亚热带北缘毛竹林群落生产力、有机碳及养分动态
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摘要
毛竹(Phyllostachys pubescens)是我国南方重要的森林资源,生长快,经营周期长,具有巨大的固碳能力。本文选取毛竹林为研究对象,通过样地调查与分析、野外定点观测、室内测定等研究亚热带北缘毛竹林群落生产力、有机碳和养分动态特征及其经营措施的影响,以揭示在全球变化背景下,亚热带北缘毛竹林生长、碳循环以及养分平衡关系。主要研究结果如下：
(1)毛竹林立竹密度为2711～3956株·hm-2；立竹度为0.33-0.64；平均眉径10.12cm,径级结构尚不合理,集中在9～15cm；平均高13.8-21.6m；毛竹林立竹整齐度都大于7,竹林很整齐；立竹均匀度为5.51,达到均匀竹林的水平；叶面积指数在5.01-13.14之间,样地1和样地3已达到丰产林水平,但年龄结构不合理,Ⅰ度和Ⅳ度以上毛竹比例偏大。
(2)毛竹单株生物量在6.99～56.89kg之间,地上部分占90%以上,随年龄增大先增加后减小(Ⅲ度以后),随径阶增大而增加。毛竹林分平均生物量为89.83t·hm-2,样地3的生物量最高,达到127.19t·hm-2；毛竹各器官生物量的大小排序为竹秆>竹枝>竹根>竹叶,各器官及单株生物量分别与眉径、株高间达到了极显著相关(p<0.01),并建立了相关方程。
(3)毛竹各器官含碳率排序为竹根(50.76%)>竹秆(49.44%)>竹枝(46.07%)>竹叶(41.71%),幼年期较稳定,老竹(3度以上)间差异显著(p<0.05)。未经营毛竹林碳贮量45.84tC·hm-2,而经营后的毛竹林碳贮量在63.66～101.11tC.hm-2。竹秆碳贮量占毛竹林碳贮量的78.22%,竹枝10.57%,竹根6.35%和竹叶4.87%；毛竹不同部位的营养元素含量存在着明显差异,叶、枝、秆和根中的N和K较多,P和Ca较少,但叶中Ca较多,而N、P、Ca、Mg元素含量叶>秆>枝>根；随年龄增加,不同器官中的钾含量降低,氮、磷与镁含量上下波动,钙含量相对稳定,且不同器官中的N、P、K、Ca、Mg元素间达到了显著(p<0.05)或极显著水平(p<0.01)。
(4)毛竹林年凋落物量为2.95～4.61t·hm-2·a-1,集中在4～5月份,呈单峰型,月份间凋落物量差异显著(p<0.05)；凋落物含碳率为32.93%～48.75%,凋落物碳年输入量以对照样地(1.85t·hm-2·a-1)最高,在4～5月份形成单峰值；凋落物中养分含量排序均为N>Ca>K>P>Mg,存在明显的季节变化。
(5)毛竹林土壤有机碳含量为21.61～42.30g·kg-1,上下层土壤DOC含量分别为0.15～0.17g·kg-1和0.11～0.15g·kg-1,均随土壤剖面深度增加而降低,变异系数分别为17.92%～26.10%和81.36%～150.26%,土层间有机碳含量均达到了显著差异水平(p<0.05),DOC均未达到显著差异(p>0.05)；土壤有机碳和DOC含量的月变化均呈波浪形,变化趋势大致相同。
毛竹林土壤呼吸日变化为较平缓的单峰曲线,峰值出现在14：00；存在明显的季节变化,月平均土壤呼吸变幅在0.5～5.12μmol CO2·m-2·s-1之间,不同月份间均差异显著(p<0.05)；土壤呼吸速率与温度(t)成正相关,与湿度(w,0～20cm土壤)成负相关,土壤呼吸强度(Y)与二者的指数关系式为：Y1=0.0615×e0.2374t(R2=0.8324)；Y0=9.8065×e-0.0449w(R2=0.8248)
(6)毛竹林土壤pH为4.62～6.89,上层(0～20cm)小于下层(20～40cm),变异系数27.81%～62.02%,仅样地3显著差异(p<0.05)；EC值为37～463μS·cm-1,上下层变异系数为4.04%～8.39%,差异均不显著(p>0.05)；上下层土壤有机质含量分别为54.64～72.92g·kg-1和37.26～50.80g·kg-1,上层均高于下层,变异系数为17.92%～26.10%,均达到了极显著差异水平(p<0.01)；上下层土壤全氮含量为0.74～4.64g·kg--和0.41～4.56g·kg-1,变异系数为22.17～46.22%,且在样地1、样地4中差异显著(p<0.05)；上下层土壤铵态氮含量为3.27-4.00g·kg-1和2.33～3.71g·kg-1,硝态氮含量2.29～6.81g·kg-1和2.00～4.35g·kg-1,上层均高于下层,但差异不显著(p>0.05)；上下层土壤全磷含量为0.05～2.68g·kg-1和0.02～2.39g·kg-1,上下层土壤速效磷含量为0.80～13.66mg·kg-1和(?)0.12～6.34mg·kg-1,上层土壤全磷和速效磷含量均高于下层,变异系数为49.53%～100.18%和40.80%～76.11%,上下层土壤全磷含量差异不显著(p>0.05)；土壤钾、钙、镁含量分别为0.52～8.58g·kg-1、0.07～4.15g·kg-1和0.63～3.69g·kg-1,变异系数分别为41.17%～53.43%、28.49%～55.58%和44.36%～50.47%,但均无显著差异(p>0.05)。
