遮光对连香树幼苗光合特性及其叶片解剖结构的影响
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摘要
以2年生连香树实生苗为材料,在田间通过黑色遮阳网设置全光照(L_0)及透光率55%(L_1)、25%(L_2)和10%(L_3)4种光环境,研究遮光对连香树幼苗光合作用及叶片解剖结构的影响。结果表明:(1)连香树幼苗叶片P_n在全光和L_1处理下呈非典型的"乁"形变化,未出现"午休"现象,中午14:00出现极值,而在L_2和L_3处理下变化相对缓和,极值出现在中午12:00;叶片G_s呈现与P_n类似的变化趋势,而C_i则呈基本一致的凹形变化。(2)各处理P_n、G_s和T_r的日均值均表现为L_0>L_1>L_2>L_3,而C_i的日均值则呈相反的顺序;P_n与G_s、T_r、气温和光合有效辐射均呈极显著正相关关系(P<0.01)。(3)全光照连香树幼苗的光补偿点(LCP)、光饱和点(LSP)、暗呼吸速率(R_d)均显著高于遮光处理,并维持较高的P_n而未出现明显的光抑制;遮光导致幼苗的LCP、LSP、R_d显著降低,有利于充分利用弱光,以满足低光环境下植株的正常生长。(4)与全光照相比,遮光下连香树叶片气孔密度显著变小,但气孔器长度、气孔器宽度、单个气孔器面积显著增加,气孔器面积百分比减少,影响幼苗细胞内外的水分和气体传递。(5)遮光条件下,连香树叶片明显变薄,表皮细胞厚度减小,栅栏组织(PT)厚度降低,排列变得疏松,海绵组织(ST)厚度增加,PT/ST相应减小。(6)与全光照相比,强度遮光下(L_2和L_3)连香树幼苗生长受阻,苗高(H)和基径(D)明显减小,生物量模型D~2H下降;而轻度遮光(L_1)下幼苗H和D、H/D和D~2H均未出现显著变化。研究发现,连香树具有一定的光忍耐性和喜光性,对光照条件的生态幅较宽,轻度遮光影响较小,但强度遮光对连香树幼苗气体交换参数和光合响应特征产生了显著影响,同时影响了叶片的解剖结构和气孔分布特征,从而影响连香树幼苗的生长形态。在育苗生产中,适度遮光有利于降低气温、减小蒸腾,但遮光后田间有效辐射强度应保持在自然光强的55%以上。
Effects of shading on photosynthesis characteristics and leaf anatomical structure of Cercidiphyllum japonicum seedlings were researched. Four shade treatments including full sunlight(L_0), 55% sunlight(L_1), 25% sunlight(L_2) and 10% sunlight(L_3) were set. The results showed that:(1) shading had an significant effect on daily variation characteristics of photosynthesis of seedlings. The changes of P_n under L_1 treatment were similar to L_0, and showed rough "乁"-shape, and the "noon break" did not appear in the course of the day, and the peak value appeared at 14:00 AM, that of other treatments was relatively mild, and the peak value appeared at 12:00 AM. The variation curve of G_s similar to P_n, and C_i showed the basically concave change.(2) The order of daily average of P_(n,) G_s and T_r was L_0 > L_1 > L_2 > L_3, and the order of C_i was versa. Correlation analysis showed that P_n was positively correlated with G_s, T_r, T_a and photosynthetically active radiation(PAR)(P < 0.01).(3) The photosynthesis-light response characteristics under different treatments were obviously different. Under full sunlight, the light compensation point(LCP), light saturation point(LSP) and dark respiration rate(R_d) of seedlings were significantly higher than that under shading. The seedlings can maintain high P_n without obvious light inhibition, and C. japonicum seedlings had the ability of light tolerance in a certain degree, and showed the property of light demander. Shading resulted in a significant decrease of LCP, LSP and R_d, which was conducive to the growth of seedings under low light conditions.(4) Shading significantly affected the stomatal distribution on leaf surface. Under shading treatments, the stomatal density was smaller compared with the control, but the length of stomata, width of stomata and single stomatal area increased significantly. The percentage of stomatal area reduced, which will affect water and gas delivery between inside and outside of the seedling cells.(5) Under shading, the leaf thickness decreased, the upper epidermis thickened and the lower epidermis thinned. The thickness of palisade tissue(PT) decreased, sponge tissue(ST) was increased, and the PT/ST decreased accordingly.(6) Shading affects the growth form of C. japonicum seedlings. Compared with the control, the seedlings growth were inhibited under intensive shading(L_2 and L_3), the height(H) and basal diameter(D) decreased, and D~2H dwindled accordingly, while the mild shading(L_1) had no significant effects on H, D, H/D and D~2H of seedlings. So the ecological width of C. japonicum seedlings to the light was wide. In forest seedlings production, moderate shading is beneficial to reduce air temperature and reduce transpiration, but the PAR in the field should be kept above 55% of natural light intensity.
