冬前镇压和灌溉对冬小麦灌浆期旗叶光合特性和叶绿体超微结构的影响
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摘要
为明确冬前镇压和灌溉对冬小麦灌浆期旗叶功能的影响,以济麦22为材料,在2016-2017小麦生长季,对其进行冬前镇压+灌溉(C+I)、灌溉(I)和镇压(C)处理,以常规种植管理为对照(CK),研究不同冬前栽培措施对冬小麦灌浆期土壤微环境、旗叶光合特性和叶绿体超微结构以及最终产量和生物量的影响。结果表明,与CK相比,C、I及C+I处理都可提高麦田土壤含水量和土壤温度,可增加花后14～28d旗叶叶绿素相对含量(SPAD)、灌浆期旗叶最大光学效率(Fv/Fm)、净光合速率(Pn)和蒸腾速率(Tr),缓解旗叶SPAD、Fv/Fm、Pn和Tr在灌浆后期的下降,其中以C+I处理的效果最佳;灌浆中后期(花后21d),CK条件下小麦植株旗叶部分叶绿体呈圆形,大部分基粒片层变形且排列紊乱,可见较多的亲锇颗粒,但C+I、C和I处理下叶绿体的破坏程度降低,较多的叶绿体呈椭圆形,单位面积内叶绿体数和叶绿体基粒数提高,基粒片层较CK清晰且排列紧密,亲锇颗粒数量减少,其中C+I和I处理对叶绿体超微结构的改善作用大于C处理。这说明在本试验条件下,冬前镇压+灌溉较单独灌溉或镇压更有利于改善小麦灌浆期旗叶光合功能,缓解灌浆后期旗叶的衰老,促进小麦产量形成。
In order to clarify the effects of pre-winter compaction and irrigation on the function of flag leaf at grain filling stage of winter wheat,a field experiment was carried out during the wheat growing season of 2016-2017,and pre-winter compaction and irrigation(C+I),irrigation(I)and compaction(C)were applied to the winter wheat Jimai 22.With the normal growing plant as the control(CK),and the chlorophyll content(SPAD),maximum photochemical efficiency(Fv/Fm),net photosynthetic rate(Pn),transpiration rate(Tr),chloroplast ultrastructure during grain filling stage and the final yield were analyzed.The result showed that water content and temperature of the soil during the lategrain filling stage were all increased under C+I,I and C treatments,among which,the maximum increment was achieved under C+I.The SPAD in the flag leaf was higher during the late grain filling stage(14-28 days after anthesis(DAA))under the C+I and C treatments than that under I and CK treatments.The Fv/Fm,Pnand Trof the flag leaf during the whole grain filling period showed highest under C+I treatment,followed by those under I and C treatments.C+I,C or I can relief the reduction of SPAD,Fv/Fm,Pnand Tr.Among them,C+I treatment showed the most effect.Under the control conditions,the chloroplast of flag leaf was circular,and most of the grana lamellaes were deformed and disorderly arranged and a lot of osmiophilic granules appeared at 21 DAA.However,under the C+I,C and I treatments,the chloroplasts was ellipsoidal,and the number of chloroplasts and chloroplasts grana were significantly higher than those under CK treatment,and the grana lamellaes were clear and tightly packed,with a much less number of osmiophilic granules.The damage degree of chloroplasts in the flag leaf was decreased under C+I,C and I treatments.Based on the result of the SPAS,Pn,Trand Fv/Fm,the C+I treatment in this experiment was more conducive than C,I and CK treatments by improving the photosynthetic function of flag leaf during grain filling and alleviating aging of flag leaves at late grain filling stage.Thus,C+I treatment is more conducive to improve wheat production.
