遮阴对骆驼刺叶性状和水分生理的影响
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摘要
以塔克拉玛干沙漠南缘荒漠绿洲过渡带优势种骆驼刺为试验材料,研究正常光照(NL)、中度遮阴(MS)和重度遮阴(SS)3种不同遮阴环境对骆驼刺(Alhagi sparsifolia Shap.)叶厚、单叶面积、比叶面积、比叶质量等叶性状参数和相对含水量、水势、失水速率、气孔导度、蒸腾速率以及水分利用效率等水分生理参数的影响。结果显示:随着光照强度的降低,骆驼刺的叶片厚度(LT)、比叶质量(LMA)、净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用效率(WUE)明显降低,而单叶面积(LA)、比叶面积(SLA)、相对含水量(RWC)、失水速率(RWL)和水势(WP)明显升高;各叶性状参数与水分生理参数之间均有极显著的相关性。说明:骆驼刺在遮阴环境下,其叶性状特征主要通过单叶面积和单叶干物质积累的变化来响应遮阴环境;其水分生理特征主要是牺牲对水分胁迫具有较好抗逆性的水分生理特征并通过气孔调节和更多的水分消耗用于维持一定的光合能力来响应遮阴环境。叶性状与水分生理参数相关性分析表明,遮阴环境下骆驼刺叶性状变化主要通过影响RWL、WP、Tr和WUE进而影响其水分生理特征的变化。因此,建议可利用遮阴措施对极端干旱区荒漠植物骆驼刺进行植被恢复,但其遮光度应设置在70%—80%自然光之间。
The southern edge of the Taklamakan Desert has a typical inland warm temperate desert climate. It is hot in summer,with little rain and drought; it has abundant light and heat,and strong evaporation; and it has a large water deficit. Desert plants in this region are often subjected to multiple stresses of drought,high temperatures,strong light,and wind and sand. Alhagi sparsifolia Shap. represents an important vegetation community on the southern edge of the Taklamakan Desert. Alhagi sparsifolia Shap. plays an important role in mitigating the effects to wind and sand and in maintaining the fragile ecological environment of the Taklamakan Desert. It is,therefore,important to study the adaptability mechanism and strategy of A. sparsifolia under environmental stress conditions such as extreme light and drought for therestoration of degraded vegetation. Current research indicates that A. sparsifolia is heliophyte. Study of its habitats,however,suggested that-when other conditions were the same-its morphology and physiology changed significantly with increased shading( from 100% natural light to 20% natural light),and the plants rarely appeared when shading increased enough.Therefore,this study assumes that the shading environment will have a significant impact on the morphology and physiological ecology of A. sparsifolia and will affect the distribution and succession of vegetation communities as the degree of shading increases. The objectives of this study were to investigate the effects of shade on the leaf traits and water physiological characteristics in A. sparsifolia on the southern edge of the Taklamakan Desert. We conducted three treatments: natural light( NL),moderate shade( MS) and severe shade( SS). The results showed that increasing shading significantly reduced the leaf thickness( LT),leaf mass per unit area( LMA),net photosynthetic rate( Pn),stomatal conductance( Gs),transpiration rate( Tr) and water use efficiency( WUE) in A. sparsifolia,while it increased the individual single leaf area( LA),specific LA( SLA),relative water content( RWC),water loss rate( RWL) and water potential( WP). We concluded that A. sparsifolia changed its leaf traits to adjust to shade by changing the single LA and single leaf dry matter accumulation; changed its water physiological characteristics to adjust to shade by attenuating characteristics that have good resistance to water stress; and adjusted its stomatal regulation and water consumption to maintain a certain photosynthetic capacity. Correlation analysis between the leaf traits and water physiological parameters show that the changes in leaf traits in A. sparsifolia in shade mainly influenced the RWL,WP,Tr and WUE,which then influenced the changes in water physiological characteristics. Therefore,this study recommends protecting and repairing A.sparsifolia in extreme arid regions using shading measures; however,the degree of shade should be between 70% and 80%natural light.
