根际氧环境对水培尖叶莴苣生长生理和品质的影响
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摘要
该研究采用自制循环营养液系统,设置循环营养液且悬根长度为0cm(T1)、静止营养液且初始悬根长度为0cm(T2)、循环营养液且悬根长度为2cm(T3)3个处理,通过测量营养液的溶解氧、pH、EC值,根系与叶片的形态学指标和生理指标,以及根系显微结构观察,以明晰根际氧环境在水培尖叶莴苣中的作用机理。结果表明:(1)尖叶莴苣幼苗的根际氧环境以T3处理最优,T2处理次之,T1处理最差,但它们营养液的pH和离子总量差异不大。(2)各处理根系和叶片的长势均表现为T3>T2>T1;其根系长度、根系表面积、根系体积、平均直径、根尖数、叶片表面积和叶片体积,以及植株茎粗、株高、根干重、地上部干重、壮苗指数、根冠比均以T3处理最大,但T3与T2间均无显著差异,而两者均显著大于T1处理,此时T3处理的壮苗指数和根冠比分别比T1显著增加了38.8%和13.4%。(3)尖叶莴苣根系维管束以及木质部的面积大小均表现为T3>T2>T1。(4)T1处理根系SOD、POD活性均高于相应的T2和T3处理,CAT活性和MDA含量均显著高于T3处理;T1处理叶片叶绿素和4种有机酸含量基本低于T2和T3,柠檬酸及琥珀酸含量以T2最大,苹果酸及丙二酸含量以T3最大。研究发现,根际低氧胁迫对水培尖叶莴苣幼苗生长、生理和品质特性均有明显抑制作用,且空气中的氧气相对于营养液中的氧气抑制作用更明显;在水培尖叶莴苣生产上设置适宜的悬根长度,可增加根系与空气接触面积,从而增加根际氧供给,解决水气矛盾,提升尖叶莴苣的产量与品质。
The self-made circulating nutrient solution system was used to set the circulating nutrient solution with a hanging root length of 0 cm(T1),a static nutrient solution and an initial hanging root length of0 cm(T2),a circulating nutrient solution and a hanging root length of 2 cm(T3).With these three treatments,we measured the dissolved oxygen,pH,EC value of nutrient solution,root and leaf morphologicaland physiological indicators,and root microstructure observation,to clarify the role of rhizosphere oxygen environment in hydroponic lettuce mechanism.The results showed that:(1)the rhizosphere oxygen environment of the lettuce was the best T3 treatment,the T2 treatment was the second,and the T1 treatment was the worst.The pH of the nutrient solution and the total amount of ions were not much different.(2)The growth of roots and leaves of each treatment showed T3>T2>T1;the root length,root surface area,root volume,average diameter,number of root tips,leaf surface area and leaf volume,and plant stem diameter and plant height,dry weight of the roots,the dry weight of the shoots,the seedling index and the root-shoot ratio were the highest in T3 treatment,but there was no significant difference between T3 and T2,and both were significantly larger than those of T1 treatment.The root-to-crown ratio was significantly increased by 38.8% and 13.4%,respectively,compared to T1.(3)The area of vascular bundles and xylem of root lettuce showed T3>T2>T1.(4)The activities of SOD and POD activities in T1 treatment were higher than those in T2 and T3 treatments.The CAT activity and MDA content were significantly higher than that of T3 treatment.The chlorophyll and four organic acids in T1 treatment were lower than that of T2 and T3.The content of acid and succinic acid is the largest with T2,and the content of malic acid and malonic acid is the largest with T3.It was found that rhizosphere hypoxia stress significantly inhibited the growth,physiology and quality of hydroponic lettuce,and the oxygen in the air was more effective than the oxygen in the nutrient solution.Setting the appropriate length of the roots of the lettuce can increase the contact area between the roots and the air,thereby increasing the supply of rhizosphere oxygen,solving the water-gas contradiction,and improving the yield and quality of lettuce.
