海岸梯度上黑松针叶化学计量特征的变化规律
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摘要
风对树木形态、生理和生长的影响一直是生态学研究的热点和难点问题,目前主要采用模拟风速或机械刺激的方式来研究树木对风胁迫的响应与适应变化,无法准确反映出树木在风环境下的长期适应机制。揭示自然环境下树木对风胁迫的长期适应机制,测定了海岸梯度上迎风面和背风面60年生黑松林(Pinus thunbergii)针叶有机碳、氮、磷、钾、钙、钠、镁元素含量,并分析元素含量及其化学计量特征的变化规律。结果表明:黑松林迎风面针叶有机碳含量随离海岸距离的减小显著降低(P<0.05);而氮、磷、钾、钙、钠、镁含量则显著增加(P<0.05);氮磷比保持相对稳定。黑松林背风面针叶除钙和钠含量随离海岸距离的减小逐渐增加外,其他元素含量变化规律均不明显;距海岸500 m内黑松针叶化学计量特征差异显著(P<0.05),其后趋于相对稳定。黑松林迎风面针叶化学计量特征普遍存在相关关系,背风面仅针叶氮和磷、钙和钠间存在相关关系,且与迎风面存在共同的SMA斜率和截距。研究表明,海岸梯度上风胁迫会显著改变黑松针叶化学元素含量,但其比值相对稳定;随海岸距离减少,风胁迫增加,针叶碳同化能力受到明显影响,黑松通过针叶氮、磷、钾、钙、钠、镁含量的增加来提高耐风胁迫的能力,这可能是自然环境下黑松对长期风胁迫的一种适应策略。
The effects of wind on the morphology,physiology and growth of trees have been a hot topic and difficulty in ecology. Previous studies examined response and adaptation of trees to wind through short-time wind simulation or mechanical stimulation,which cannot reflect the long-term adaptation mechanism of trees in the natural windy environment. To fill such knowledge gap,we measured the concentrations of carbon,nitrogen,phosphorus,potassium,calcium,sodium and magnesium in needles of a 60-year-old Pinus thunbergii forest along a coastal-inland gradient,and analyzed the variations of element concentration and stoichiometric characteristics. The results showed that organic carbon concentration in needle on windward of P. thunbergii decreased from inland to coast,the concentrations of nitrogen,phosphorus,potassium,calcium,sodium and magnesium increased( P<0.05),and N ∶ P ratio remained stable. However,there were no significant differences for these variables on leeward along the coastal-inland gradient,except that calcium and sodium concentrations decreased. Needle stoichiometric characters were significantly different between windward and leeward within 500 m from coastline,and then kept stable over distance. The correlations among windward needle stoichiometry were pervasive,but on the leeward side,only significant relationships between nitrogen and phosphorus and between calcium and sodium existed. Standardized major axis( SMA) fitted between windward and leeward were not different in slope and intercept( P > 0.05). In sum,the needle chemical composition of P.thunbergii was significantly affected by wind stress,but the ratio was relatively stable. The needle carbon assimilation function was significantly affected with the increases of sea wind stress. Meanwhile,P. thunbergii improved its tolerance to wind stress by increasing needle nitrogen,phosphorus,potassium,calcium,sodium and magnesium concentrations,which may be an adaptive strategy for P. thunbergii to long-term wind stress in natural environment.
The effects of wind on the morphology,physiology and growth of trees have been a hot topic and difficulty in ecology. Previous studies examined response and adaptation of trees to wind through short-time wind simulation or mechanical stimulation,which cannot reflect the long-term adaptation mechanism of trees in the natural windy environment. To fill such knowledge gap,we measured the concentrations of carbon,nitrogen,phosphorus,potassium,calcium,sodium and magnesium in needles of a 60-year-old Pinus thunbergii forest along a coastal-inland gradient,and analyzed the variations of element concentration and stoichiometric characteristics. The results showed that organic carbon concentration in needle on windward of P. thunbergii decreased from inland to coast,the concentrations of nitrogen,phosphorus,potassium,calcium,sodium and magnesium increased( P<0.05),and N ∶ P ratio remained stable. However,there were no significant differences for these variables on leeward along the coastal-inland gradient,except that calcium and sodium concentrations decreased. Needle stoichiometric characters were significantly different between windward and leeward within 500 m from coastline,and then kept stable over distance. The correlations among windward needle stoichiometry were pervasive,but on the leeward side,only significant relationships between nitrogen and phosphorus and between calcium and sodium existed. Standardized major axis( SMA) fitted between windward and leeward were not different in slope and intercept( P > 0.05). In sum,the needle chemical composition of P.thunbergii was significantly affected by wind stress,but the ratio was relatively stable. The needle carbon assimilation function was significantly affected with the increases of sea wind stress. Meanwhile,P. thunbergii improved its tolerance to wind stress by increasing needle nitrogen,phosphorus,potassium,calcium,sodium and magnesium concentrations,which may be an adaptive strategy for P. thunbergii to long-term wind stress in natural environment.
