelochemicals,on photosynthesis and antioxidant enzymes in cucumber.Biochem Syst Ecol,31(2):129-139
Yu XB,Luo L,Zeng XT,et al(2015).Response of roots morphology and physiology to shading in maize-soybean relay strip intercropping system.Chin J Oil Crop Sci,37(2):185-193(in Chinese with English abstract)[于晓波,罗玲,曾宪堂等(2015).套作弱光胁迫对大豆苗期根系形态和生理活性的影响.中国油料作物学报,37(2):185-193]
Zha LY,Liu WK(2017).Effect of continuous light with red and blue LED lamps on growth and characteristics of photosynthesis and chlorophyll fluorescence of five lettuce cultivars.Plant Physiol J,53(9):1735-1741(in Chinese with English abstract)[查凌雁,刘文科(2017).LED红蓝光连续光照对五种生菜生长、光合和叶绿素荧光特性的影响.植物生理学报,53(9):1735-1741]
Zhong PF,Chen NL(2008).Research progress of the influence of light intensity on photosynthesis of horticultural plants.J Gansu Agr Univ,43(5):104-109(in Chinese with English abstract)[种培芳,陈年来(2008).光照强度对园艺植物光合作用影响的研究进展.甘肃农业大学学报,43(5):104-109]
Zhou CB,Zhang X,Cui QQ,et al(2017).Effects of supplementary light quality on growth and photosynthesis of pakchoi(Brassica campestris).Plant Physiol J,53(6):1030-1038(in Chinese with English abstract)[周成波,张旭,崔青青等(2017).LED补光光质对小白菜生长及光合作用的影响.植物生理学报,53(6):1030-1038]
Cluis CP,Mouchel CF,Hardtke CS(2004).The Arabidopsis transcription factor HY5 integrates light and hormone signaling pathways.Plant J,38(2):332-347
Duan K(2009).Experiments of Instrument Analysis.Beijing:Chemical Industry Press(in Chinese)[段科欣(2009).仪器分析实验.北京:化学工业出版社]
Fu YR,Zhang WM,Chen GM,et al(2006).Determination of gallic acid and total phenolic acid in Sedum aizoon L.Chin Tradit Pat Med,28(7):1016-1018(in Chinese with English abstract)[付煜荣,张万明,陈桂敏等(2006),景天三七中没食子酸和总酚酸含量测定.中成药,28(7):1016-1018]
Gao J,Shi JG,Dong ST,et al(2017).Effect of different light intensities on root characteristics and grain yield of summer maize(Zea mays L.).Sci Agr Sin,50(11):2104-2113(in Chinese with English abstract)[高佳,史建国,董树亭等(2017).花粒期光照强度对夏玉米根系生长和产量的影响.中国农业科学,50(11):2104-2113]
Guo YS,Gu AS,Cui J(2011).Effects of light quality on rice seedlings growth and physiological characteristics.Chin J Appl Ecol,22(6):1485-1492(in Chinese with English abstract)[郭银生,谷艾素,崔瑾(2011).光质对水稻幼苗生长及生理特性的影响.应用生态学报,22(6):1485-1492]
Hierro JL,Callaway RW(2003).Allelopathy and exotic plant invasion.Plant Soil,256:29-39
Hogewoning SW,Trouwborst G,Maljaars H,et al(2010).Blue light dose-esponses of leaf photosynthesis,morphology,and chemical composition of Cucumis sativus grown under different combinations of red and blue light.J Exp Bot,61(11):3107-3117
Jiang N,Liu Z,Ren H,et al(2010).Changes of phenolic acids and total phenolic acids contents in cell walls of Moso bamboos at different ages.J Bamboo Res,29(1):24-28(in Chinese with English abstract)[蒋乃翔,刘志明,任海青等(2010),不同竹龄毛竹细胞壁总酚酸和酚酸类物质的含量变化.竹子研究汇刊,29(1):24-28]
Lee JG,Lee BY,Lee,HJ(2006).Accumulation of phytotoxic organic acids in reused nutrient solution during hydroponic cultivation of lettuce(Lactuca sativa L.).Sci Hortic,110(2):119-128
Li S,Pan R(1994).Effects of blue light on the growth of rice seedlings.Chin J Rice Sci,8(2):115-118(in Chinese with English abstract)[李韶山,潘瑞炽(1994).蓝光对水稻幼苗生长效应的研究.中国水稻科学,8(2):115-118]
Liu W,Yang Q,Wei L(2012).Light-Emitting Diodes(LEDs)and Their Applications in Protected Horticulture as Light Sources.Beijing:China Agricultural Science and Technology Press(in Chinese)[刘文科,杨其长,魏灵玲(2012).LED光源及其设施园艺应用.北京:中国农业科学技术出版社]
Ma G,Lian H,Yan M(2002).Effects of different mulching materials on the growth and development of tomato in greenhouse.J Jilin Agr Sci,27(4):41-43(in Chinese)[马光恕,廉华,闫明伟(2002).不同覆盖材料对大棚内番茄生长发育的影响.吉林农业科学,27(4):41-43]
Molas ML,Kiss JZ,Correll MJ(2006).Gene profiling of the red light signalling pathways in roots.J Exp Bot,57(12):3217-3229
Neumann G,Bott S,Ohler MA,et al(2014).Root exudation and root development of lettuce(Lactuca sativa L.cv.Tizian)as affected by different soils.Front Microbiol,5:2