In order to clarify the effects of pre-winter compaction and irrigation on the function of flag leaf at grain filling stage of winter wheat,a field experiment was carried out during the wheat growing season of 2016-2017,and pre-winter compaction and irrigation(C+I),irrigation(I)and compaction(C)were applied to the winter wheat Jimai 22.With the normal growing plant as the control(CK),and the chlorophyll content(SPAD),maximum photochemical efficiency(Fv/Fm),net photosynthetic rate(Pn),transpiration rate(Tr),chloroplast ultrastructure during grain filling stage and the final yield were analyzed.The result showed that water content and temperature of the soil during the lategrain filling stage were all increased under C+I,I and C treatments,among which,the maximum increment was achieved under C+I.The SPAD in the flag leaf was higher during the late grain filling stage(14-28 days after anthesis(DAA))under the C+I and C treatments than that under I and CK treatments.The Fv/Fm,Pnand Trof the flag leaf during the whole grain filling period showed highest under C+I treatment,followed by those under I and C treatments.C+I,C or I can relief the reduction of SPAD,Fv/Fm,Pnand Tr.Among them,C+I treatment showed the most effect.Under the control conditions,the chloroplast of flag leaf was circular,and most of the grana lamellaes were deformed and disorderly arranged and a lot of osmiophilic granules appeared at 21 DAA.However,under the C+I,C and I treatments,the chloroplasts was ellipsoidal,and the number of chloroplasts and chloroplasts grana were significantly higher than those under CK treatment,and the grana lamellaes were clear and tightly packed,with a much less number of osmiophilic granules.The damage degree of chloroplasts in the flag leaf was decreased under C+I,C and I treatments.Based on the result of the SPAS,Pn,Trand Fv/Fm,the C+I treatment in this experiment was more conducive than C,I and CK treatments by improving the photosynthetic function of flag leaf during grain filling and alleviating aging of flag leaves at late grain filling stage.Thus,C+I treatment is more conducive to improve wheat production.
引文
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[9]孙岩,张宏纪,辛文利,等.苗期镇压与矮壮素结合处理对小麦生育及其基部节生长的影响[J].黑龙江农业科学,2013(2):14.SUN Y,ZHANG H J,YIN W L,et al.Effects of combination treatment of seedling-compaction and chlormequat on growth and basis festival growth of wheat[J].Heilongjiang Agricultural Sciences,2013(2):14.
[10]张迪,王红光,贾彬,等.播后镇压和冬前灌溉对冬小麦干物质转移和氮素利用效率的影响[J].麦类作物学报,2017,37(4):535.ZHANG D,WANG H G,JIA B,et al.Effect of post-sowing soil compaction and pre-winter irrigation on dry matter translocation and nitrogen use efficiency of winter wheat[J].Journal of Triticeae Crops,2017,37(4):535.
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[12]张永平,王志敏,黄琴,等.不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响[J].作物学报,2008,34(7):1213.ZHANG Y P,WANG Z M,HUANG Q,et al.Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation[J].Acta Agronomica Sinica,2008,34(7):1213.
[13]闫丽霞,于振文,石玉,等.测墒补灌对2个小麦品种旗叶叶绿素荧光及衰老特性的影响[J].中国农业科学,2017,50(8):1416.YAN L X,YU Z W,SHI Y,et al.Effects of supplemental irrigation based on soil moisture measurement on flag leaf chlorophyll fluorescence and senescence characteristics in two wheat cultivars[J].Scientia Agricultural Sinica,2017,50(8):1416.
[14]蔡瑞国,张敏,戴忠民,等.施氮水平对优质小麦旗叶光合特性和子粒生长发育的影响[J].植物营养与肥料学报,2006,12(1):49.CAI R G,ZHANG M,DAI Z M,et al.Effects of nitrogen application rate on flag leaf photosynthetic characteristics and grain growth and development of high-quality wheat[J].Plant Nutrition and Fertilizer Science,2006,12(1):49.
[15]徐雯,杨景,郑明明,等.低温对小麦植株形态、生理生化的影响及其防御研究[J].金陵科技学院学报,2013,29(2):62.XU W,YANG J,ZHENG M M,et al.The effects of low temperature on morphological and physiological characteristics of wheat plants and prevention[J].Journal of Jinling Institute of Technology,2013,29(2):62.