Effects of shading on photosynthesis characteristics and leaf anatomical structure of Cercidiphyllum japonicum seedlings were researched. Four shade treatments including full sunlight(L_0), 55% sunlight(L_1), 25% sunlight(L_2) and 10% sunlight(L_3) were set. The results showed that:(1) shading had an significant effect on daily variation characteristics of photosynthesis of seedlings. The changes of P_n under L_1 treatment were similar to L_0, and showed rough "乁"-shape, and the "noon break" did not appear in the course of the day, and the peak value appeared at 14:00 AM, that of other treatments was relatively mild, and the peak value appeared at 12:00 AM. The variation curve of G_s similar to P_n, and C_i showed the basically concave change.(2) The order of daily average of P_(n,) G_s and T_r was L_0 > L_1 > L_2 > L_3, and the order of C_i was versa. Correlation analysis showed that P_n was positively correlated with G_s, T_r, T_a and photosynthetically active radiation(PAR)(P < 0.01).(3) The photosynthesis-light response characteristics under different treatments were obviously different. Under full sunlight, the light compensation point(LCP), light saturation point(LSP) and dark respiration rate(R_d) of seedlings were significantly higher than that under shading. The seedlings can maintain high P_n without obvious light inhibition, and C. japonicum seedlings had the ability of light tolerance in a certain degree, and showed the property of light demander. Shading resulted in a significant decrease of LCP, LSP and R_d, which was conducive to the growth of seedings under low light conditions.(4) Shading significantly affected the stomatal distribution on leaf surface. Under shading treatments, the stomatal density was smaller compared with the control, but the length of stomata, width of stomata and single stomatal area increased significantly. The percentage of stomatal area reduced, which will affect water and gas delivery between inside and outside of the seedling cells.(5) Under shading, the leaf thickness decreased, the upper epidermis thickened and the lower epidermis thinned. The thickness of palisade tissue(PT) decreased, sponge tissue(ST) was increased, and the PT/ST decreased accordingly.(6) Shading affects the growth form of C. japonicum seedlings. Compared with the control, the seedlings growth were inhibited under intensive shading(L_2 and L_3), the height(H) and basal diameter(D) decreased, and D~2H dwindled accordingly, while the mild shading(L_1) had no significant effects on H, D, H/D and D~2H of seedlings. So the ecological width of C. japonicum seedlings to the light was wide. In forest seedlings production, moderate shading is beneficial to reduce air temperature and reduce transpiration, but the PAR in the field should be kept above 55% of natural light intensity.
引文
[1]李颖娇,李国防,王志博,等.中午适度遮阴对苹果叶片光合特性的影响[J].西北农业学报,2015,24(2):91-96.LI Y J,LI G F,WANG Z B,et al.Effect of moderately shade on photosynthetic characteristic in Malus domestica Borkh.leaves duing Middy[J].Acta Agriculturae Boreali-occidentalis Sinica,2015,24(2):91-96.
[2]KHAN S R,ROSE R,HAASE D,et al.Effects of shade on morphology,chlorophyll concentration,and chlorophyll fluorescence of four pacific northwest conifer species[J].New Forests,2000,19:171-186.