毛竹林上下层土壤pH、EC值随时间先升高后降低再升高；土壤有机质先降低后升高又降低；全氮含量在秋季(8、9、10月份)相对较低,其他时间相对较高；上层土壤铵态氮含量以春季后期最高,秋冬季最低,下层则以夏季最高,其他三季较低；上下层土壤硝态氮含量均是春季前期最高,秋季较高,其他月份较低；上下两层土壤全磷和速效磷含量均是秋季较高,冬季次之,春夏季较低；钾、钙和镁含量均是1、9、10月份含量较低,其他月份含量较高。
毛竹根际土(取样时间为10月份)的pH值、全氮含量小于非根际土；EC值、全磷、速效磷、钾、钙、镁以及有机质含量均高于非根际土,且毛竹根际土养分含量有随毛竹年龄先增加后减小的趋势。
(7)毛竹林原位土壤培养的净DOC转化速率在秋冬季多为负值,春夏季多为正值；两层土壤净DOC转化速率的峰值均出现在8～9月份,趋势相似,大小相近；净氨化速率在2009年的秋冬季和2010年的春夏季多为负值,2010年的秋冬季则多为正值,下层土壤要比上层土壤净氨化速率要大；净硝化速率与净氨化速率相反,峰值出现在5-8月份,上层土壤净硝化速率比下层大；上下层土壤净矿化速率的变化规律基本一致,多为正值,峰值出现在5-8月份,土壤矿化以硝化作用为主；各样地两层土壤的年平均氨化速率基本为负值,而年平均硝化速率和年平均矿化速率均为正值；土壤的净氨化速率与土壤的pH值呈极显著正相关(p<0.01),与温度、铵态氮含量呈极显著负相关(p<0.01)；净硝化速率与铵态氮含量呈极显著正相关(p<0.01)；净矿化速率与土壤物理因子呈显著相关性。
(8)毛竹秆茎流率为3.99%～23.44%,穿透率为22.65%～105.71%。不同年龄毛竹的秆茎流量不存在显著差异(p>0.05)；林外雨、秆茎流和穿透雨均呈酸性,秆茎流酸性最强,分别与林外雨和穿透雨呈极显著差异(p<0.01)；电导率以穿透雨最高,林外雨最小,且穿透雨的电导率分别与秆茎流、林外雨的电导率达到极显著差异(p<0.01)；林外雨中的养分元素浓度差异较大,Ca> K> N>Mg> P,钙年输入量为67.98kg·hm-2,磷年输入量仅为0.12kg·hm-2;秆茎流的营养元素浓度明显低于穿透雨,以K和Ca减少最明显；除秆茎流中Ca外,二者中的其他元素浓度均高于林外雨：秆茎流和穿透雨中营养元素浓度排序为K>Ca>NO3--N>NH4+-N>Mg>P;秆茎流养分年输入量很小,穿透雨养分年输入量除钙外均最高,而毛竹林降水的养分年净输入量以氮最高,13.03kg-hm-2,磷为0.06kg·hm-2,钙为负值,营养元素浓度存在明显的季节变化；不同年龄的毛竹秆茎流的年养分平均含量差异不显著(p>0.05)；不同降雨时间的秆茎流、穿透雨和大气降雨中养分含量存在显著(p<0.05)或极显著差异(p<0.01),特别是秋季以后降水中输入的养分减少；降水中的养分元素与降水量存在着明显的相关性。
(9)经营后,毛竹林的立竹密度、立竹度、平均眉径、平均高分别增加了32.9%～94.9%、34.5%～80.7%、20.4%～52.8%和23.1%～58.8%,经营措施明显提高了密度、立竹度、叶面积和上层土壤碳氮比,施肥和埋青增加了毛竹林生物量、碳贮量以及上层土壤硝态氮含量和5月份土壤全氮含量,且随抚育措施种类增多,叶面积指数和秆茎流量增大,碳贮量也达到了101.11tC·hm-2；经营措施降低了上层土壤有机质、全氮、铵态氮含量毛竹林年凋落物量、秆茎流量、毛竹林凋落物养分含量、年碳输入量和月碳输入量,且经营措施越多,上层土壤铵态氮含量越小,但对土壤pH值、EC值、全磷、速效磷、钾、钙、镁含量、土壤净DOC转化速率、净氨化速率、土壤净硝化速率、穿透雨量、秆茎流与穿透雨的养分含量等影响均不明显,因此,施肥、埋青等经营措施能够有效提高毛竹林群落生产力和碳贮量。
The Phyllostachys pubescens (Moso bamboo) that is a fast-growing plant is an important forest resource in southern China, which has a long management period and a huge fixing CO2ability from the atmosphere. The Phyllostachys pubescens forest was selected in this study. Based on plot investigation and analysis, fixed point measurement and in-lab determination, the productivity, organic carbon and nutrient dynamics of Phyllostachys pubescens communities after stand improvement and the effects of the different management measures were studied. The main results are as follows.