The southern edge of the Taklamakan Desert has a typical inland warm temperate desert climate. It is hot in summer,with little rain and drought; it has abundant light and heat,and strong evaporation; and it has a large water deficit. Desert plants in this region are often subjected to multiple stresses of drought,high temperatures,strong light,and wind and sand. Alhagi sparsifolia Shap. represents an important vegetation community on the southern edge of the Taklamakan Desert. Alhagi sparsifolia Shap. plays an important role in mitigating the effects to wind and sand and in maintaining the fragile ecological environment of the Taklamakan Desert. It is,therefore,important to study the adaptability mechanism and strategy of A. sparsifolia under environmental stress conditions such as extreme light and drought for therestoration of degraded vegetation. Current research indicates that A. sparsifolia is heliophyte. Study of its habitats,however,suggested that-when other conditions were the same-its morphology and physiology changed significantly with increased shading( from 100% natural light to 20% natural light),and the plants rarely appeared when shading increased enough.Therefore,this study assumes that the shading environment will have a significant impact on the morphology and physiological ecology of A. sparsifolia and will affect the distribution and succession of vegetation communities as the degree of shading increases. The objectives of this study were to investigate the effects of shade on the leaf traits and water physiological characteristics in A. sparsifolia on the southern edge of the Taklamakan Desert. We conducted three treatments: natural light( NL),moderate shade( MS) and severe shade( SS). The results showed that increasing shading significantly reduced the leaf thickness( LT),leaf mass per unit area( LMA),net photosynthetic rate( Pn),stomatal conductance( Gs),transpiration rate( Tr) and water use efficiency( WUE) in A. sparsifolia,while it increased the individual single leaf area( LA),specific LA( SLA),relative water content( RWC),water loss rate( RWL) and water potential( WP). We concluded that A. sparsifolia changed its leaf traits to adjust to shade by changing the single LA and single leaf dry matter accumulation; changed its water physiological characteristics to adjust to shade by attenuating characteristics that have good resistance to water stress; and adjusted its stomatal regulation and water consumption to maintain a certain photosynthetic capacity. Correlation analysis between the leaf traits and water physiological parameters show that the changes in leaf traits in A. sparsifolia in shade mainly influenced the RWL,WP,Tr and WUE,which then influenced the changes in water physiological characteristics. Therefore,this study recommends protecting and repairing A.sparsifolia in extreme arid regions using shading measures; however,the degree of shade should be between 70% and 80%natural light.
引文
[1]刘建锋,杨文娟,江泽平,郭泉水,金江群,薛亮.遮荫对濒危植物崖柏光合作用和叶绿素荧光参数的影响.生态学报,2011,31(20):5999-6004.
[2]Sofo A,Dichio B,Montanaro G,Xiloyannis C.Shade effect on photosynthesis and photoinhibition in olive during drought and rewatering.Agricultural Water Management,2009,96(8):1201-1206.
[3]Dai Y J,Shen Z G,Ying L,Wang L L,Hannaway D,Lu H F.Effects of shade treatments on the photosynthetic capacity,chlorophyll fluorescence,and chlorophyll content of Tetrastigma hemsleyanum Diels et Gilg.Environmental and Experimental Botany,2009,65(2/3):177-182.
[4]Sarijeva G,Knapp M,Lichtenthaler H K.Differences in photosynthetic activity,chlorophyll and carotenoid levels,and in chlorophyll fluorescence parameters in green sun and shade leaves of Ginkgo and Fagus.Journal of Plant Physiology,2007,164(7):950-955.
[5]张林,罗天祥.植物叶寿命及其相关叶性状的生态学研究进展.植物生态学报,2004,28(6):844-852.
[6]秦舒浩,李玲玲.遮光处理对西葫芦幼苗形态特征及光合生理特性的影响.应用生态学报,2006,17(4):653-656.
[7]王蕊,孙备,李建东,王国骄,孙佳楠,王欣若,钟日亭.不同光强对入侵种三裂叶豚草表型可塑性的影响.应用生态学报,2012,23(7):1797-1802.
[8]涂璟,王克勤.干旱地区造林树种的水分生理生态的研究进展.西北林学院学报,2003,18(3):26-30.
[9]张硕,刘勇,李国雷,陈晓,孙巧玉,许飞.稳定碳同位素在森林植物水分利用效率研究中的应用.世界林业研究,2013,26(3):39-45.
[10]刘建平,李志军,韩路,龚卫江,周正立.胡杨、灰叶胡杨P-V曲线水分参数的初步研究.西北植物学报,2004,24(7):1255-1259.
[11]蔡海霞,吴福忠,杨万勤.干旱胁迫对高山柳和沙棘幼苗光合生理特征的影响.生态学报,2011,31(9):2430-2436.
[12]Liu F,Stützel H.Biomass partitioning,specific leaf area,and water use efficiency of vegetable amaranth(Amaranthus spp.)in response to drought stress.Scientia Horticulturae,2004,102(1):15-27.
[13]张晓蕾,曾凡江,刘波,张慧,刘镇,安桂香.不同地下水埋深下骆驼刺幼苗叶片生理参数光响应特性.干旱区地理,2011,34(2):229-235.
[14]罗维成,曾凡江,刘波,张利刚,宋聪,彭守兰,Arndt S K.疏叶骆驼刺根系对土壤异质性和种间竞争的响应.植物生态学报,2012,36(10):1015-1023.