[3]颜超,王中生,安树青,等.濒危植物银缕梅(Parrotia subaequalis)不同径级个体的光合能力差异与更新限制[J].生态学报,2008,28(9):4 153-4 161.YAN C,WANG Z S,AN S Q,et al.Differences in photosynthetic capacity among different diameter-classes of Parrotia subaequalispopulations and their implications to regeneration limitation[J].Acta Ecologica Sinica,2008,28(9):4 153-4 161.
[4]郭连金,杜佳朋,吴艳萍,等.香果树实生苗的光合特性及其与环境因子的关系[J].应用生态学报,2017,28(5):1 473-1 481.GUO L J,DU J P,WU Y P,et al.Photosynthetic characteristics of Emmenopterys henryi seedlings and their relationships with environmental factors[J].Chinese Journal of Applied Ecology,2017,28(5):1 473-1 481
[5]任全进,于金平.古老稀有植物:连香树[J].中国野生植物资源,1998,17(4):37-38.REN Q J,YU J P.Ancient rare plant:Cercidiphyllum japonicum[J].Chinese Wild Plant Resources,1998,17(4):37-38.
[6]傅立国,金鉴明.中国植物红皮书:稀有濒危植物(第一册)[M].北京:科学出版社,1992:212-213.
[7]袁丽洁,方向民,崔波,等.濒危植物连香树的传粉生物学研究[J].河南农业大学学报,2007,41(6):650-654.YUAN L J,FANG X M,CUI B,et al.Study on pollination biology of endangered plant Cercidiphyllum japonicum[J].Journal of Henan Agricultural University,2007,41(6):650-654.
[8]杨荣慧,孙宝胜,刘守阳.秦岭地区连香树分布现状与濒危机理[J].东北林业大学学报,2012,40(6):19-22.YANG R H,SUN B S,LIU S Y.Shouyang resource distribution and endangered mechanism of genus Cercidiphyllumin Qinling Region[J].Journal of Northeast Forestry University,2012,40(6):19-22.
[9]何东,魏新增,李连发,等.神农架山地河岸带连香树的种群结构与动态[J].植物生态学报,2009,33(3):469-481.HE D,WEI X Z,LI L F,et al.Population structure and dynamics of Cercidiphyllum japonicumin riparian zones of the shennongjia mountainous region,Central China[J].Chinese Journal of Plant Ecology,2009,33(3):469-481.
[10]刘光华,操国兴,张望,等.濒危植物连香树种群结构与动态研究[J].生物学杂志,2008,25(5):24-27.LIU G H,CAO G X,ZHANG W,et al.Population structure and dynamics of endangered plant Cercidiphyllum japonicum[J].Journal of Biology,2008,25(5):24-27.
[11]麦苗苗,石大兴,王米力,等.PEG处理对连香树种子萌发与芽苗生长的影响[J].林业科学,2009,45(10):94-99.MAI M M,SHI D X,WANG M L,et al.Seed germination and seedling growth of Cercidiphyllum japonicum with PEGtreatment[J].Scientia Silvae Sinicae,2009,45(10):94-99.
[12]麦苗苗,石大兴,王米力.连香树离体快繁初步研究[J].园艺学报,2006,33(1):186-189.MAI M M,SHI D X,WANG M L.A preliminary study on the invitro oulture of Cercidiphyllum japonicum Sieb.et Zucc.[J].Acta Horticulturae Sinica,2006,33(1):186-189.
[13]王静,张小平,李文良,等.濒危植物连香树居群的遗传多样性和遗传分化研究[J].植物研究,2010,30(2):208-214.WANG J,ZHANG X P,LI W L,et al.Genetic diversity and genetic variation of populations endangered tree Cercidiphyllum japonicum[J].Bulletin of Botanical Research,2010,30(2):208-214.
[14]黎明,段增强,李莲枝,等.连香树营养器官的解剖学研究[J].河南农业大学学报,2005,39(2):178-181.LI M,DUAN Z Q,LI L Z,et al.Anatomical study on the nutritive organs of Cercidiphyllum japonicum[J].Journal of Henan Agricultural University,2005,39(2):178-181.