(1) The density of Moso bamboo forest is2711～3956individuals@hm-2; the bamboo stocking density is0.33～0.64; the mean diameter at the eyebrow height is10.12cm, the size structue is unreasonable, most of them are9～15cm; the average height is13.8-21.6m; the bamboo uniformity is more than7, which is uniform; the bamboo eveness is5.51, and the leaf area index (LAI) is5.01～13.14, but the age structure is unreasonable, the proportion of two-year and more than six-year individuals is bigger.
(2) The individual biomass of Moso bamboo is6.99～56.89kg, the aboveground part accounts for above90%, and increased and then decreased with the age increased (more than six years), and enhanced with the diameter grades increasing. The mean biomass of Moso bamboo forest is89.83t·hm-2, and the biomass of plot3is highest, which reaches127.19t·hm-2. The order of biomass for the different organs is ranked as culms> branches> roots> leaves, every organ and individual significantly related with the diameter at the eyebrow height and culm height (p<0.01), and the related equation was built.
(3) The order of carban concentrations in the different organs of Moso bamboo is roots (50.76%)> culms (49.44%)> branches (46.07%)> leaves (41.71%). It is more stable in young stage for carban concentrations, but there is notable difference among older bamboos (more than6-year)(p<0.05). The carbon storage of Moso bamboo forest is63.66～101.11t C·hm-2for the intensive management, while is only45.84t C·hm-2for the extensive management. The carbon storage of culms, branches, roots and leaves account for78.22%,10.57%,6.35%and4.87%of the total carbon storage of the stand respectively. The difference of nutrient contents is obvious in different organs. The contents of N and K in leaves, branchs, culms and roots are higher, and the contents of P and Ca are lower, but the content of Ca in the leaves is higher. The order of N, P, Ca, Mg contents is leaves> culms> branches> roots. The content of potassium in different organs decreased, with age, the content of nitrogen, phosphorus and magnesium fluctuated, and the content of calcium is relatively stable. The content of N, P, K, Ca, Mg among different organs reached significant (p<0.05) or marked (p<0.01) correlation.
(4) The annual amounts of litterfall were2.95～4.61t·ha-1·a-1, which mostly fell in April and May with a single-peak. The amounts of litterfall in the different months differed significantly (p<0.05). The carban concentrations of litterfall are32.93%～48.75%, the annual carbon input from litterfall in CK is highest (1.85t·hm-2·a-1), which peak occurred in April or May. The order of nutrient contents in litterfall is N> Ca> K> P> Mg. The nutrient content of litterfall in unimproved plot is largest, with obviously seasonal variation.
(5)The soil organic carbon contents are21.61～42.30g·kg-1, and the dissolved organic carban (DOC) contents of upper (0～20cm) and lower (20-40cm) soils were, respectively,0.15～0.17g·kg-1and0.11～0.15g·kg-1, reduced with the increasing soil depth, the coefficient of variation is17.92%～26.10%and81.36%～150.26%. The organic carbon content between upper and lower layers have extremely difference (p <0.05), while DOC content has not significant difference (p>0.05). In the same soil layer, soil organic carbon content among different plots have reached notable or extremely notable difference, but DOC content has not significant difference (p>0.05). The change of soil organic carbon and DOC content fluctuated with month. The change trend is similar.
The diurnal change of soil respiration in Moso bamboo forest showed a single-peak which appeared in14:00. There is an obvious seasonal dynamics. The average monthly rates of soil respiration varied from0.5to5.12μmol CO2·m-2·s-1, with a significant difference among months (p<0.05). The correlation between soil respiration rate and temperature (t) is positive, and it is negative with humidity (w,0～20cm soil depth), the correlated equation among soil respiration intensity (Y), soil temperature and humidity is: Y1=0.0615×e0.2374t(R2=0.8324); Y2=9.8065×e-0.0449w (R2=0.8248)
(6) In Moso bamboo forest, soil pH is4.62～6.89, the upper layer (0～20cm) is smaller than the lower layer (20～40cm), the coefficient of variation is27.81%～62.02%, there is a significant difference (p<0.05) only in plot3. EC values are37～463μS·cm-1with the coefficient of variation of4.04%～8.39%, there is not significant difference (p>0.05). The soil organic matter contents in two layers are54.64～72.92g·kg-1and37.26～50.80g·kg-1, the upper layer is larger than the lower layer, the coefficient of variation is17.92%～26.10%, and the difference have reached extremely notable level (p<0.01); the total nitrogen content in upper and lower soils is0.74～4.64g·kg-1and0.41～4.56g·kg-1, the coefficient of variation is22.17～46.22%, which has a significant difference between the upper and lower layers in plot1and4(p<0.05). The soil ammonium contents in upper and lower layers are3.27～4.00g·kg-1and2.33～3.71g·kg-1. The nitrate contents are2.29～6.81g·kg-1and2.00～4.35g·kg-1, the upper layer is higher than the lower layer, but it has not significant difference (p>0.05). The soil total phosphorus contents in upper and lower layer are0.05～2.68g·kg-1and0.02～2.39g·kg-1, and the soil available phosphorus contents are0.80～13.66m g·kg-1and0.12～6.34m g·kg-1, the coefficient of variation is49.53%～100.18%and40.80%～76.11%, the upper layer is higher than the lower layer, but it has not significant difference of total phosphorus content (p>0.05). The soil potassium, calcium, magnesium contents are, respectively,0.52～8.58g·kg-1,0.074.15g·kg-1and0.63～3.69g·kg-1, and the coefficient of variation are41.17%～55.58%53.43%,28.49%～and44.36%～50.47%, but there are not significant difference (p>0.05).