[15]李磊.温度及光强对疏叶骆驼刺表型可塑性及光系统II活性的影响[D].乌鲁木齐:中国科学院新疆生态与地理研究所,2015.
[16]薛伟,李向义,朱军涛,林丽莎,王迎菊.遮阴对疏叶骆驼刺叶形态和光合参数的影响.植物生态学报,2011,35(1):82-90.
[17]张晓蕾,曾凡江,刘波,刘镇,安桂香,孙旭伟.不同土壤水分处理对疏叶骆驼刺幼苗光合特性及干物质积累的影响.干旱区研究,2010,27(4):649-655.
[18]朱军涛,李向义,张希明,曾凡江,闫海龙,杨尚功.灌溉对疏叶骆驼刺(Alhagi sparsifolia)幼苗光合生理指标及渗透物质的影响.中国沙漠,2009,29(4):697-702.
[19]李向义,张希明,何兴元,曾凡江,Foetzki A,Thomas F M.沙漠-绿洲过渡带四种多年生植物水分关系特征.生态学报,2004,24(6):1164-1171.
[20]唐钢梁,李向义,林丽莎,李磊,鲁建荣.骆驼刺在不同遮阴下的水分状况变化及其生理响应.植物生态学报,2013,37(4):354-364.
[21]Reich P B,Buschena C,Tjoelker M G,Wrage K,Knops J,Tilman D,Machado J L.Variation in growth rate and ecophysiology among 34grassland and savanna species under contrasting N supply:a test of functional group differences.New Phytologist,2010,157(3):617-631.
[22]Ackerly D,Knight C,Weiss S,Barton K,Starmer K.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:contrasting patterns in species level and community level analyses.Oecologia,2002,130(3):449-457.
[23]Davi H,Barbaroux C,Dufrêne E,Fran9ois C,Montpied P,Bréda N,Badeck F.Modelling leaf mass per area in forest canopy as affected by prevailing radiation conditions.Ecological Modelling,2008,211(3/4):339-349.
[24]李芳兰,包维楷.植物叶片形态解剖结构对环境变化的响应与适应.植物学通报,2005,22(S1):118-127.
[25]Lusk C H,Reich P B,Montgomery R A,Ackerly D D,Cavender-Bares J.Why are evergreen leaves so contrary about shade?Trends in Ecology&Evolution,2008,23(6):299-303.
[26]付爱红,陈亚宁,李卫红,张宏锋.干旱、盐胁迫下的植物水势研究与进展.中国沙漠,2005,25(5):744-749.
[27]González-Rodríguez A M,Martín-Olivera A,Morales D,Jiménez M S.Physiological responses of tagasaste to a progressive drought in its native environment on the Canary Islands.Environmental and Experimental Botany,2005,53(2):195-204.
[28]魏晓霞,呼和牧仁,周梅,翟洪波,李良,邵仁旭,宋吉祥.不同年龄华北落叶松叶水势及其影响因素的研究.干旱区资源与环境,2010,24(7):144-148.
[29]罗永忠,成自勇.水分胁迫对紫花苜蓿叶水势、蒸腾速率和气孔导度的影响.草地学报,2011,19(2):215-221.
[30]Sack,L,Cowan P D,Jaikumar N,Holbrook N M.The‘hydrology’of leaves:co-ordination of structure and function in temperate woody species.Plant,Cell&Environment,2003,26(8):1343-1356.
[2]Sofo A,Dichio B,Montanaro G,Xiloyannis C.Shade effect on photosynthesis and photoinhibition in olive during drought and rewatering.Agricultural Water Management,2009,96(8):1201-1206.
[3]Dai Y J,Shen Z G,Ying L,Wang L L,Hannaway D,Lu H F.Effects of shade treatments on the photosynthetic capacity,chlorophyll fluorescence,and chlorophyll content of Tetrastigma hemsleyanum Diels et Gilg.Environmental and Experimental Botany,2009,65(2/3):177-182.
[4]Sarijeva G,Knapp M,Lichtenthaler H K.Differences in photosynthetic activity,chlorophyll and carotenoid levels,and in chlorophyll fluorescence parameters in green sun and shade leaves of Ginkgo and Fagus.Journal of Plant Physiology,2007,164(7):950-955.
[5]张林,罗天祥.植物叶寿命及其相关叶性状的生态学研究进展.植物生态学报,2004,28(6):844-852.
[6]秦舒浩,李玲玲.遮光处理对西葫芦幼苗形态特征及光合生理特性的影响.应用生态学报,2006,17(4):653-656.
[7]王蕊,孙备,李建东,王国骄,孙佳楠,王欣若,钟日亭.不同光强对入侵种三裂叶豚草表型可塑性的影响.应用生态学报,2012,23(7):1797-1802.
[8]涂璟,王克勤.干旱地区造林树种的水分生理生态的研究进展.西北林学院学报,2003,18(3):26-30.