[15]王东,高淑珍.中国连香树科的系统研究.叶的宏观结构和叶柄维管束变化[J].西北植物学报,1990,10(1):37-41.WANG D,GAO S Z.A study on the relationships of Cercidiphyllaceae.I.Leaf architecture and vasculature changes of petiole[J].Acta Botanica Boreali-Occidentalia Sinica,1990,10(1):37-41.
[16]张旺锋,樊大勇,谢宗强,等.濒危植物银杉幼树对生长光强的季节性光合响应[J].生物多样性,2005,13(5):387-397.ZANG W F,FAN D Y,XIE Z Q,et al.The seasonal photosynthetic responses of seedlings of the endangered plant Cathaya argyrophyllato different growth light environments[J].Biodiversity Science,2005,13(5):387-397.
[17]刘建锋,杨文娟,江泽平,等.遮光对濒危植物崖柏光合作用和叶绿素荧光参数的影响[J].生态学报,2011,31(20):5 999-6 004.LIU J F,YANG W J,JIANG Z P,et al.Effects of shading on photosynthetic characteristics and chlorophyll fluorescence parameters in leaves of the endangered plant Thuja sutchuenensis[J].Acta Ecologica Sinica,2011,31(20):5 999-6 004.
[18]RICHARDSON A,BERLYN G P.Changes in foliar spectral reflectanceand chlorophyll fluorescence of four temperate species followingbranch cutting[J].Tree Physiology.2002,22(7):499-506.
[19]李正理.植物制片技术(第二版)[M].北京:科学出版社,1987.
[20]蔡艳飞,李世峰,王继华,等.遮光对油用牡丹植株生长和光合特性的影响[J].西北植物学报,2016,36(8):1 623-1 631.CAI Y F,LI S F,WANG J H,et al.Effects of shading on growth and photosynthetic characteristic of oil peony[J].Acta Botanica Boreali-Occidentalia Sinica,2016,36(8):1 623-1 631.
[21]王文栋,白志强,阿里木·买买提,等.天山林区6种优势种灌木林生物量比较及估测模型[J].生态学报,2016,36(9):2 695-2 704.WANG W D,BAI Z Q,ALIMU M M T,et al.Biomass comparison and estimation models for six dominant species of woody shrubs in the forest zones of the Tianshan Mountains[J].Acta Ecologica Sinica,2016,36(9):2 695-2 704.
[22]刘江涛,刘立佳.SPSS数据统计与分析应用教程[M].北京:清华大学出版社,2017,74-99;143-147.
[23]PHILIPPE M,CLAUDIO P,MATTHEW P.Limitations to photosynthesis under light and heat stress in three high-yielding wheat genotypes[J].J Plant Physiol,2003,160:657-666.
[24]许克章,史跃林,许贵民,等.保护地黄瓜叶片光合作用温度特性的研究[J].园艺学报,1993,20(1):51-55.XU K Z,SHI Y L,XU G M,et al.Studies on photosynthetic temperature characteristic of cucumber leaves in protective field[J].Acta Horticulturae Sinica,1993,20(1):51-55.
[25]施建敏,郭起荣,杨光耀.毛竹光合动态研究[J].林业科学研究,2005,18(5):551-555.SHI J M,GUO Q R,YANG G Y.Study on thephotosynthetic dynamic variation of Phyllostachys edulis[J].Forest Research,2005,18(5):551-555.
[26]梁开明,曹洪麟,徐志防,等.台湾青枣及野生种的光合作用日变化及光响应特征[J].园艺学报,2008,35(6):793-798.LIANG K M,CAO H L,XU Z F,et al.Diurnal variations and light response of photosynthesis in three cultivars and wild type of Ziziphusm auritiana Lam[J].Acta Horticulturae Sinica,2008,35(6):793-798.
[27]PEARCY RW,SIMS DA.Photosynthetic acclimation to changing light environments:Scaling from the leaf to the whole plant//Caldwell MM,Pearcy RW,eds.Exploitation of environmental heterogeneity by plants:ecophysiological processes above-and belowground.San Diego:Academic Press,1994:145-174.