The soil pH and EC value first rised, then decreased and rised again, while soil organic matter content is contrary. The total nitrogen contents are relatively lower in autumn (August, September and October), higher in others. The ammonium nitrogen content in the upper soils is highest in latter spring and lowest in autumn and winter; while for the lower soils it is highest in summer and lower in other season. The soil nitrate nitrogen content in two layers is highest in early spring, higher in autumn and lowest in summer and winter. The soil total and available phosphorus contents in two layers are highest in autumn, higher in winter and lowest in spring and summer. The soil contents of potassium, calcium and magnesium are lower in January, September and October, higher in other months.
The rhizosphere soil (sampling time in October) pH and total nitrogen is smaller than non-rhizosphere soil; and the EC, total phosphorus, available phosphorus, potassium, calcium, magnesium in rhizosphere soil is higher than non-rhizosphere soil, the nutrient in rhizosphere soil first increased then decreased with the age.
(7) The net rate of soil DOC in situ incubation in Moso bamboo forest is negative mostly in autumn and winter, while is positive in spring and summer. The peak value of the net DOC translation rate in two layer appears in August and September, the trend and the size is similar; The net rate of soil ammonification is negative in autumn and winter in2009and spring and summer in2010while it is positive in autumn and winter in2010, which is higher in the upper layer than in the lower layer. The net rate of soil nitrifieation is contrary to the ammonification, its peak value appears from May to August, with a greater value in the upper layer than in the lower layer.
It is not obvious among the net rate of soil DOC, ammonification, nitrifieation under different mamagement measures, but it is positive to ammonification and negative to nitrifieation. The soil net mineralization rates between the upper and the lower layers are similar, and are positive several times, the peak value appears from May to August, it is mainly from the nitrification in soil mineralization. The annual average soil ammonification rate is negative in all plots, but the annual average soil nitrifieation and mineralization rate is positive. The net ammonification rate in soil is extremely positive related with the soil pH (p<0.01), and negative extremely related with the soil temperature and the ammonium nitrogen content (p<0.01); the net nitrification rate is extremely positive related with ammonium nitrogen contents (p<0.01); the net mineralization rate is related with soil physical characteristics.
(8) The rate of stemflow and throughfall in Moso bamboo forest is3.99%-23.44%and22.65%～105.71%, respectively. The stemflow from the different aged bamboo has not signifecant difference (p>0.05). The pH values of stemflow, throughfall and rainfall are all acidic. The acidity of stemflow is strongest, which has extremely notable difference with rainfall and throughfall (p<0.01). The electrical conductivity (EC) in throughfall is highest, the rainfall is lowest, and it has extremely significant difference with the EC in stemflow and rainfall (p<0.01. It has a high variance in nutrients in rainfall, the order of nutrient content is Ca> K> N> Mg> P; the annual calcium input is67.98kg·hm-2, and the annual phosphorus input is only0.12kg·hm-2. The nutrient elements in stemflow are lower than throughfall obviously, especialy the decrease of K and Ca is the most obvious. The order of nutrient input in stemflow and throughfall is K> Ca> NO3—N>NH4+-N> Mg> P, which is higher than rainfall except calcium. The annual nutrient input by stemflow of Moso bamboo forest is very small, while it is large by throughfall except for calcium. The annual nitrogen input by precipitation in Moso bamboo forest is highest with annual mean of13.03kg·hm-2. However, the phosphorus input is0.06kg·hm-2, and the calcium input is negative. There exists obviously seasonal variation in nutrient fluxes. The annual average nutrient contents of stemflow from different aged bamboo individuals has not significant difference (p>0.05), but the nutrinet content among stemflow, throughfall and precipitation in different raining time exist notable difference (p<0.05) or extremely notable difference (p<0.01), especialy the nutrient input from precipitation reduced after autumn. The nutrient element content in precipitation is related with the amount of precipitation.
(9) The density, stocking density, average diameter at the eyebrow height and height of Moso bamboo after taking measurements has increased by32.9%～94.9%,34.5%～80.7%,20.4%～52.8%and23.1%～58.8%respectively, the measurements have obviously increased the density, stocking density, leaf area and upper soil carbon-nitrogen ratio. The fertilization and burying weeds and shrubs increased the biomass, carbon storage, the upper soil nitrate-N contents and the total nitrogen content in May. With the multiple measurements, the LAI and the stemflow amounts enhanced, and the carbon storage reached101.11t C· hm-2. While the measurements has reduced the upper soil organic content, the soil total nitrogen and ammonium nitrogen contents, annual littterfall, stemflow amounts, the nutrient in litterfall, annual and monthl carbon input amounts. The upper soil ammonium nitrogen contents reduced with the measure kinds increased. It is not obvious that the management measures affect the soil pH, EC value, total phosphorus, available phosphorus, potassium, calcium, magnesium, net rate of soil DOC in situ incubation, net rate of soil ammonification and nitrifieation, throughfall, nutrient in stemflow and throughfall. Therefore, fertilization and burying weeds and shrubs can raise the productivity and carbon storage of Moso bamboo forest.