The self-made circulating nutrient solution system was used to set the circulating nutrient solution with a hanging root length of 0 cm(T1),a static nutrient solution and an initial hanging root length of0 cm(T2),a circulating nutrient solution and a hanging root length of 2 cm(T3).With these three treatments,we measured the dissolved oxygen,pH,EC value of nutrient solution,root and leaf morphologicaland physiological indicators,and root microstructure observation,to clarify the role of rhizosphere oxygen environment in hydroponic lettuce mechanism.The results showed that:(1)the rhizosphere oxygen environment of the lettuce was the best T3 treatment,the T2 treatment was the second,and the T1 treatment was the worst.The pH of the nutrient solution and the total amount of ions were not much different.(2)The growth of roots and leaves of each treatment showed T3>T2>T1;the root length,root surface area,root volume,average diameter,number of root tips,leaf surface area and leaf volume,and plant stem diameter and plant height,dry weight of the roots,the dry weight of the shoots,the seedling index and the root-shoot ratio were the highest in T3 treatment,but there was no significant difference between T3 and T2,and both were significantly larger than those of T1 treatment.The root-to-crown ratio was significantly increased by 38.8% and 13.4%,respectively,compared to T1.(3)The area of vascular bundles and xylem of root lettuce showed T3>T2>T1.(4)The activities of SOD and POD activities in T1 treatment were higher than those in T2 and T3 treatments.The CAT activity and MDA content were significantly higher than that of T3 treatment.The chlorophyll and four organic acids in T1 treatment were lower than that of T2 and T3.The content of acid and succinic acid is the largest with T2,and the content of malic acid and malonic acid is the largest with T3.It was found that rhizosphere hypoxia stress significantly inhibited the growth,physiology and quality of hydroponic lettuce,and the oxygen in the air was more effective than the oxygen in the nutrient solution.Setting the appropriate length of the roots of the lettuce can increase the contact area between the roots and the air,thereby increasing the supply of rhizosphere oxygen,solving the water-gas contradiction,and improving the yield and quality of lettuce.
引文
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[5]胡继杰.溶解氧对水稻生长和氮素吸收及根际氮素转化的影响[D].北京:中国农业科学院,2017.
[6]付利波,贾菲,杨志新,等.悬根长度对水培生菜产量质量和水溶氧的影响[J].西南农业学报,2015,28(4):1 775-1 779.FU L B,JIA F,YANG Z X,et al.Effects of hanging roots with different length on yield,quality and water dissolved oxygen of romaine lettuce[J].Southwest China Journal of Agricultural Sciences,2015,28(4):1 775-1 779.
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[8]师恺.果菜类蔬菜根系限制下能量代谢变化及其调控机制[D].杭州:浙江大学,2007.
[9]郭世荣.营养液溶氧浓度对黄瓜和番茄根系呼吸强度的影响[J].园艺学报,2000,27(2):141-142.GUO S R.Effect of dissolved oxygen concentrations in nutrient solution on the respiratory intensity of cucumber and tomato roots[J].Acta Horticulturae Sinica,2000,27(2):141-142.
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[11] ZHAO F,XU C M,ZHANG W J,et al.Effects of rhizosphere dissolved oxygen content and nitrogen form on root traits and nitrogen accumulation in rice[J].Rice Science,2011,18(4):304-310.
[12]韩素芹,王秀峰.氮磷对甜椒穴盘苗壮苗指数的影响[J].西北农业学报,2004,13(2):128-132.HAN S Q,WANG X F.Research on effects of nitrogen and phosphorous norm on seedling index of sweet pepper plug seedlings[J].Acta Agriculturae Boreali-Occidentalis Sinica,2004,13(2):128-132.
[13]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[14] TURNER L R,DONAGHY D J,LANE P A,et al.Effect of defoliation interval on water-soluble carbohydrate and nitrogen energy reserves,regrowth of leaves and roots,and tiller number of cocksfoot(Dactylis glomerata L.)plants[J].Australian Journal of Agricultural Research,2006,57(2):243-249.
[15] TAMAIO N,NEVES M F,BRANDES A F N,et al.Quantitative analyses establish the central vascular cylinder as the standard for wood-anatomy studies in lianas having compound stems(Paullinieae:Sapindaceae)[J].Flora-Morphology,Distribution,Functional Ecology of Plants,2011,206(11):987-996.
