海拔对毛竹林结构及生理生态学特性的影响研究
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摘要
本文对不同海拔梯度毛竹(Phyllostachys edulis)林主要气象因子变化和土壤理化性质、种群结构、经济性状、光合生理等方面进行了较为系统的研究,揭示了不同海拔梯度毛竹林生长更新规律,探讨了主要环境因子对毛竹生理生态过程的作用机理及其生态适应机制。主要结论如下:
1.随着海拔的升高,Tair明显下降,在3个海拔梯度不同季节,海拔升高100m的降温幅度无显著差异;RH较大辐度提高,以季节性干旱的秋冬季增湿作用强;PAR一定幅度增强;海拔升高100m,Tair下降0.61～0.69℃,RH提高1.42～2.47%,PAR增大23.61～60.36μmol·m~(-2)·s~(-1)。
2.同一海拔梯度毛竹林土壤容重、总孔隙度、非毛管孔隙度、毛管孔隙度、饱和持水量、田间持水量、粘粒和砂粒含量随土层加深而增大,粉粒含量减少。总孔隙度与容重呈负相关;海拔对土壤容重、毛管孔隙度、田间持水量无显著影响,总孔隙度、非毛管孔隙度、饱和持水量随海拔升高而显著增大;不同海拔梯度毛竹林土壤微结构颗粒组成砂粒＞粉粒＞粘粒,粘粒、粉粒含量随海拔升高而减少,砂粒含量趋于增加。
3.海拔与土壤pH值、全氮、速效氮呈极显著负相关,与有机质呈极显著正相关,对全磷、速效磷、全钾、速效钾影响不显著;土壤酶活性与海拔呈正相关,蔗糖酶、脲酶、磷酸酶在中海拔发生转折性变化,蛋白酶随海拔升高缓慢提高,过氧化氢酶呈均匀上升趋势。土壤物理性质对土壤酶活性无显著影响。有机质、全氮、速效氮与土壤酶活性呈显著或极显著正相关,pH值对土壤酶活性具正效应,对全磷、全钾、速磷、速钾影响不显著。不同种类土壤酶活性间呈显著或极显著相关,酶促反应具专一性和共同性特点。
4.立竹密度、立竹胸径、枝下高、全高随海拔升高显著增大,立竹整齐度、均匀度显著下降,种群个体生长存在“利己”行为;始出笋时间、出笋终止时间随海拔升高向后推移,出笋持续时间随海拔升高而缩短;退笋率随海拔升高而降低;竹笋——幼竹高生长划分为初期、上升期、盛期、末期4个时期,遵循“慢——快——慢”节律。高生长差异主要体现在随海拔升高而增大的盛期日平均高生长量上,相同出笋阶段的竹笋高生长期随海拔升高而延长。相同海拔各出笋时期竹笋高生长期前期最长,后期最短,高生长时间接近,具有“利他”性生态适应机制。
5.海拔与立竹胸高壁厚、壁厚率呈负相关。相同径级立竹重量、材积随海拔升高而降低,竹林竹材产量、材积随海拔升高而增长;立竹尖削度值随海拔升高显著增大;编制了不同海拔梯度毛竹林立竹材积与胸径的对应表;竹笋产量、品质随海拔升高而提高,海拔对竹笋品质影响主要反映在总糖、可溶性糖、P、Fe、含水率、可食比例随海拔升高而提高,纤维、热量、灰分、Ca随海拔升高而降低。
6.不同年龄立竹竹叶叶绿素含量随海拔升高有差异不显著的提高;Pn随海拔升高而增大,夏季＞秋季＞冬季＞春季,夏秋季中、低海拔呈双峰型,出现光合“午休”,高海拔毛竹林和冬季、春季呈单峰型;不同海拔毛竹林Tr夏季＞秋季＞春季＞冬季,春季、秋季、冬季呈单峰型,夏季呈双峰型,春季、夏季随海拔升高而降低,秋冬季变化趋势相反;WUE春季、夏季随海拔升高而提高,秋冬季变化不明显,季节性水分亏缺能提高WUE。
7.PAR、Tair、RH是影响毛竹光合的主要环境因子,Tair、RH分别与Ci、Cond极显著负相关和正相关,PAR与Pn、Tr、Vpdl极显著正相关,与Ci极显著负相关。环境因子通过Cond影响毛竹光合作用,对Cond产生显著影响的是夏季、初秋的高温和秋冬季的季节性水分亏缺,光合“午休”由气孔限制引起;不同海拔毛竹林LCP春季＞夏季＞秋季＞冬季,LSP秋季＞夏季＞冬季＞春季,均随海拔升高而降低。P_(max)夏季＞秋季＞冬季＞春季,夏季、秋季随海拔升高而增大,春季、冬季呈相反变化趋势,年平均P_(max)随海拔升高而增大。AQY秋季、夏季差异小,高于冬季、春季,随海拔升高而增大。季节性水分亏缺及春季竹叶老化是影响AQY的主要原因。
In the dissertation,the author has investigated changes of main weather factors, soil properties,population structure,economic characters,photosynthetic physiology of Moso bamboo forest at different altitudes.The growth and regeneration characteristics,effects of main environmental factors and adaptive mechanism were also revealed.The main results were described as follows:
1.With the increase of the altitude,Tair definitely decreased.When the altitude increased for 100m,there is no notable decrease range of temperature in three altitude grads among different seasons.RH has a bigger increment with the humidity increasing most obviously both in Autumn and Winter.PAR is also high in some scale.With the increase of altitude for 100m.Tair decreased at a range from 0.61℃to 0.69℃,RH increased at a range from 1.42%to 2.47%,and PAR increased at an extent from 23.61μmol·m~(-2)·s~(-1) to 60.36μmol·m~(-2)·s~(-1).The weather is more suitable for the growth of Moso bamboo forest when the altitude increased.