Qian H,Xu X,Chen W,et al(2009).Allelochemical stress causes oxidative damage and inhibition of photosynthesis in Chlorella vulgaris.Chemosphere,75(3):368-375
Qiu ZP,Yang QC,Liu WK(2013).Effect of nano-TiO2 photocatalysis on yield and quality of hydroponic lettuce.Acta Agr Boreali Sin,28(2):103-107(in Chinese with English abstract)[邱志平,杨其长,刘文科(2013).纳米TiO2光催化处理对水培生菜产量及品质的影响.华北农学报,28(2):103-107]
S?b?A,Krekling T,Appelgren M(1995).Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell Tissue Organ Cult,41(2):177-185
Savvides A,Fanourakis D,van Ieperen W(2012).Co-ordination of hydraulic and stomatal conductances across light qualities in cucumber leaves.J Exp Bot,63(3):1135-1143
Talukder MR,Asaduzzaman M,Tanaka H,et al(2019).Electro-degradation of culture solution improves growth,yield and quality of strawberry plants grown in closed hydroponics.Sci Hortic,243:243-251
Ullah N,Haq IU,Safdar N,et al(2015).Physiological and biochemical mechanisms of allelopathy mediated by the allelochemical extracts of Phytolacca latbenia(Moq.)H.Walter.Toxicol Ind Health,31(10):931-937
Weir TL,Park SW,Vivanco JM(2004).Biochemical and physiological mechanisms mediated by allelochemicals.Curr Opin Plant Biol,7:472-479
Wu FZ,Wang W(1999).Research on soil microflora of greenhouse tomato.North Hortic,(3):1-2(in Chinese)[吴凤芝,王伟(1999).大棚番茄土壤微生物区系研究.北方园艺,(3):1-2]
Xu DQ,Gao W,Ruan J(2015).Effects of light quality on plant growth and development.Plant Physiol J,51(8):1217-1234(in Chinese with English abstract)[许大全,高伟,阮军(2015).光质对植物生长发育的影响.植物生理学报,51(8):1217-1234]
Xu J,Ren S,Zhang L,et al(2015).Effects of additional light irradiation on growth of tobacco seedling root system in a multilayer seedling culturing system.Tob Sci Tech,48(5):9-13(in Chinese with English abstract)[徐经年,任四海,张林等(2015).立体育苗模式下补光光源对烟苗根系生长的影响.烟草科技,48(5):9-13]
Yan X,Yu J,Xie J(2016).Effects of supplemental light quality and durations of illumination on growth and root activity of cucumber seedling.J Nucl Agr Sci,30(6):1211-1217(in Chinese with English abstract)[闫晓花,郁继华,颉建明(2016).补光时间及光质对黄瓜幼苗生长及根系活力的影响.核农学报,30(6):1211-1217]
Yang J,Pan T,Wang J,et al(2018).Effects of different light quality on tomato seedlings growth and roots.North Hortic,(12):77-82(in Chinese with English abstract)[杨俊伟,潘铜华,王嘉维等(2018).不同光质对番茄幼苗生长和根系的影响.北方园艺,(12):77-82]
Yu JQ,Matsui Y(1994).Phytotoxic substances in root exudates of cucumber(Cucumis sativus L.).J Chem Ecol,20(1):21-31
Yu JQ,Ye SF,Zhang MF,et al(2003).Effects of root exudates and aqueous root extracts of cucumber(Cucumis sativus)and allelochemicals,on photosynthesis and antioxidant enzymes in cucumber.Biochem Syst Ecol,31(2):129-139
Yu XB,Luo L,Zeng XT,et al(2015).Response of roots morphology and physiology to shading in maize-soybean relay strip intercropping system.Chin J Oil Crop Sci,37(2):185-193(in Chinese with English abstract)[于晓波,罗玲,曾宪堂等(2015).套作弱光胁迫对大豆苗期根系形态和生理活性的影响.中国油料作物学报,37(2):185-193]
Zha LY,Liu WK(2017).Effect of continuous light with red and blue LED lamps on growth and characteristics of photosynthesis and chlorophyll fluorescence of five lettuce cultivars.Plant Physiol J,53(9):1735-1741(in Chinese with English abstract)[查凌雁,刘文科(2017).LED红蓝光连续光照对五种生菜生长、光合和叶绿素荧光特性的影响.植物生理学报,53(9):1735-1741]
Zhong PF,Chen NL(2008).Research progress of the influence of light intensity on photosynthesis of horticultural plants.J Gansu Agr Univ,43(5):104-109(in Chinese with English abstract)[种培芳,陈年来(2008).光照强度对园艺植物光合作用影响的研究进展.甘肃农业大学学报,43(5):104-109]
Zhou CB,Zhang X,Cui QQ,et al(2017).Effects of supplementary light quality on growth and photosynthesis of pakchoi(Brassica campestris).Plant Physiol J,53(6):1030-1038(in Chinese with English abstract)[周成波,张旭,崔青青等(2017).LED补光光质对小白菜生长及光合作用的影响.植物生理学报,53(6):1030-1038]