[16] JANG G,CHOI Y D.Drought stress promotes xylem differentiation by modulating the interaction between cytokinin and jasmonic acid[J].Plant Signaling&Behavior,2018:1-10.
[17]逯明辉,宋慧,李晓明,等.冷害过程中黄瓜叶片SOD、CAT和POD活性的变化[J].西北植物学报,2005,25(8):1 570-1 573.LU M H,SONG H,LI X M,et al.Changes of SOD,CAT and POD activities in cucumber leaves during cold damage[J].Acta Botanica Boreali-Occidentalia Sinica,2005,25(8):1 570-1 573.
[18]白青云.低氧胁迫和盐胁迫下发芽粟谷γ氨基丁酸富集机理及抗氧化性研究[D].南京:南京农业大学,2009.
[19]黄琴.有机酸对发酵蔬菜品质影响的研究[D].重庆:西南大学,2009.
[20]张昊,陈金湘,刘海荷,等.营养液漂浮育苗棉苗根系适应水生环境的机理研究[J].中国农业科学,2014,47(17):3 372-3 381.ZHANG H,CHEN J X,LIU H H,et al.Adaptation mechanism to aquatic environment of cotton seedlings roots in Floating Nursing Seedlings in Nutrient Water-Bed(FNSNW)[J].Scientia Agricultura Sinica,2014,47(17):3 372-3 381.
[21]谢云韵,余常兵,侯加佳,等.低氧胁迫对油菜幼苗不定根生长的影响[J].中国油料作物学报,2013,35(3):284-289.XIE Y Y,YU C B,HOU J J,et al.Effect of low oxygen stress on adventitious roots of rape seedlings[J].Chinese Journal of Oil Crop Sciences,2013,35(3):284-289.
[22]徐春梅,谢涛,王丹英,等.根际氧浓度对水稻分蘖期养分吸收和根系形态的影响[J].中国水稻科学,2015,29(6):619-627.XU C M,XIE T,WANG D Y,et al.Effects of rhizosphere oxygen concentration on nutrient uptake and root morphology of rice at tillering stage[J].Chinese Journal of Rice Science,2015,29(6):619-627.
[23]马月花,郭世荣,杜南山,等.低氧胁迫对黄瓜幼苗生长和形态结构及有关酶活性的影响[J].南京农业大学学报,2016,39(2):213-219.MA Y H,GUO S R,DU N S,et al.Effect of hypoxia stress on growth,morpho-anatomical acclimation and activity of involved enzymes of cucumber seedlings[J].Journal of Nanjing Agricultural University,2016,39(2):213-219.
[24]邓丹,吴可为,邓泓.根区通氧状况对水稻幼苗生长及吸收镉的影响[J].生态学报,2009,29(5):2 520-2 526.DENG D,WU K W,DENG H,et al.Effects of root zone oxygenation on growth and Cd accumulation in paddy rice(Oryza sativa L.)[J].Acta Ecologica Sinica,2009,29(5):2 520-2 526.
[25]宋卫堂,高丽红,张树阁,等.深液流栽培番茄根际氧环境的试验研究[J].中国农学通报,2005,21(1):219-223.SONG W T,GAO L H,ZHANG S G,et al.EX study on root-zone Oxygen environment in DFT for tomato cultured[J].Chinese Agricultural Science Bulletin,2005,21(1):219-223.
[26]徐志豪,张德威.改善水培作物根际氧气供给的原理和实践[J].浙江农业学报,1994,(1):45-49.XU Z H,ZHANG D W.Improving aeration in hydroponicsPrinciple and practice[J].Acta Agriculturae Zhejiangensis,1994,(1):45-49.
[27]刘义玲,李天来,孙周平,等.根际低氧胁迫对网纹甜瓜生长、根呼吸代谢及抗氧化酶活性的影响[J].应用生态学报,2010,21(6):1 439-1 445.LIU Y L,LI T L,SUN Z P,et al.Impacts of root zone hypoxia stress on muskmelon growth,its root respiratory metabolism and antioxidative enzyme activities[J].Chinese Journal of Applied Ecology,2010,21(6):1 439-1 445.