[16]史占良,郭进考.冷害对小麦生长发育及产量影响的研究[J].河北农业科学,1997(1):1.SHI Z L,GUO J K.Effects of cold damage on growth and yield of wheat[J].Journal of Hebei Agricultural Sciences,1997(1):1.
[17]REYNOLDS M,FOULKES M J,SLAFER G A,et al.Raising yield potential in wheat[J].Journal of Experimental Botany,2009,60(7):1899.
[18]QU X F,LU S M,WANG L H,et al.The response of different drought-resistance of wheat varieties under drought stress and the regulating role of nitric oxygen[J].Agricultural Science&Technology,2010,11(4):30.
[19]薛汉文,谢惠民,刘立明.冬灌对陕西关中地区小麦增产的作用[J].陕西农业科学,2003(1):26.XUE H W,XIE H M,LIU L M,et al.Effects of winter irrigation on increasing wheat production in Guanzhong region of Shaanxi Province[J].Journal of Shaanxi Agricultural Sciences,2003(1):26.
[20]郭建平,高素华.土壤水分对冬小麦影响机制研究[J].气象学报,2003,67(4):501.GUO J P,GAO S H.Mechanism study on impact of soil water on winter wheat[J].Acta Meteorologica Sinica,2003,67(4):501.
[21]魏晓东,陈国祥,徐艳丽,等.银杏叶片衰老过程中光合生理特性及叶绿体超微结构的变化[J].植物研究,2008(4):433.WEI X D,CHEN G X,XU Y L,et al.Changes in photosynthesis and ultrastructure of chloroplasts during leaf senescence of ginkgo[J].Bulletin of Botanical Research,2008(4):433.
[22]吴沅英,方志伟,程在全,等.小麦叶片、叶肉细胞、叶绿体的形态、结构与光合速率的关系[J].电子显微学报,1988,7(3):45.WU Y Y,FANG Z W,CHENG Z Q,et al.Relationships among morphology,structure and photosynthetic rate of wheat leaf,mesophyll cells and chloroplast[J].Journal of Electron Microscopy,1988,7(3):45.
[23]冀天会,张灿军,杨子光,等.冬小麦叶绿素荧光参数与品种抗旱性的关系[J].麦类作物学报,2005,25(4):64.JI T H,ZHANG C J,YANG Z G,et al.Study on the relationship of chlorophyll fluorescence characters and drought resistance of winter wheat varieties under drought stress[J].Journal of Triticeae Crops,2005,25(4):64.
[24]SUBRAHMANYAM D,SUBASH N,HARIS A,et al.Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought[J].Photosynthetica,2006,44(1):125.
[25]王瑞霞,闫长生,张秀英,等.春季低温对小麦产量和光合特性的影响[J].作物学报,2018,44(2):288.WANG R X,YAN C S,ZHANG X Y,et al.Effects of low temperature in spring on yield and photosynthetic characteristics of wheat[J].Acta Agronomica Sinica,2018,44(2):288.
[26]陈思思,李春燕,杨景,等.拔节期低温冻害对扬麦16光合特性及产量形成的影响[J].扬州大学学报,2014,35(3):59.CHEN S S,LI C Y,YANG J,et al.Effect of low temperature at jointing stage on photosynthetic characteristics and yield in wheat cultivar Yangmai 16[J].Journal of Yangzhou University,2014,35(3):59.
[2]李少昆,王克如,冯聚凯,等.玉米秸秆还田与不同耕作方式下影响小麦出苗的因素[J].作物学报,2006,32(3):464.LI S K,WANG K R,FENG J K,et al.Factors affecting seedling emergence in winter wheat under different tillage patterns with maize stalk mulching returned to the field[J].Acta Agromomica Sinica,2006,32(3):464.
[3]冯冬丽.冬灌对小麦生长的影响及应注意的问题[J].种业导刊,2010(12):26.FENG D L.Effects of winter irrigation on wheat growth and problems we should pay attention to[J].Journal of Seed Industry Guide,2010(12):26.