[28]郑玉龙,姜春玲,冯玉龙.植物的气孔发生[J].植物生理学通讯,2005,41(6):847-850.ZHENG Y L,JIANG C L,FENG Y L.Stomatal generation in plants[J].Plant Physiology Communications,2005,41(6):847-850.
[29]HETHERINGTON A M,WOODWARD F I.The role of stomata in sensing and driving environmental change[J].Nature,2003,424:901-908.
[30]ONWUEME I C,JOHNSTON M.Influence of shade on stomatal density,leaf size and other leaf characteristics in the major tropical root crops,tannia,sweet potato,yam,cassava and taro[J].Experimental Agriculture,2000,36(4):509-516.
[31]徐坤,邹琦,赵燕.土壤水分胁迫与遮光对生姜生长特性的影响[J].应用生态学报,2003,14(10):1 645-1 648.XU K,ZOU Q,ZHAO Y.Effects of soil water stress and shading on growth characteristics of ginger[J].Chinese Journal of Applied Ecolog,2003,14(10):1 645-1 648.
[32]JAMES S A,BELL D T.Influence of light avilability on lesf structure and growth of two Euclyptus ssp.Globulus Provenances[J].Tree physiology,2000,20(15):1 007-1 018.
[33]王荣,郭志华.不同光环境下枫香幼苗的叶片解剖结构[J].生态学杂志,2007,26(11):1 719-1 724.WANG R,GUO Z H.Responses of seedling leaf anatomical structure of Liquidambar formosana,a deciduous broadleaf tree,to different light regimes[J].Chinese Journal of Ecology,2007,26(11):1 719-1 724.
[34]杨文权,褚继鹏,寇建村,等.遮阴对白三叶叶片解剖结构和光合特性的影响[J].草地科学,2015,23(3):653-657.YANG W Q,CHU J P,KOU J C,et al.Effects of shading on the leave anatomical structure and photosynthetic characteristics of white clover[J].Acta Agrestia Sinica,2015,23(3):653-657.
[2]KHAN S R,ROSE R,HAASE D,et al.Effects of shade on morphology,chlorophyll concentration,and chlorophyll fluorescence of four pacific northwest conifer species[J].New Forests,2000,19:171-186.
[3]颜超,王中生,安树青,等.濒危植物银缕梅(Parrotia subaequalis)不同径级个体的光合能力差异与更新限制[J].生态学报,2008,28(9):4 153-4 161.YAN C,WANG Z S,AN S Q,et al.Differences in photosynthetic capacity among different diameter-classes of Parrotia subaequalispopulations and their implications to regeneration limitation[J].Acta Ecologica Sinica,2008,28(9):4 153-4 161.
[4]郭连金,杜佳朋,吴艳萍,等.香果树实生苗的光合特性及其与环境因子的关系[J].应用生态学报,2017,28(5):1 473-1 481.GUO L J,DU J P,WU Y P,et al.Photosynthetic characteristics of Emmenopterys henryi seedlings and their relationships with environmental factors[J].Chinese Journal of Applied Ecology,2017,28(5):1 473-1 481
[5]任全进,于金平.古老稀有植物:连香树[J].中国野生植物资源,1998,17(4):37-38.REN Q J,YU J P.Ancient rare plant:Cercidiphyllum japonicum[J].Chinese Wild Plant Resources,1998,17(4):37-38.
[6]傅立国,金鉴明.中国植物红皮书:稀有濒危植物(第一册)[M].北京:科学出版社,1992:212-213.
[7]袁丽洁,方向民,崔波,等.濒危植物连香树的传粉生物学研究[J].河南农业大学学报,2007,41(6):650-654.YUAN L J,FANG X M,CUI B,et al.Study on pollination biology of endangered plant Cercidiphyllum japonicum[J].Journal of Henan Agricultural University,2007,41(6):650-654.
[8]杨荣慧,孙宝胜,刘守阳.秦岭地区连香树分布现状与濒危机理[J].东北林业大学学报,2012,40(6):19-22.YANG R H,SUN B S,LIU S Y.Shouyang resource distribution and endangered mechanism of genus Cercidiphyllumin Qinling Region[J].Journal of Northeast Forestry University,2012,40(6):19-22.