2.At the same altitude,with the increase of soil depth,soil bulk density,total porosity,non-capillary porosity,capillary porosity,saturated water capacity,field water-holding capacity,clay and sand particle content increased,but silt content decreased.There existed a negative correlation between total porosity and soil bulk density.Altitude had no significant effect on soil bulk density,capillary porosity, field water-holding capacity.With the increase of altitude,total porosity, non-capillary porosity,saturated water capacity increased significantly.Particle composition of soil in different altitudes varied in such a sequence:sand＞silt＞clay particle.With the increase of altitude,clay and silt particle content decreased,but sand content increased gradually.
3.There were a highly significant negative correlation among altitude and soil pH,total nitrogen and available nitrogen content,and a significant positive correlation between altitude and organic matter,but there was no significant effect on the contents of total phosphorus,available phosphorus,total potassium,and available potassium.There was a positive correlation between altitude and soil enzyme activities.At middle level of altitude,the great changes took place in phosphatase,urease and invertase.With the increase of altitude,protease activities slightly increased,and calatase activities also rise smoothly.Soil physical properties had no significant effect on soil enzyme activities.Soil enzyme activities had significant or definitely significant correlation with organic matter,total nitrogen and available nitrogen,and no significant correlation with total phosphorus, available phosphorus,total potassium,available potassium.There was a significant correlation among soil enzyme activities with specifiying and concomitant characteristics.
4.With the increase of altitude,the density,DBH,height under branch and total height increased significantly,but uniformity and evenness decreased significantly. Individual growth showed egoistic behavior to some extent.With increasing altitude, the time of beginning and termination of shooting were backward,whereas,the time of duration and degenerated shoot ratio decreased.The duration of earlier stage and later stage of shoot emergence was similar both at middle and lower altitude,but longer than that of at higher altitude.Height growth period from shoot to young culm could be divided into four stages,such as initial stage,rising stage,rapid growth stage and telophase,which showed a rhythm of slow growth followed by fast growth.With the increasing altitude,the difference of height growth existed mainly in the increase of mean height growth increment at rapid growth stage.The height growth period of shoot at the same stage was delayed and the duration of height growth shortened.Among all shooting stages,the earlier stage was longest,and the later stage was shortest at the same altitude.The high growth nearly ended in the same time which exhibited ecological adaptive mechanism of altruism.