引文
鲍士旦.2007.土壤农化分析.北京:中国农业出版社.
韩广轩,王光美,张志东,等.2008.烟台海岸黑松防护林种群结构及其随离岸距离的变化.林业科学,44(10):8-13.
季子敬,全先奎,王传宽.2013.兴安落叶松针叶解剖结构变化及其光合能力对气候变化的适应性.生态学报,33(21):6967-6974.
郎志红.2008.盐碱胁迫对植物种子萌发和幼苗生长的影响(硕士学位论文).兰州:兰州交通大学.
李雪峰,欧阳玉祝,张晓旭,等.2015.火焰原子吸收分光光度法测定野葛根中5种金属元素含量.应用化工,44(5):963-966.
毛伟,李玉霖,张铜会,等.2012.不同尺度生态学中植物叶性状研究概述.中国沙漠,32(1):33-41.
王林权,邵明安.2005.高等植物对钠离子的吸收、运输和累积.干旱地区农业研究,23(5):244-249.
肖遥,陶冶,张元明.2014.古尔班通古特沙漠4种荒漠草本植物不同生长期的生物量分配与叶片化学计量特征.植物生态学报,38(9):929-940.
许景伟,李传荣,王卫东,等.2005.沿海沙质岸黑松防护林的生物量及生产力.东北林业大学学报,33(6):29-32.
张丹,李传荣,许景伟,等.2011.沙质海岸黑松分枝格局特征及其抗风折能力分析.植物生态学报,35(9):926-936.
张志东,韩广轩,毛培利,等.2009.成年个体密度、距海远近及下层植被对烟台黑松海防林天然更新的影响.自然资源学报,24(5):782-790.
赵哈林,何玉惠,岳广阳,等.2010.风吹、沙埋对沙地植物幼苗生长和光合蒸腾特性的影响.生态学杂志,29(3):413-419.
Anten NPR,Alcala-Herrera R,Schieving F,et al.2010.Wind and mechanical stimuli differentially affect leaf traits in Plantago major.New Phytologist,188:554-564.
Buchanan B,Gruissem W,Jones R.2000.Biochemistry and Molecular Biology of Plants.American Society of Plant Physiologists:Rockeweel,MD,USA.
Chen Y,Han W,Tang L,et al.2013.Leaf nitrogen and phosphorus concentrations of woody plants differ in responses to climate,soil and plant growth form.Ecography,36:178-184.
Cordero RA.1999.Ecophysiology of Cecropia schreberiana saplings in two wind regimes in an elfin cloud forest:Growth,gas exchange,architecture and stem biomechanics.Tree Physiology,19:153-163.
Coutand C,Dupraz C,Jaouen G,et al.2008.Mechanical stimuli regulate the allocation of biomass in trees:Demonstration with young Prunus avium trees.Annals of Botany,101:1421-1432.
Hidaka A,Kitayama K.2015.Physiological linkage in co-variation of foliar nitrogen and phosphorus in tropical tree species along a gradient of soil phosphorus availability.Journal of Tropical Ecology,31:221-229.
Huang P,Wan X,Lieffers VJ.2016.Daytime and nighttime wind differentially affects hydraulic properties and thigmomorphogenic response of poplar saplings.Physiologia Plantarum,157:85-94.
Jaffe MJ.1973.Thigmomorphogenesis:The response of plant growth and development to mechanical stimulation:With special reference to Bryonia dioica.Planta,114:143-157.