[28]赵宽.几种植物光合及有机酸特征及其在喀斯特逆境检测中的应用[D].江苏镇江:江苏大学,2014.
[2]董洁,范双喜,陈青君,等.叶用莴苣遗传多样性的初步研究[J].北京农学院学报,2009,24(4):7-10.DONG J,FAN S X,CHEN Q J,et al.Study on genetic diversity of lettuce germplasm[J].Journal of Beijing University of Agriculture,2009,24(4):7-10.
[3]王晋,周相助,胡海非,等.硝态和铵态氮配比对水培油麦菜苗期生长及生理特性的影响[J].西北植物学报,2016,36(3):542-550.WANG J,ZHOU X Z,HU H F,et al.Effect of nitrogen forms on the growth and physiological characteristics of Lactuca sativa L.seedlings[J].Acta Bot.Boreal-Occident.Sin.,2016,36(3):542-550.
[4]刘士哲.现代实用无土栽培技术[M].北京:中国农业出版社,2001.
[5]胡继杰.溶解氧对水稻生长和氮素吸收及根际氮素转化的影响[D].北京:中国农业科学院,2017.
[6]付利波,贾菲,杨志新,等.悬根长度对水培生菜产量质量和水溶氧的影响[J].西南农业学报,2015,28(4):1 775-1 779.FU L B,JIA F,YANG Z X,et al.Effects of hanging roots with different length on yield,quality and water dissolved oxygen of romaine lettuce[J].Southwest China Journal of Agricultural Sciences,2015,28(4):1 775-1 779.
[7]陈艳丽.热带高温季节叶用莴苣水培关键技术体系研究[D].海口:海南大学,2014.
[8]师恺.果菜类蔬菜根系限制下能量代谢变化及其调控机制[D].杭州:浙江大学,2007.
[9]郭世荣.营养液溶氧浓度对黄瓜和番茄根系呼吸强度的影响[J].园艺学报,2000,27(2):141-142.GUO S R.Effect of dissolved oxygen concentrations in nutrient solution on the respiratory intensity of cucumber and tomato roots[J].Acta Horticulturae Sinica,2000,27(2):141-142.
[10] LIANG H,QI J W,JI Y H,et al.Response of lettuce to the coupling electrical conductivity and circulation rate of the nutrient solution in hydroponics[J].Journal of Life Sciences(ISSN1934-7391),2017,34(4):202-205.
[11] ZHAO F,XU C M,ZHANG W J,et al.Effects of rhizosphere dissolved oxygen content and nitrogen form on root traits and nitrogen accumulation in rice[J].Rice Science,2011,18(4):304-310.
[12]韩素芹,王秀峰.氮磷对甜椒穴盘苗壮苗指数的影响[J].西北农业学报,2004,13(2):128-132.HAN S Q,WANG X F.Research on effects of nitrogen and phosphorous norm on seedling index of sweet pepper plug seedlings[J].Acta Agriculturae Boreali-Occidentalis Sinica,2004,13(2):128-132.
[13]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[14] TURNER L R,DONAGHY D J,LANE P A,et al.Effect of defoliation interval on water-soluble carbohydrate and nitrogen energy reserves,regrowth of leaves and roots,and tiller number of cocksfoot(Dactylis glomerata L.)plants[J].Australian Journal of Agricultural Research,2006,57(2):243-249.
[15] TAMAIO N,NEVES M F,BRANDES A F N,et al.Quantitative analyses establish the central vascular cylinder as the standard for wood-anatomy studies in lianas having compound stems(Paullinieae:Sapindaceae)[J].Flora-Morphology,Distribution,Functional Ecology of Plants,2011,206(11):987-996.
[16] JANG G,CHOI Y D.Drought stress promotes xylem differentiation by modulating the interaction between cytokinin and jasmonic acid[J].Plant Signaling&Behavior,2018:1-10.