5.There was a negative correlation between altitude and cell wall thickness at DBH,wall thickness ratio.With the increasing altitude,the weight and timber volume of Moso bamboo with the same DBH decreased,while the timber yield and volume increased.There was no significant difference in timber volume between middle and higher altitude due to multiple changes of structure factors of bamboo stands.With the increasing altitude,taper increased greatly.The corresponding table was established which could reflected the relationship between timber volume and DBH of bamboo in different altitude grads.With increase of altitude,the yield and quality of shoots enhanced.Eeffects of altitude on the quality of bamboo shoots depended on the rise of total sugar content,soluble sugar content,P,Fe,water content and edible proportion of shoot.But heat,fibre content,ash content and Ca all decreased with increasing altitude.
6.With the increasing altitude,the leaf chlorophyll content of Moso bamboo with different ages represented a small increasing trend,and Pn also increased in the sequence of Summer＞Autumn＞Winter＞Spring.At lower and middle altitude,Pn showed a double-peak curve,and midday depression of photosynthesis rate took place.But at higher altitude,Pn showed a single-peak curve in Spring and Autumn. Tr showed a single-peak curve in Spring,Autumn and Winter,but double-peak curve in Summer and its value was in a sequence of Summer＞Autumn＞Spring＞Winter. With the increasing altitude,Tr decreased in Summer and Spring,but increased in Autumn and Winter.WUE increased in Summer and Spring,but did not significantly varied in Autumn and Winter.Seasonal drought could improve WUE.
7.Main environmental factors that influenced photosynthetic rate were PAR, Tair and RH.There was a highly significant negative correlation between Tair and Ci and Cond.RH represented a highly significant positive correlation with Ci and Cond. PAR had a highly significant positive correlation between Pn、Tr and Vpdl,and a highly significant negative correlation between Ci.Environmental factors influenced Moso bamboo photosynthesis due to Cond variation.High temperature in Summer and seasonal drought in Autumn and Winter had significant effects on Cond and caused midday depression of photosynthetic rate decline by means of stomatal limitation.Seasonal variation of LCP was in the sequence of Spring＞Summer＞Autumn＞Winter,and that of LSP was in the sequence of Autumn＞Summer＞Winter＞Spring.With the increasing altitude,LCP and LSP both decreased.Seasonal variation of P_(max) was in the sequence of Summer＞Autumn＞Winter＞Spring.With the increasing altitude,annual average P_(max) increased.P_(max) and annual average P_(max) increased in Summer and Autumn,but decreased in Winter and Spring.AQY in Summer and Autumn was higher than that both in Winter and Spring.With the increasing altitude,seasonal drought and leaf aging in Spring were the main reason affecting AQY.
1.随着海拔的升高,Tair明显下降,在3个海拔梯度不同季节,海拔升高100m的降温幅度无显著差异;RH较大辐度提高,以季节性干旱的秋冬季增湿作用强;PAR一定幅度增强;海拔升高100m,Tair下降0.61～0.69℃,RH提高1.42～2.47%,PAR增大23.61～60.36μmol·m~(-2)·s~(-1)。