[9]何东,魏新增,李连发,等.神农架山地河岸带连香树的种群结构与动态[J].植物生态学报,2009,33(3):469-481.HE D,WEI X Z,LI L F,et al.Population structure and dynamics of Cercidiphyllum japonicumin riparian zones of the shennongjia mountainous region,Central China[J].Chinese Journal of Plant Ecology,2009,33(3):469-481.
[10]刘光华,操国兴,张望,等.濒危植物连香树种群结构与动态研究[J].生物学杂志,2008,25(5):24-27.LIU G H,CAO G X,ZHANG W,et al.Population structure and dynamics of endangered plant Cercidiphyllum japonicum[J].Journal of Biology,2008,25(5):24-27.
[11]麦苗苗,石大兴,王米力,等.PEG处理对连香树种子萌发与芽苗生长的影响[J].林业科学,2009,45(10):94-99.MAI M M,SHI D X,WANG M L,et al.Seed germination and seedling growth of Cercidiphyllum japonicum with PEGtreatment[J].Scientia Silvae Sinicae,2009,45(10):94-99.
[12]麦苗苗,石大兴,王米力.连香树离体快繁初步研究[J].园艺学报,2006,33(1):186-189.MAI M M,SHI D X,WANG M L.A preliminary study on the invitro oulture of Cercidiphyllum japonicum Sieb.et Zucc.[J].Acta Horticulturae Sinica,2006,33(1):186-189.
[13]王静,张小平,李文良,等.濒危植物连香树居群的遗传多样性和遗传分化研究[J].植物研究,2010,30(2):208-214.WANG J,ZHANG X P,LI W L,et al.Genetic diversity and genetic variation of populations endangered tree Cercidiphyllum japonicum[J].Bulletin of Botanical Research,2010,30(2):208-214.
[14]黎明,段增强,李莲枝,等.连香树营养器官的解剖学研究[J].河南农业大学学报,2005,39(2):178-181.LI M,DUAN Z Q,LI L Z,et al.Anatomical study on the nutritive organs of Cercidiphyllum japonicum[J].Journal of Henan Agricultural University,2005,39(2):178-181.
[15]王东,高淑珍.中国连香树科的系统研究.叶的宏观结构和叶柄维管束变化[J].西北植物学报,1990,10(1):37-41.WANG D,GAO S Z.A study on the relationships of Cercidiphyllaceae.I.Leaf architecture and vasculature changes of petiole[J].Acta Botanica Boreali-Occidentalia Sinica,1990,10(1):37-41.
[16]张旺锋,樊大勇,谢宗强,等.濒危植物银杉幼树对生长光强的季节性光合响应[J].生物多样性,2005,13(5):387-397.ZANG W F,FAN D Y,XIE Z Q,et al.The seasonal photosynthetic responses of seedlings of the endangered plant Cathaya argyrophyllato different growth light environments[J].Biodiversity Science,2005,13(5):387-397.
[17]刘建锋,杨文娟,江泽平,等.遮光对濒危植物崖柏光合作用和叶绿素荧光参数的影响[J].生态学报,2011,31(20):5 999-6 004.LIU J F,YANG W J,JIANG Z P,et al.Effects of shading on photosynthetic characteristics and chlorophyll fluorescence parameters in leaves of the endangered plant Thuja sutchuenensis[J].Acta Ecologica Sinica,2011,31(20):5 999-6 004.
[18]RICHARDSON A,BERLYN G P.Changes in foliar spectral reflectanceand chlorophyll fluorescence of four temperate species followingbranch cutting[J].Tree Physiology.2002,22(7):499-506.
[19]李正理.植物制片技术(第二版)[M].北京:科学出版社,1987.
[20]蔡艳飞,李世峰,王继华,等.遮光对油用牡丹植株生长和光合特性的影响[J].西北植物学报,2016,36(8):1 623-1 631.CAI Y F,LI S F,WANG J H,et al.Effects of shading on growth and photosynthetic characteristic of oil peony[J].Acta Botanica Boreali-Occidentalia Sinica,2016,36(8):1 623-1 631.