(1)毛竹林立竹密度为2711～3956株·hm-2；立竹度为0.33-0.64；平均眉径10.12cm,径级结构尚不合理,集中在9～15cm；平均高13.8-21.6m；毛竹林立竹整齐度都大于7,竹林很整齐；立竹均匀度为5.51,达到均匀竹林的水平；叶面积指数在5.01-13.14之间,样地1和样地3已达到丰产林水平,但年龄结构不合理,Ⅰ度和Ⅳ度以上毛竹比例偏大。
(2)毛竹单株生物量在6.99～56.89kg之间,地上部分占90%以上,随年龄增大先增加后减小(Ⅲ度以后),随径阶增大而增加。毛竹林分平均生物量为89.83t·hm-2,样地3的生物量最高,达到127.19t·hm-2；毛竹各器官生物量的大小排序为竹秆>竹枝>竹根>竹叶,各器官及单株生物量分别与眉径、株高间达到了极显著相关(p<0.01),并建立了相关方程。
(3)毛竹各器官含碳率排序为竹根(50.76%)>竹秆(49.44%)>竹枝(46.07%)>竹叶(41.71%),幼年期较稳定,老竹(3度以上)间差异显著(p<0.05)。未经营毛竹林碳贮量45.84tC·hm-2,而经营后的毛竹林碳贮量在63.66～101.11tC.hm-2。竹秆碳贮量占毛竹林碳贮量的78.22%,竹枝10.57%,竹根6.35%和竹叶4.87%；毛竹不同部位的营养元素含量存在着明显差异,叶、枝、秆和根中的N和K较多,P和Ca较少,但叶中Ca较多,而N、P、Ca、Mg元素含量叶>秆>枝>根；随年龄增加,不同器官中的钾含量降低,氮、磷与镁含量上下波动,钙含量相对稳定,且不同器官中的N、P、K、Ca、Mg元素间达到了显著(p<0.05)或极显著水平(p<0.01)。
(4)毛竹林年凋落物量为2.95～4.61t·hm-2·a-1,集中在4～5月份,呈单峰型,月份间凋落物量差异显著(p<0.05)；凋落物含碳率为32.93%～48.75%,凋落物碳年输入量以对照样地(1.85t·hm-2·a-1)最高,在4～5月份形成单峰值；凋落物中养分含量排序均为N>Ca>K>P>Mg,存在明显的季节变化。
(5)毛竹林土壤有机碳含量为21.61～42.30g·kg-1,上下层土壤DOC含量分别为0.15～0.17g·kg-1和0.11～0.15g·kg-1,均随土壤剖面深度增加而降低,变异系数分别为17.92%～26.10%和81.36%～150.26%,土层间有机碳含量均达到了显著差异水平(p<0.05),DOC均未达到显著差异(p>0.05)；土壤有机碳和DOC含量的月变化均呈波浪形,变化趋势大致相同。
毛竹林土壤呼吸日变化为较平缓的单峰曲线,峰值出现在14：00；存在明显的季节变化,月平均土壤呼吸变幅在0.5～5.12μmol CO2·m-2·s-1之间,不同月份间均差异显著(p<0.05)；土壤呼吸速率与温度(t)成正相关,与湿度(w,0～20cm土壤)成负相关,土壤呼吸强度(Y)与二者的指数关系式为：Y1=0.0615×e0.2374t(R2=0.8324)；Y0=9.8065×e-0.0449w(R2=0.8248)
(6)毛竹林土壤pH为4.62～6.89,上层(0～20cm)小于下层(20～40cm),变异系数27.81%～62.02%,仅样地3显著差异(p<0.05)；EC值为37～463μS·cm-1,上下层变异系数为4.04%～8.39%,差异均不显著(p>0.05)；上下层土壤有机质含量分别为54.64～72.92g·kg-1和37.26～50.80g·kg-1,上层均高于下层,变异系数为17.92%～26.10%,均达到了极显著差异水平(p<0.01)；上下层土壤全氮含量为0.74～4.64g·kg--和0.41～4.56g·kg-1,变异系数为22.17～46.22%,且在样地1、样地4中差异显著(p<0.05)；上下层土壤铵态氮含量为3.27-4.00g·kg-1和2.33～3.71g·kg-1,硝态氮含量2.29～6.81g·kg-1和2.00～4.35g·kg-1,上层均高于下层,但差异不显著(p>0.05)；上下层土壤全磷含量为0.05～2.68g·kg-1和0.02～2.39g·kg-1,上下层土壤速效磷含量为0.80～13.66mg·kg-1和(?)0.12～6.34mg·kg-1,上层土壤全磷和速效磷含量均高于下层,变异系数为49.53%～100.18%和40.80%～76.11%,上下层土壤全磷含量差异不显著(p>0.05)；土壤钾、钙、镁含量分别为0.52～8.58g·kg-1、0.07～4.15g·kg-1和0.63～3.69g·kg-1,变异系数分别为41.17%～53.43%、28.49%～55.58%和44.36%～50.47%,但均无显著差异(p>0.05)。