Knight MR,Smith SM,Trewavas AJ.1992.Wind-induced plant motion immediately increases cytosolic calcium.Proceedings of the National Academy of Sciences of the United States of America,89:4967-4971.
Mcdowell NG.2011.Mechanisms linking drought,hydraulics,carbon metabolism,and vegetation mortality.Plant Physiology,155:1051-1059.
Murren CJ,Pigliucci M.2005.Morphological responses to simulated wind in the genus Brassica(Brassicaceae):Allopolyploids and their parental species.American Journal of Botany,92:810-818.
Niklas KJ.1996.Differences between Acer saccharum leaves from open and wind-protected sites.Annals of Botany,78:61-66.
Onoda Y,Hikosaka K,Hirose T.2004.Allocation of nitrogen to cell walls decreases photosynthetic nitrogen-use efficiency.Functional Ecology,18:419-425.
Sadiqov ST,Akbulut M,Ehmedov V.2002.Role of Ca2+in drought stress signaling in wheat seedlings.Biochemistry Biokhimiia,67:491-497.
Schymanski SJ,Or D.2016.Wind increases leaf water use efficiency.Plant,Cell and Environment,39:1448-1459.
Shi Z,Tang J,Cheng R,et al.2015.A review of nitrogen allocation in leaves and factors in its effects.Acta Ecologica Sinica,35:5909-5919.
Smith VC,Ennos AR.2003.The effects of air flow and stem flexure on the mechanical and hydraulic properties of the stems of sunflowers Helianthus annuus L.Journal of Experimental Botany,54:845-849.
Woodward FI.1986.Ecophysiological studies on the shrub Vaccinium myrtillus L.taken from a wide altitudinal range.Oecologia,70:580-586.
Washusen R,Baker T,Menz D,et al.2005.Effect of thinning and fertilizer on the cellulose crystallite width of Eucalyptus globulus.Wood Science&Technology,39:569-578.
Wu TG,Yu MK,Wang GG,et al.2012.Leaf nitrogen and phosphorus stoichiometry across forty-two woody species in Southeast China.Biochemical Systematics&Ecology,44:255-263.
Zhang H,Guo WH,Wang GG,et al.2016.Effects of environment and genetics on leaf N and P stoichiometry for Quercus acutissima across China.European Journal of Forest Research,135:795-802.
Zhang P,Wang H,Wu QT,et al.2018.Effect of wind on the relation of leaf N,P stoichiometry with leaf morphology in Quercus species.Forests,9:110.
韩广轩,王光美,张志东,等.2008.烟台海岸黑松防护林种群结构及其随离岸距离的变化.林业科学,44(10):8-13.
季子敬,全先奎,王传宽.2013.兴安落叶松针叶解剖结构变化及其光合能力对气候变化的适应性.生态学报,33(21):6967-6974.
郎志红.2008.盐碱胁迫对植物种子萌发和幼苗生长的影响(硕士学位论文).兰州:兰州交通大学.
李雪峰,欧阳玉祝,张晓旭,等.2015.火焰原子吸收分光光度法测定野葛根中5种金属元素含量.应用化工,44(5):963-966.
毛伟,李玉霖,张铜会,等.2012.不同尺度生态学中植物叶性状研究概述.中国沙漠,32(1):33-41.
王林权,邵明安.2005.高等植物对钠离子的吸收、运输和累积.干旱地区农业研究,23(5):244-249.
肖遥,陶冶,张元明.2014.古尔班通古特沙漠4种荒漠草本植物不同生长期的生物量分配与叶片化学计量特征.植物生态学报,38(9):929-940.
许景伟,李传荣,王卫东,等.2005.沿海沙质岸黑松防护林的生物量及生产力.东北林业大学学报,33(6):29-32.
张丹,李传荣,许景伟,等.2011.沙质海岸黑松分枝格局特征及其抗风折能力分析.植物生态学报,35(9):926-936.
张志东,韩广轩,毛培利,等.2009.成年个体密度、距海远近及下层植被对烟台黑松海防林天然更新的影响.自然资源学报,24(5):782-790.