[17]逯明辉,宋慧,李晓明,等.冷害过程中黄瓜叶片SOD、CAT和POD活性的变化[J].西北植物学报,2005,25(8):1 570-1 573.LU M H,SONG H,LI X M,et al.Changes of SOD,CAT and POD activities in cucumber leaves during cold damage[J].Acta Botanica Boreali-Occidentalia Sinica,2005,25(8):1 570-1 573.
[18]白青云.低氧胁迫和盐胁迫下发芽粟谷γ氨基丁酸富集机理及抗氧化性研究[D].南京:南京农业大学,2009.
[19]黄琴.有机酸对发酵蔬菜品质影响的研究[D].重庆:西南大学,2009.
[20]张昊,陈金湘,刘海荷,等.营养液漂浮育苗棉苗根系适应水生环境的机理研究[J].中国农业科学,2014,47(17):3 372-3 381.ZHANG H,CHEN J X,LIU H H,et al.Adaptation mechanism to aquatic environment of cotton seedlings roots in Floating Nursing Seedlings in Nutrient Water-Bed(FNSNW)[J].Scientia Agricultura Sinica,2014,47(17):3 372-3 381.
[21]谢云韵,余常兵,侯加佳,等.低氧胁迫对油菜幼苗不定根生长的影响[J].中国油料作物学报,2013,35(3):284-289.XIE Y Y,YU C B,HOU J J,et al.Effect of low oxygen stress on adventitious roots of rape seedlings[J].Chinese Journal of Oil Crop Sciences,2013,35(3):284-289.
[22]徐春梅,谢涛,王丹英,等.根际氧浓度对水稻分蘖期养分吸收和根系形态的影响[J].中国水稻科学,2015,29(6):619-627.XU C M,XIE T,WANG D Y,et al.Effects of rhizosphere oxygen concentration on nutrient uptake and root morphology of rice at tillering stage[J].Chinese Journal of Rice Science,2015,29(6):619-627.
[23]马月花,郭世荣,杜南山,等.低氧胁迫对黄瓜幼苗生长和形态结构及有关酶活性的影响[J].南京农业大学学报,2016,39(2):213-219.MA Y H,GUO S R,DU N S,et al.Effect of hypoxia stress on growth,morpho-anatomical acclimation and activity of involved enzymes of cucumber seedlings[J].Journal of Nanjing Agricultural University,2016,39(2):213-219.
[24]邓丹,吴可为,邓泓.根区通氧状况对水稻幼苗生长及吸收镉的影响[J].生态学报,2009,29(5):2 520-2 526.DENG D,WU K W,DENG H,et al.Effects of root zone oxygenation on growth and Cd accumulation in paddy rice(Oryza sativa L.)[J].Acta Ecologica Sinica,2009,29(5):2 520-2 526.
[25]宋卫堂,高丽红,张树阁,等.深液流栽培番茄根际氧环境的试验研究[J].中国农学通报,2005,21(1):219-223.SONG W T,GAO L H,ZHANG S G,et al.EX study on root-zone Oxygen environment in DFT for tomato cultured[J].Chinese Agricultural Science Bulletin,2005,21(1):219-223.
[26]徐志豪,张德威.改善水培作物根际氧气供给的原理和实践[J].浙江农业学报,1994,(1):45-49.XU Z H,ZHANG D W.Improving aeration in hydroponicsPrinciple and practice[J].Acta Agriculturae Zhejiangensis,1994,(1):45-49.
[27]刘义玲,李天来,孙周平,等.根际低氧胁迫对网纹甜瓜生长、根呼吸代谢及抗氧化酶活性的影响[J].应用生态学报,2010,21(6):1 439-1 445.LIU Y L,LI T L,SUN Z P,et al.Impacts of root zone hypoxia stress on muskmelon growth,its root respiratory metabolism and antioxidative enzyme activities[J].Chinese Journal of Applied Ecology,2010,21(6):1 439-1 445.
[28]赵宽.几种植物光合及有机酸特征及其在喀斯特逆境检测中的应用[D].江苏镇江:江苏大学,2014.