毛竹林上下层土壤pH、EC值随时间先升高后降低再升高；土壤有机质先降低后升高又降低；全氮含量在秋季(8、9、10月份)相对较低,其他时间相对较高；上层土壤铵态氮含量以春季后期最高,秋冬季最低,下层则以夏季最高,其他三季较低；上下层土壤硝态氮含量均是春季前期最高,秋季较高,其他月份较低；上下两层土壤全磷和速效磷含量均是秋季较高,冬季次之,春夏季较低；钾、钙和镁含量均是1、9、10月份含量较低,其他月份含量较高。
毛竹根际土(取样时间为10月份)的pH值、全氮含量小于非根际土；EC值、全磷、速效磷、钾、钙、镁以及有机质含量均高于非根际土,且毛竹根际土养分含量有随毛竹年龄先增加后减小的趋势。
(7)毛竹林原位土壤培养的净DOC转化速率在秋冬季多为负值,春夏季多为正值；两层土壤净DOC转化速率的峰值均出现在8～9月份,趋势相似,大小相近；净氨化速率在2009年的秋冬季和2010年的春夏季多为负值,2010年的秋冬季则多为正值,下层土壤要比上层土壤净氨化速率要大；净硝化速率与净氨化速率相反,峰值出现在5-8月份,上层土壤净硝化速率比下层大；上下层土壤净矿化速率的变化规律基本一致,多为正值,峰值出现在5-8月份,土壤矿化以硝化作用为主；各样地两层土壤的年平均氨化速率基本为负值,而年平均硝化速率和年平均矿化速率均为正值；土壤的净氨化速率与土壤的pH值呈极显著正相关(p<0.01),与温度、铵态氮含量呈极显著负相关(p<0.01)；净硝化速率与铵态氮含量呈极显著正相关(p<0.01)；净矿化速率与土壤物理因子呈显著相关性。
(8)毛竹秆茎流率为3.99%～23.44%,穿透率为22.65%～105.71%。不同年龄毛竹的秆茎流量不存在显著差异(p>0.05)；林外雨、秆茎流和穿透雨均呈酸性,秆茎流酸性最强,分别与林外雨和穿透雨呈极显著差异(p<0.01)；电导率以穿透雨最高,林外雨最小,且穿透雨的电导率分别与秆茎流、林外雨的电导率达到极显著差异(p<0.01)；林外雨中的养分元素浓度差异较大,Ca> K> N>Mg> P,钙年输入量为67.98kg·hm-2,磷年输入量仅为0.12kg·hm-2;秆茎流的营养元素浓度明显低于穿透雨,以K和Ca减少最明显；除秆茎流中Ca外,二者中的其他元素浓度均高于林外雨：秆茎流和穿透雨中营养元素浓度排序为K>Ca>NO3--N>NH4+-N>Mg>P;秆茎流养分年输入量很小,穿透雨养分年输入量除钙外均最高,而毛竹林降水的养分年净输入量以氮最高,13.03kg-hm-2,磷为0.06kg·hm-2,钙为负值,营养元素浓度存在明显的季节变化；不同年龄的毛竹秆茎流的年养分平均含量差异不显著(p>0.05)；不同降雨时间的秆茎流、穿透雨和大气降雨中养分含量存在显著(p<0.05)或极显著差异(p<0.01),特别是秋季以后降水中输入的养分减少；降水中的养分元素与降水量存在着明显的相关性。
(9)经营后,毛竹林的立竹密度、立竹度、平均眉径、平均高分别增加了32.9%～94.9%、34.5%～80.7%、20.4%～52.8%和23.1%～58.8%,经营措施明显提高了密度、立竹度、叶面积和上层土壤碳氮比,施肥和埋青增加了毛竹林生物量、碳贮量以及上层土壤硝态氮含量和5月份土壤全氮含量,且随抚育措施种类增多,叶面积指数和秆茎流量增大,碳贮量也达到了101.11tC·hm-2；经营措施降低了上层土壤有机质、全氮、铵态氮含量毛竹林年凋落物量、秆茎流量、毛竹林凋落物养分含量、年碳输入量和月碳输入量,且经营措施越多,上层土壤铵态氮含量越小,但对土壤pH值、EC值、全磷、速效磷、钾、钙、镁含量、土壤净DOC转化速率、净氨化速率、土壤净硝化速率、穿透雨量、秆茎流与穿透雨的养分含量等影响均不明显,因此,施肥、埋青等经营措施能够有效提高毛竹林群落生产力和碳贮量。
The Phyllostachys pubescens (Moso bamboo) that is a fast-growing plant is an important forest resource in southern China, which has a long management period and a huge fixing CO2ability from the atmosphere. The Phyllostachys pubescens forest was selected in this study. Based on plot investigation and analysis, fixed point measurement and in-lab determination, the productivity, organic carbon and nutrient dynamics of Phyllostachys pubescens communities after stand improvement and the effects of the different management measures were studied. The main results are as follows.