[4]张起刚,何昌永,王化国,等.应用15 N示踪技术研究冬灌对小麦生长发育及吸收N素的影响[J].核农学通报,1993(4):22.ZHANG Q G,HE C Y,WANG H G,et al.Application of 15 N tracer technique to study the growth and development of winter irrigation and absorption effect of N peptide on wheat[J].Nuclear Agronomy Bulletin,1993(4):22.
[5]王付江.小麦越冬水的作用与科学冬灌[J].种业导刊,2012,(11):21.WANG F J.Effect of overwinter irrigation on wheat and how to irrigation scientifically[J].Journal of Seed Industry Guide,2012,(11):21.
[6]YAO Z J,LI B L,CHEN R Y,et al.Effects of water and nitrogen application on photosynthetic characteristics of flag leaves and grain yield of wheat[J].Agricultural Science&Technology,2011,12(2):258.
[7]闫翠萍,肖俊红,张晶,等.冬水前移对冬小麦生长及水分利用效率的影响[J].中国生态农业学报,2018,26(5):677.YAN C P,XIAO J H,ZHANG J,et al.The effect of early winter watering on the growth and water use efficiency of winter wheat sowing under threshed maize straw by rotary machine[J].Chinese Journal of Eco-Agriculture,2018,26(5):677.
[8]毛凤梧,蒋向,王策,等.不同时期镇压对豫西旱地小麦生长发育的影响[J].农业科技通讯,2017(4):48.MAO F W,JIANG X,WANG C,et al.Effects of compaction on the growth and development of dryland wheat in western Henan[J].Agricultural Science and Technology Newsletter,2017(4):48.
[9]孙岩,张宏纪,辛文利,等.苗期镇压与矮壮素结合处理对小麦生育及其基部节生长的影响[J].黑龙江农业科学,2013(2):14.SUN Y,ZHANG H J,YIN W L,et al.Effects of combination treatment of seedling-compaction and chlormequat on growth and basis festival growth of wheat[J].Heilongjiang Agricultural Sciences,2013(2):14.
[10]张迪,王红光,贾彬,等.播后镇压和冬前灌溉对冬小麦干物质转移和氮素利用效率的影响[J].麦类作物学报,2017,37(4):535.ZHANG D,WANG H G,JIA B,et al.Effect of post-sowing soil compaction and pre-winter irrigation on dry matter translocation and nitrogen use efficiency of winter wheat[J].Journal of Triticeae Crops,2017,37(4):535.
[11]张迪,王红光,马伯威,等.播后镇压和冬前灌溉对土壤条件和冬小麦生育特性的影响[J].麦类作物学报,2014,34(6):787.ZHANG D,WANG H G,MA B W,et al.Effects of post-sowing soil compaction and pre-winter irrigation on soil conditions and growth and development of winter wheat[J].Journal of Triticeae Crops,2014,34(6):787.
[12]张永平,王志敏,黄琴,等.不同水分供给对小麦叶与非叶器官叶绿体结构和功能的影响[J].作物学报,2008,34(7):1213.ZHANG Y P,WANG Z M,HUANG Q,et al.Changes of chloroplast ultramicrostructure and function of different green organs in wheat under limited irrigation[J].Acta Agronomica Sinica,2008,34(7):1213.
[13]闫丽霞,于振文,石玉,等.测墒补灌对2个小麦品种旗叶叶绿素荧光及衰老特性的影响[J].中国农业科学,2017,50(8):1416.YAN L X,YU Z W,SHI Y,et al.Effects of supplemental irrigation based on soil moisture measurement on flag leaf chlorophyll fluorescence and senescence characteristics in two wheat cultivars[J].Scientia Agricultural Sinica,2017,50(8):1416.