2.同一海拔梯度毛竹林土壤容重、总孔隙度、非毛管孔隙度、毛管孔隙度、饱和持水量、田间持水量、粘粒和砂粒含量随土层加深而增大,粉粒含量减少。总孔隙度与容重呈负相关;海拔对土壤容重、毛管孔隙度、田间持水量无显著影响,总孔隙度、非毛管孔隙度、饱和持水量随海拔升高而显著增大;不同海拔梯度毛竹林土壤微结构颗粒组成砂粒＞粉粒＞粘粒,粘粒、粉粒含量随海拔升高而减少,砂粒含量趋于增加。
3.海拔与土壤pH值、全氮、速效氮呈极显著负相关,与有机质呈极显著正相关,对全磷、速效磷、全钾、速效钾影响不显著;土壤酶活性与海拔呈正相关,蔗糖酶、脲酶、磷酸酶在中海拔发生转折性变化,蛋白酶随海拔升高缓慢提高,过氧化氢酶呈均匀上升趋势。土壤物理性质对土壤酶活性无显著影响。有机质、全氮、速效氮与土壤酶活性呈显著或极显著正相关,pH值对土壤酶活性具正效应,对全磷、全钾、速磷、速钾影响不显著。不同种类土壤酶活性间呈显著或极显著相关,酶促反应具专一性和共同性特点。
4.立竹密度、立竹胸径、枝下高、全高随海拔升高显著增大,立竹整齐度、均匀度显著下降,种群个体生长存在“利己”行为;始出笋时间、出笋终止时间随海拔升高向后推移,出笋持续时间随海拔升高而缩短;退笋率随海拔升高而降低;竹笋——幼竹高生长划分为初期、上升期、盛期、末期4个时期,遵循“慢——快——慢”节律。高生长差异主要体现在随海拔升高而增大的盛期日平均高生长量上,相同出笋阶段的竹笋高生长期随海拔升高而延长。相同海拔各出笋时期竹笋高生长期前期最长,后期最短,高生长时间接近,具有“利他”性生态适应机制。
5.海拔与立竹胸高壁厚、壁厚率呈负相关。相同径级立竹重量、材积随海拔升高而降低,竹林竹材产量、材积随海拔升高而增长;立竹尖削度值随海拔升高显著增大;编制了不同海拔梯度毛竹林立竹材积与胸径的对应表;竹笋产量、品质随海拔升高而提高,海拔对竹笋品质影响主要反映在总糖、可溶性糖、P、Fe、含水率、可食比例随海拔升高而提高,纤维、热量、灰分、Ca随海拔升高而降低。
6.不同年龄立竹竹叶叶绿素含量随海拔升高有差异不显著的提高;Pn随海拔升高而增大,夏季＞秋季＞冬季＞春季,夏秋季中、低海拔呈双峰型,出现光合“午休”,高海拔毛竹林和冬季、春季呈单峰型;不同海拔毛竹林Tr夏季＞秋季＞春季＞冬季,春季、秋季、冬季呈单峰型,夏季呈双峰型,春季、夏季随海拔升高而降低,秋冬季变化趋势相反;WUE春季、夏季随海拔升高而提高,秋冬季变化不明显,季节性水分亏缺能提高WUE。
7.PAR、Tair、RH是影响毛竹光合的主要环境因子,Tair、RH分别与Ci、Cond极显著负相关和正相关,PAR与Pn、Tr、Vpdl极显著正相关,与Ci极显著负相关。环境因子通过Cond影响毛竹光合作用,对Cond产生显著影响的是夏季、初秋的高温和秋冬季的季节性水分亏缺,光合“午休”由气孔限制引起;不同海拔毛竹林LCP春季＞夏季＞秋季＞冬季,LSP秋季＞夏季＞冬季＞春季,均随海拔升高而降低。P_(max)夏季＞秋季＞冬季＞春季,夏季、秋季随海拔升高而增大,春季、冬季呈相反变化趋势,年平均P_(max)随海拔升高而增大。AQY秋季、夏季差异小,高于冬季、春季,随海拔升高而增大。季节性水分亏缺及春季竹叶老化是影响AQY的主要原因。
In the dissertation,the author has investigated changes of main weather factors, soil properties,population structure,economic characters,photosynthetic physiology of Moso bamboo forest at different altitudes.The growth and regeneration characteristics,effects of main environmental factors and adaptive mechanism were also revealed.The main results were described as follows:
1.With the increase of the altitude,Tair definitely decreased.When the altitude increased for 100m,there is no notable decrease range of temperature in three altitude grads among different seasons.RH has a bigger increment with the humidity increasing most obviously both in Autumn and Winter.PAR is also high in some scale.With the increase of altitude for 100m.Tair decreased at a range from 0.61℃to 0.69℃,RH increased at a range from 1.42%to 2.47%,and PAR increased at an extent from 23.61μmol·m~(-2)·s~(-1) to 60.36μmol·m~(-2)·s~(-1).The weather is more suitable for the growth of Moso bamboo forest when the altitude increased.
2.At the same altitude,with the increase of soil depth,soil bulk density,total porosity,non-capillary porosity,capillary porosity,saturated water capacity,field water-holding capacity,clay and sand particle content increased,but silt content decreased.There existed a negative correlation between total porosity and soil bulk density.Altitude had no significant effect on soil bulk density,capillary porosity, field water-holding capacity.With the increase of altitude,total porosity, non-capillary porosity,saturated water capacity increased significantly.Particle composition of soil in different altitudes varied in such a sequence:sand＞silt＞clay particle.With the increase of altitude,clay and silt particle content decreased,but sand content increased gradually.