(1) The density of Moso bamboo forest is2711～3956individuals@hm-2; the bamboo stocking density is0.33～0.64; the mean diameter at the eyebrow height is10.12cm, the size structue is unreasonable, most of them are9～15cm; the average height is13.8-21.6m; the bamboo uniformity is more than7, which is uniform; the bamboo eveness is5.51, and the leaf area index (LAI) is5.01～13.14, but the age structure is unreasonable, the proportion of two-year and more than six-year individuals is bigger.
(2) The individual biomass of Moso bamboo is6.99～56.89kg, the aboveground part accounts for above90%, and increased and then decreased with the age increased (more than six years), and enhanced with the diameter grades increasing. The mean biomass of Moso bamboo forest is89.83t·hm-2, and the biomass of plot3is highest, which reaches127.19t·hm-2. The order of biomass for the different organs is ranked as culms> branches> roots> leaves, every organ and individual significantly related with the diameter at the eyebrow height and culm height (p<0.01), and the related equation was built.
(3) The order of carban concentrations in the different organs of Moso bamboo is roots (50.76%)> culms (49.44%)> branches (46.07%)> leaves (41.71%). It is more stable in young stage for carban concentrations, but there is notable difference among older bamboos (more than6-year)(p<0.05). The carbon storage of Moso bamboo forest is63.66～101.11t C·hm-2for the intensive management, while is only45.84t C·hm-2for the extensive management. The carbon storage of culms, branches, roots and leaves account for78.22%,10.57%,6.35%and4.87%of the total carbon storage of the stand respectively. The difference of nutrient contents is obvious in different organs. The contents of N and K in leaves, branchs, culms and roots are higher, and the contents of P and Ca are lower, but the content of Ca in the leaves is higher. The order of N, P, Ca, Mg contents is leaves> culms> branches> roots. The content of potassium in different organs decreased, with age, the content of nitrogen, phosphorus and magnesium fluctuated, and the content of calcium is relatively stable. The content of N, P, K, Ca, Mg among different organs reached significant (p<0.05) or marked (p<0.01) correlation.
(4) The annual amounts of litterfall were2.95～4.61t·ha-1·a-1, which mostly fell in April and May with a single-peak. The amounts of litterfall in the different months differed significantly (p<0.05). The carban concentrations of litterfall are32.93%～48.75%, the annual carbon input from litterfall in CK is highest (1.85t·hm-2·a-1), which peak occurred in April or May. The order of nutrient contents in litterfall is N> Ca> K> P> Mg. The nutrient content of litterfall in unimproved plot is largest, with obviously seasonal variation.
(5)The soil organic carbon contents are21.61～42.30g·kg-1, and the dissolved organic carban (DOC) contents of upper (0～20cm) and lower (20-40cm) soils were, respectively,0.15～0.17g·kg-1and0.11～0.15g·kg-1, reduced with the increasing soil depth, the coefficient of variation is17.92%～26.10%and81.36%～150.26%. The organic carbon content between upper and lower layers have extremely difference (p <0.05), while DOC content has not significant difference (p>0.05). In the same soil layer, soil organic carbon content among different plots have reached notable or extremely notable difference, but DOC content has not significant difference (p>0.05). The change of soil organic carbon and DOC content fluctuated with month. The change trend is similar.
The diurnal change of soil respiration in Moso bamboo forest showed a single-peak which appeared in14:00. There is an obvious seasonal dynamics. The average monthly rates of soil respiration varied from0.5to5.12μmol CO2·m-2·s-1, with a significant difference among months (p<0.05). The correlation between soil respiration rate and temperature (t) is positive, and it is negative with humidity (w,0～20cm soil depth), the correlated equation among soil respiration intensity (Y), soil temperature and humidity is: Y1=0.0615×e0.2374t(R2=0.8324); Y2=9.8065×e-0.0449w (R2=0.8248)
(6) In Moso bamboo forest, soil pH is4.62～6.89, the upper layer (0～20cm) is smaller than the lower layer (20～40cm), the coefficient of variation is27.81%～62.02%, there is a significant difference (p<0.05) only in plot3. EC values are37～463μS·cm-1with the coefficient of variation of4.04%～8.39%, there is not significant difference (p>0.05). The soil organic matter contents in two layers are54.64～72.92g·kg-1and37.26～50.80g·kg-1, the upper layer is larger than the lower layer, the coefficient of variation is17.92%～26.10%, and the difference have reached extremely notable level (p<0.01); the total nitrogen content in upper and lower soils is0.74～4.64g·kg-1and0.41～4.56g·kg-1, the coefficient of variation is22.17～46.22%, which has a significant difference between the upper and lower layers in plot1and4(p<0.05). The soil ammonium contents in upper and lower layers are3.27～4.00g·kg-1and2.33～3.71g·kg-1. The nitrate contents are2.29～6.81g·kg-1and2.00～4.35g·kg-1, the upper layer is higher than the lower layer, but it has not significant difference (p>0.05). The soil total phosphorus contents in upper and lower layer are0.05～2.68g·kg-1and0.02～2.39g·kg-1, and the soil available phosphorus contents are0.80～13.66m g·kg-1and0.12～6.34m g·kg-1, the coefficient of variation is49.53%～100.18%and40.80%～76.11%, the upper layer is higher than the lower layer, but it has not significant difference of total phosphorus content (p>0.05). The soil potassium, calcium, magnesium contents are, respectively,0.52～8.58g·kg-1,0.074.15g·kg-1and0.63～3.69g·kg-1, and the coefficient of variation are41.17%～55.58%53.43%,28.49%～and44.36%～50.47%, but there are not significant difference (p>0.05).