[9]张硕,刘勇,李国雷,陈晓,孙巧玉,许飞.稳定碳同位素在森林植物水分利用效率研究中的应用.世界林业研究,2013,26(3):39-45.
[10]刘建平,李志军,韩路,龚卫江,周正立.胡杨、灰叶胡杨P-V曲线水分参数的初步研究.西北植物学报,2004,24(7):1255-1259.
[11]蔡海霞,吴福忠,杨万勤.干旱胁迫对高山柳和沙棘幼苗光合生理特征的影响.生态学报,2011,31(9):2430-2436.
[12]Liu F,Stützel H.Biomass partitioning,specific leaf area,and water use efficiency of vegetable amaranth(Amaranthus spp.)in response to drought stress.Scientia Horticulturae,2004,102(1):15-27.
[13]张晓蕾,曾凡江,刘波,张慧,刘镇,安桂香.不同地下水埋深下骆驼刺幼苗叶片生理参数光响应特性.干旱区地理,2011,34(2):229-235.
[14]罗维成,曾凡江,刘波,张利刚,宋聪,彭守兰,Arndt S K.疏叶骆驼刺根系对土壤异质性和种间竞争的响应.植物生态学报,2012,36(10):1015-1023.
[15]李磊.温度及光强对疏叶骆驼刺表型可塑性及光系统II活性的影响[D].乌鲁木齐:中国科学院新疆生态与地理研究所,2015.
[16]薛伟,李向义,朱军涛,林丽莎,王迎菊.遮阴对疏叶骆驼刺叶形态和光合参数的影响.植物生态学报,2011,35(1):82-90.
[17]张晓蕾,曾凡江,刘波,刘镇,安桂香,孙旭伟.不同土壤水分处理对疏叶骆驼刺幼苗光合特性及干物质积累的影响.干旱区研究,2010,27(4):649-655.
[18]朱军涛,李向义,张希明,曾凡江,闫海龙,杨尚功.灌溉对疏叶骆驼刺(Alhagi sparsifolia)幼苗光合生理指标及渗透物质的影响.中国沙漠,2009,29(4):697-702.
[19]李向义,张希明,何兴元,曾凡江,Foetzki A,Thomas F M.沙漠-绿洲过渡带四种多年生植物水分关系特征.生态学报,2004,24(6):1164-1171.
[20]唐钢梁,李向义,林丽莎,李磊,鲁建荣.骆驼刺在不同遮阴下的水分状况变化及其生理响应.植物生态学报,2013,37(4):354-364.
[21]Reich P B,Buschena C,Tjoelker M G,Wrage K,Knops J,Tilman D,Machado J L.Variation in growth rate and ecophysiology among 34grassland and savanna species under contrasting N supply:a test of functional group differences.New Phytologist,2010,157(3):617-631.
[22]Ackerly D,Knight C,Weiss S,Barton K,Starmer K.Leaf size,specific leaf area and microhabitat distribution of chaparral woody plants:contrasting patterns in species level and community level analyses.Oecologia,2002,130(3):449-457.
[23]Davi H,Barbaroux C,Dufrêne E,Fran9ois C,Montpied P,Bréda N,Badeck F.Modelling leaf mass per area in forest canopy as affected by prevailing radiation conditions.Ecological Modelling,2008,211(3/4):339-349.
[24]李芳兰,包维楷.植物叶片形态解剖结构对环境变化的响应与适应.植物学通报,2005,22(S1):118-127.
[25]Lusk C H,Reich P B,Montgomery R A,Ackerly D D,Cavender-Bares J.Why are evergreen leaves so contrary about shade?Trends in Ecology&Evolution,2008,23(6):299-303.
[26]付爱红,陈亚宁,李卫红,张宏锋.干旱、盐胁迫下的植物水势研究与进展.中国沙漠,2005,25(5):744-749.
[27]González-Rodríguez A M,Martín-Olivera A,Morales D,Jiménez M S.Physiological responses of tagasaste to a progressive drought in its native environment on the Canary Islands.Environmental and Experimental Botany,2005,53(2):195-204.
[28]魏晓霞,呼和牧仁,周梅,翟洪波,李良,邵仁旭,宋吉祥.不同年龄华北落叶松叶水势及其影响因素的研究.干旱区资源与环境,2010,24(7):144-148.
[29]罗永忠,成自勇.水分胁迫对紫花苜蓿叶水势、蒸腾速率和气孔导度的影响.草地学报,2011,19(2):215-221.
[30]Sack,L,Cowan P D,Jaikumar N,Holbrook N M.The‘hydrology’of leaves:co-ordination of structure and function in temperate woody species.Plant,Cell&Environment,2003,26(8):1343-1356.