3.There were a highly significant negative correlation among altitude and soil pH,total nitrogen and available nitrogen content,and a significant positive correlation between altitude and organic matter,but there was no significant effect on the contents of total phosphorus,available phosphorus,total potassium,and available potassium.There was a positive correlation between altitude and soil enzyme activities.At middle level of altitude,the great changes took place in phosphatase,urease and invertase.With the increase of altitude,protease activities slightly increased,and calatase activities also rise smoothly.Soil physical properties had no significant effect on soil enzyme activities.Soil enzyme activities had significant or definitely significant correlation with organic matter,total nitrogen and available nitrogen,and no significant correlation with total phosphorus, available phosphorus,total potassium,available potassium.There was a significant correlation among soil enzyme activities with specifiying and concomitant characteristics.
4.With the increase of altitude,the density,DBH,height under branch and total height increased significantly,but uniformity and evenness decreased significantly. Individual growth showed egoistic behavior to some extent.With increasing altitude, the time of beginning and termination of shooting were backward,whereas,the time of duration and degenerated shoot ratio decreased.The duration of earlier stage and later stage of shoot emergence was similar both at middle and lower altitude,but longer than that of at higher altitude.Height growth period from shoot to young culm could be divided into four stages,such as initial stage,rising stage,rapid growth stage and telophase,which showed a rhythm of slow growth followed by fast growth.With the increasing altitude,the difference of height growth existed mainly in the increase of mean height growth increment at rapid growth stage.The height growth period of shoot at the same stage was delayed and the duration of height growth shortened.Among all shooting stages,the earlier stage was longest,and the later stage was shortest at the same altitude.The high growth nearly ended in the same time which exhibited ecological adaptive mechanism of altruism.
5.There was a negative correlation between altitude and cell wall thickness at DBH,wall thickness ratio.With the increasing altitude,the weight and timber volume of Moso bamboo with the same DBH decreased,while the timber yield and volume increased.There was no significant difference in timber volume between middle and higher altitude due to multiple changes of structure factors of bamboo stands.With the increasing altitude,taper increased greatly.The corresponding table was established which could reflected the relationship between timber volume and DBH of bamboo in different altitude grads.With increase of altitude,the yield and quality of shoots enhanced.Eeffects of altitude on the quality of bamboo shoots depended on the rise of total sugar content,soluble sugar content,P,Fe,water content and edible proportion of shoot.But heat,fibre content,ash content and Ca all decreased with increasing altitude.
6.With the increasing altitude,the leaf chlorophyll content of Moso bamboo with different ages represented a small increasing trend,and Pn also increased in the sequence of Summer＞Autumn＞Winter＞Spring.At lower and middle altitude,Pn showed a double-peak curve,and midday depression of photosynthesis rate took place.But at higher altitude,Pn showed a single-peak curve in Spring and Autumn. Tr showed a single-peak curve in Spring,Autumn and Winter,but double-peak curve in Summer and its value was in a sequence of Summer＞Autumn＞Spring＞Winter. With the increasing altitude,Tr decreased in Summer and Spring,but increased in Autumn and Winter.WUE increased in Summer and Spring,but did not significantly varied in Autumn and Winter.Seasonal drought could improve WUE.
7.Main environmental factors that influenced photosynthetic rate were PAR, Tair and RH.There was a highly significant negative correlation between Tair and Ci and Cond.RH represented a highly significant positive correlation with Ci and Cond. PAR had a highly significant positive correlation between Pn、Tr and Vpdl,and a highly significant negative correlation between Ci.Environmental factors influenced Moso bamboo photosynthesis due to Cond variation.High temperature in Summer and seasonal drought in Autumn and Winter had significant effects on Cond and caused midday depression of photosynthetic rate decline by means of stomatal limitation.Seasonal variation of LCP was in the sequence of Spring＞Summer＞Autumn＞Winter,and that of LSP was in the sequence of Autumn＞Summer＞Winter＞Spring.With the increasing altitude,LCP and LSP both decreased.Seasonal variation of P_(max) was in the sequence of Summer＞Autumn＞Winter＞Spring.With the increasing altitude,annual average P_(max) increased.P_(max) and annual average P_(max) increased in Summer and Autumn,but decreased in Winter and Spring.AQY in Summer and Autumn was higher than that both in Winter and Spring.With the increasing altitude,seasonal drought and leaf aging in Spring were the main reason affecting AQY.
引文
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