The soil pH and EC value first rised, then decreased and rised again, while soil organic matter content is contrary. The total nitrogen contents are relatively lower in autumn (August, September and October), higher in others. The ammonium nitrogen content in the upper soils is highest in latter spring and lowest in autumn and winter; while for the lower soils it is highest in summer and lower in other season. The soil nitrate nitrogen content in two layers is highest in early spring, higher in autumn and lowest in summer and winter. The soil total and available phosphorus contents in two layers are highest in autumn, higher in winter and lowest in spring and summer. The soil contents of potassium, calcium and magnesium are lower in January, September and October, higher in other months.
The rhizosphere soil (sampling time in October) pH and total nitrogen is smaller than non-rhizosphere soil; and the EC, total phosphorus, available phosphorus, potassium, calcium, magnesium in rhizosphere soil is higher than non-rhizosphere soil, the nutrient in rhizosphere soil first increased then decreased with the age.
(7) The net rate of soil DOC in situ incubation in Moso bamboo forest is negative mostly in autumn and winter, while is positive in spring and summer. The peak value of the net DOC translation rate in two layer appears in August and September, the trend and the size is similar; The net rate of soil ammonification is negative in autumn and winter in2009and spring and summer in2010while it is positive in autumn and winter in2010, which is higher in the upper layer than in the lower layer. The net rate of soil nitrifieation is contrary to the ammonification, its peak value appears from May to August, with a greater value in the upper layer than in the lower layer.
It is not obvious among the net rate of soil DOC, ammonification, nitrifieation under different mamagement measures, but it is positive to ammonification and negative to nitrifieation. The soil net mineralization rates between the upper and the lower layers are similar, and are positive several times, the peak value appears from May to August, it is mainly from the nitrification in soil mineralization. The annual average soil ammonification rate is negative in all plots, but the annual average soil nitrifieation and mineralization rate is positive. The net ammonification rate in soil is extremely positive related with the soil pH (p<0.01), and negative extremely related with the soil temperature and the ammonium nitrogen content (p<0.01); the net nitrification rate is extremely positive related with ammonium nitrogen contents (p<0.01); the net mineralization rate is related with soil physical characteristics.
(8) The rate of stemflow and throughfall in Moso bamboo forest is3.99%-23.44%and22.65%～105.71%, respectively. The stemflow from the different aged bamboo has not signifecant difference (p>0.05). The pH values of stemflow, throughfall and rainfall are all acidic. The acidity of stemflow is strongest, which has extremely notable difference with rainfall and throughfall (p<0.01). The electrical conductivity (EC) in throughfall is highest, the rainfall is lowest, and it has extremely significant difference with the EC in stemflow and rainfall (p<0.01. It has a high variance in nutrients in rainfall, the order of nutrient content is Ca> K> N> Mg> P; the annual calcium input is67.98kg·hm-2, and the annual phosphorus input is only0.12kg·hm-2. The nutrient elements in stemflow are lower than throughfall obviously, especialy the decrease of K and Ca is the most obvious. The order of nutrient input in stemflow and throughfall is K> Ca> NO3—N>NH4+-N> Mg> P, which is higher than rainfall except calcium. The annual nutrient input by stemflow of Moso bamboo forest is very small, while it is large by throughfall except for calcium. The annual nitrogen input by precipitation in Moso bamboo forest is highest with annual mean of13.03kg·hm-2. However, the phosphorus input is0.06kg·hm-2, and the calcium input is negative. There exists obviously seasonal variation in nutrient fluxes. The annual average nutrient contents of stemflow from different aged bamboo individuals has not significant difference (p>0.05), but the nutrinet content among stemflow, throughfall and precipitation in different raining time exist notable difference (p<0.05) or extremely notable difference (p<0.01), especialy the nutrient input from precipitation reduced after autumn. The nutrient element content in precipitation is related with the amount of precipitation.
(9) The density, stocking density, average diameter at the eyebrow height and height of Moso bamboo after taking measurements has increased by32.9%～94.9%,34.5%～80.7%,20.4%～52.8%and23.1%～58.8%respectively, the measurements have obviously increased the density, stocking density, leaf area and upper soil carbon-nitrogen ratio. The fertilization and burying weeds and shrubs increased the biomass, carbon storage, the upper soil nitrate-N contents and the total nitrogen content in May. With the multiple measurements, the LAI and the stemflow amounts enhanced, and the carbon storage reached101.11t C· hm-2. While the measurements has reduced the upper soil organic content, the soil total nitrogen and ammonium nitrogen contents, annual littterfall, stemflow amounts, the nutrient in litterfall, annual and monthl carbon input amounts. The upper soil ammonium nitrogen contents reduced with the measure kinds increased. It is not obvious that the management measures affect the soil pH, EC value, total phosphorus, available phosphorus, potassium, calcium, magnesium, net rate of soil DOC in situ incubation, net rate of soil ammonification and nitrifieation, throughfall, nutrient in stemflow and throughfall. Therefore, fertilization and burying weeds and shrubs can raise the productivity and carbon storage of Moso bamboo forest.
引文
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