[14]蔡瑞国,张敏,戴忠民,等.施氮水平对优质小麦旗叶光合特性和子粒生长发育的影响[J].植物营养与肥料学报,2006,12(1):49.CAI R G,ZHANG M,DAI Z M,et al.Effects of nitrogen application rate on flag leaf photosynthetic characteristics and grain growth and development of high-quality wheat[J].Plant Nutrition and Fertilizer Science,2006,12(1):49.
[15]徐雯,杨景,郑明明,等.低温对小麦植株形态、生理生化的影响及其防御研究[J].金陵科技学院学报,2013,29(2):62.XU W,YANG J,ZHENG M M,et al.The effects of low temperature on morphological and physiological characteristics of wheat plants and prevention[J].Journal of Jinling Institute of Technology,2013,29(2):62.
[16]史占良,郭进考.冷害对小麦生长发育及产量影响的研究[J].河北农业科学,1997(1):1.SHI Z L,GUO J K.Effects of cold damage on growth and yield of wheat[J].Journal of Hebei Agricultural Sciences,1997(1):1.
[17]REYNOLDS M,FOULKES M J,SLAFER G A,et al.Raising yield potential in wheat[J].Journal of Experimental Botany,2009,60(7):1899.
[18]QU X F,LU S M,WANG L H,et al.The response of different drought-resistance of wheat varieties under drought stress and the regulating role of nitric oxygen[J].Agricultural Science&Technology,2010,11(4):30.
[19]薛汉文,谢惠民,刘立明.冬灌对陕西关中地区小麦增产的作用[J].陕西农业科学,2003(1):26.XUE H W,XIE H M,LIU L M,et al.Effects of winter irrigation on increasing wheat production in Guanzhong region of Shaanxi Province[J].Journal of Shaanxi Agricultural Sciences,2003(1):26.
[20]郭建平,高素华.土壤水分对冬小麦影响机制研究[J].气象学报,2003,67(4):501.GUO J P,GAO S H.Mechanism study on impact of soil water on winter wheat[J].Acta Meteorologica Sinica,2003,67(4):501.
[21]魏晓东,陈国祥,徐艳丽,等.银杏叶片衰老过程中光合生理特性及叶绿体超微结构的变化[J].植物研究,2008(4):433.WEI X D,CHEN G X,XU Y L,et al.Changes in photosynthesis and ultrastructure of chloroplasts during leaf senescence of ginkgo[J].Bulletin of Botanical Research,2008(4):433.
[22]吴沅英,方志伟,程在全,等.小麦叶片、叶肉细胞、叶绿体的形态、结构与光合速率的关系[J].电子显微学报,1988,7(3):45.WU Y Y,FANG Z W,CHENG Z Q,et al.Relationships among morphology,structure and photosynthetic rate of wheat leaf,mesophyll cells and chloroplast[J].Journal of Electron Microscopy,1988,7(3):45.
[23]冀天会,张灿军,杨子光,等.冬小麦叶绿素荧光参数与品种抗旱性的关系[J].麦类作物学报,2005,25(4):64.JI T H,ZHANG C J,YANG Z G,et al.Study on the relationship of chlorophyll fluorescence characters and drought resistance of winter wheat varieties under drought stress[J].Journal of Triticeae Crops,2005,25(4):64.
[24]SUBRAHMANYAM D,SUBASH N,HARIS A,et al.Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought[J].Photosynthetica,2006,44(1):125.
[25]王瑞霞,闫长生,张秀英,等.春季低温对小麦产量和光合特性的影响[J].作物学报,2018,44(2):288.WANG R X,YAN C S,ZHANG X Y,et al.Effects of low temperature in spring on yield and photosynthetic characteristics of wheat[J].Acta Agronomica Sinica,2018,44(2):288.
[26]陈思思,李春燕,杨景,等.拔节期低温冻害对扬麦16光合特性及产量形成的影响[J].扬州大学学报,2014,35(3):59.CHEN S S,LI C Y,YANG J,et al.Effect of low temperature at jointing stage on photosynthetic characteristics and yield in wheat cultivar Yangmai 16[J].Journal of Yangzhou University,2014,35(3):59.