[21]王文栋,白志强,阿里木·买买提,等.天山林区6种优势种灌木林生物量比较及估测模型[J].生态学报,2016,36(9):2 695-2 704.WANG W D,BAI Z Q,ALIMU M M T,et al.Biomass comparison and estimation models for six dominant species of woody shrubs in the forest zones of the Tianshan Mountains[J].Acta Ecologica Sinica,2016,36(9):2 695-2 704.
[22]刘江涛,刘立佳.SPSS数据统计与分析应用教程[M].北京:清华大学出版社,2017,74-99;143-147.
[23]PHILIPPE M,CLAUDIO P,MATTHEW P.Limitations to photosynthesis under light and heat stress in three high-yielding wheat genotypes[J].J Plant Physiol,2003,160:657-666.
[24]许克章,史跃林,许贵民,等.保护地黄瓜叶片光合作用温度特性的研究[J].园艺学报,1993,20(1):51-55.XU K Z,SHI Y L,XU G M,et al.Studies on photosynthetic temperature characteristic of cucumber leaves in protective field[J].Acta Horticulturae Sinica,1993,20(1):51-55.
[25]施建敏,郭起荣,杨光耀.毛竹光合动态研究[J].林业科学研究,2005,18(5):551-555.SHI J M,GUO Q R,YANG G Y.Study on thephotosynthetic dynamic variation of Phyllostachys edulis[J].Forest Research,2005,18(5):551-555.
[26]梁开明,曹洪麟,徐志防,等.台湾青枣及野生种的光合作用日变化及光响应特征[J].园艺学报,2008,35(6):793-798.LIANG K M,CAO H L,XU Z F,et al.Diurnal variations and light response of photosynthesis in three cultivars and wild type of Ziziphusm auritiana Lam[J].Acta Horticulturae Sinica,2008,35(6):793-798.
[27]PEARCY RW,SIMS DA.Photosynthetic acclimation to changing light environments:Scaling from the leaf to the whole plant//Caldwell MM,Pearcy RW,eds.Exploitation of environmental heterogeneity by plants:ecophysiological processes above-and belowground.San Diego:Academic Press,1994:145-174.
[28]郑玉龙,姜春玲,冯玉龙.植物的气孔发生[J].植物生理学通讯,2005,41(6):847-850.ZHENG Y L,JIANG C L,FENG Y L.Stomatal generation in plants[J].Plant Physiology Communications,2005,41(6):847-850.
[29]HETHERINGTON A M,WOODWARD F I.The role of stomata in sensing and driving environmental change[J].Nature,2003,424:901-908.
[30]ONWUEME I C,JOHNSTON M.Influence of shade on stomatal density,leaf size and other leaf characteristics in the major tropical root crops,tannia,sweet potato,yam,cassava and taro[J].Experimental Agriculture,2000,36(4):509-516.
[31]徐坤,邹琦,赵燕.土壤水分胁迫与遮光对生姜生长特性的影响[J].应用生态学报,2003,14(10):1 645-1 648.XU K,ZOU Q,ZHAO Y.Effects of soil water stress and shading on growth characteristics of ginger[J].Chinese Journal of Applied Ecolog,2003,14(10):1 645-1 648.
[32]JAMES S A,BELL D T.Influence of light avilability on lesf structure and growth of two Euclyptus ssp.Globulus Provenances[J].Tree physiology,2000,20(15):1 007-1 018.
[33]王荣,郭志华.不同光环境下枫香幼苗的叶片解剖结构[J].生态学杂志,2007,26(11):1 719-1 724.WANG R,GUO Z H.Responses of seedling leaf anatomical structure of Liquidambar formosana,a deciduous broadleaf tree,to different light regimes[J].Chinese Journal of Ecology,2007,26(11):1 719-1 724.
[34]杨文权,褚继鹏,寇建村,等.遮阴对白三叶叶片解剖结构和光合特性的影响[J].草地科学,2015,23(3):653-657.YANG W Q,CHU J P,KOU J C,et al.Effects of shading on the leave anatomical structure and photosynthetic characteristics of white clover[J].Acta Agrestia Sinica,2015,23(3):653-657.