赵哈林,何玉惠,岳广阳,等.2010.风吹、沙埋对沙地植物幼苗生长和光合蒸腾特性的影响.生态学杂志,29(3):413-419.
Anten NPR,Alcala-Herrera R,Schieving F,et al.2010.Wind and mechanical stimuli differentially affect leaf traits in Plantago major.New Phytologist,188:554-564.
Buchanan B,Gruissem W,Jones R.2000.Biochemistry and Molecular Biology of Plants.American Society of Plant Physiologists:Rockeweel,MD,USA.
Chen Y,Han W,Tang L,et al.2013.Leaf nitrogen and phosphorus concentrations of woody plants differ in responses to climate,soil and plant growth form.Ecography,36:178-184.
Cordero RA.1999.Ecophysiology of Cecropia schreberiana saplings in two wind regimes in an elfin cloud forest:Growth,gas exchange,architecture and stem biomechanics.Tree Physiology,19:153-163.
Coutand C,Dupraz C,Jaouen G,et al.2008.Mechanical stimuli regulate the allocation of biomass in trees:Demonstration with young Prunus avium trees.Annals of Botany,101:1421-1432.
Hidaka A,Kitayama K.2015.Physiological linkage in co-variation of foliar nitrogen and phosphorus in tropical tree species along a gradient of soil phosphorus availability.Journal of Tropical Ecology,31:221-229.
Huang P,Wan X,Lieffers VJ.2016.Daytime and nighttime wind differentially affects hydraulic properties and thigmomorphogenic response of poplar saplings.Physiologia Plantarum,157:85-94.
Jaffe MJ.1973.Thigmomorphogenesis:The response of plant growth and development to mechanical stimulation:With special reference to Bryonia dioica.Planta,114:143-157.
Knight MR,Smith SM,Trewavas AJ.1992.Wind-induced plant motion immediately increases cytosolic calcium.Proceedings of the National Academy of Sciences of the United States of America,89:4967-4971.
Mcdowell NG.2011.Mechanisms linking drought,hydraulics,carbon metabolism,and vegetation mortality.Plant Physiology,155:1051-1059.
Murren CJ,Pigliucci M.2005.Morphological responses to simulated wind in the genus Brassica(Brassicaceae):Allopolyploids and their parental species.American Journal of Botany,92:810-818.
Niklas KJ.1996.Differences between Acer saccharum leaves from open and wind-protected sites.Annals of Botany,78:61-66.
Onoda Y,Hikosaka K,Hirose T.2004.Allocation of nitrogen to cell walls decreases photosynthetic nitrogen-use efficiency.Functional Ecology,18:419-425.
Sadiqov ST,Akbulut M,Ehmedov V.2002.Role of Ca2+in drought stress signaling in wheat seedlings.Biochemistry Biokhimiia,67:491-497.
Schymanski SJ,Or D.2016.Wind increases leaf water use efficiency.Plant,Cell and Environment,39:1448-1459.
Shi Z,Tang J,Cheng R,et al.2015.A review of nitrogen allocation in leaves and factors in its effects.Acta Ecologica Sinica,35:5909-5919.
Smith VC,Ennos AR.2003.The effects of air flow and stem flexure on the mechanical and hydraulic properties of the stems of sunflowers Helianthus annuus L.Journal of Experimental Botany,54:845-849.
Woodward FI.1986.Ecophysiological studies on the shrub Vaccinium myrtillus L.taken from a wide altitudinal range.Oecologia,70:580-586.
Washusen R,Baker T,Menz D,et al.2005.Effect of thinning and fertilizer on the cellulose crystallite width of Eucalyptus globulus.Wood Science&Technology,39:569-578.
Wu TG,Yu MK,Wang GG,et al.2012.Leaf nitrogen and phosphorus stoichiometry across forty-two woody species in Southeast China.Biochemical Systematics&Ecology,44:255-263.
Zhang H,Guo WH,Wang GG,et al.2016.Effects of environment and genetics on leaf N and P stoichiometry for Quercus acutissima across China.European Journal of Forest Research,135:795-802.
Zhang P,Wang H,Wu QT,et al.2018.Effect of wind on the relation of leaf N,P stoichiometry with leaf morphology in Quercus species.Forests,9:110.