摘要
首次对灵石山不同海拔米槠林优势种叶片δ~(13)C值以及叶片养分含量、SLA、LDMC等各叶属性特征不同优势种间、不同叶龄间、不同海拔梯度间的差异性及其沿海拔梯度的变化规律进行分析;对δ~(13)C值与各叶属性因子的关系、各环境因子对叶片δ~(13)C值的影响以及各叶属性因子之间的关系及其与环境的关系等方面进行研究。
1.灵石山米槠林各海拔优势种叶δ~(13)C值介于-34.52‰～-28.377‰之间,平均值为-30.885‰。叶δ~(13)C值种间以及叶龄间的差异性分析表明,除A7、A8海拔梯度上优势种间叶δ~(13)C值差异性不显著外,其它各海拔梯度种间叶δ~(13)C有显著差异;各海拔梯度上不同叶龄间δ~(13)C值差异性不显著。各海拔梯度上叶δ~(13)C值分布特征表现为,随海拔升高,叶δ~(13)C值分组逐渐单一。优势种叶δ~(13)C值海拔间的差异性显著,灵石山米槠林优势种叶δ~(13)C值随海拔升高而逐渐升高,平均变化量为3.2‰·km-1。
2.分别对10种优势种不同叶龄叶绿体色素的差异分析表明,除少数优势种在某海拔梯度以外,叶绿素a、叶绿素b和叶绿素(a+b)存在显著或极显著叶龄间的差异,一般在中高海拔梯度差异最大,且2年生叶片含量高于1年生叶片。而不同年龄叶片间叶绿素a/b和类胡萝卜素含量的差异性表现形式复杂。不同优势种的叶绿素(叶绿素a、叶绿素b和叶绿素(a+b))在海拔梯度间存在显著的差异,并且沿海拔梯度的变化呈现一定的规律性,有7种优势种不同叶龄叶绿素含量均随海拔的升高先升高,一般在A5或A6达到最大值,然后随海拔的升高逐渐降低,而叶绿素a/b随海拔升高变化趋势比较多样化。类胡罗卜素作为功能色素由于其功能比较复杂,其含量对海拔的响应规律也不尽相同。说明海拔作为重要的环境因素对于植物叶绿体色素产生了显著的影响。
3.优势种养分含量的分布特征分析表明,P和Na含量的分布范围较窄,分别为0.259～2.026 g·kg-1和0.150～1.804 g·kg-1;其它养分中,Ca含量变化范围最广,且除Mn以外,养分浓度在其分布范围内的低中部频数较高,说明优势种养分含量较低的居多。1年生叶片N、P、K含量高于2年生叶片,2年生叶片Na、Ca含量高1年生叶片,微量元素Fe、Mn、Zn的平均含量也为2年生叶片高于1年生叶片。不同叶龄P含量、2年生叶片Na含量、2年生叶片Ca含量、1年生叶片Mg含量、1年生叶片Fe含量、不同叶龄Zn含量符合对数正态分布。四级分类法对优势种养分含量进行分类,不同优势种对不同养分的“排斥”和“累积”效应不同。围涎树和石栎对N、狗骨柴和鹅掌柴对K和Na有明显的“累积”效应,厚叶冬青对P、青冈栎对Zn则表现出明显的“排斥”效应。除2年生叶片Fe含量优势种间差异性不显著外,其它养分含量种间差异性显著(p<0.05),多重比较表明,养分含量处于不同分类等级中的优势种差异显著,部分处于相同等级中的优势种也有显著的差异性。
4.对不同叶龄SLA、LDMC以及养分特征等叶片属性沿海拔梯度的变化规律进行分析,各叶龄叶片SLA、LDMC、C、K、Mg含量以及2年生叶N、Mn含量均沿海拔梯度的变化表现出明显的规律性(P<0.05),其中LDMC随海拔梯度升高逐渐升高,其它指标则随海拔梯度的升高而降低。这种表现规律显示了植物叶片对生境的适应策略,随海拔升高养分含量降低,同时SLA降低、LDMC升高是对养分相对不足适应性的表现。
5.对各海拔以及综合所有海拔梯度上优势种叶δ~(13)C值进行回归分析,除叶片有机碳含量在各海拔均与δ~(13)C值不存在相关关系外,其它叶属性指标均与叶片δ~(13)C值有显著的线性回归关系。综合所有海拔梯度,不同叶龄δ~(13)C值与P、Ca、LDMC含量显著正相关;不同年龄叶δ~(13)C值与SLA以及N、K、Na、Mg、Ash含量、2年生叶片叶绿素含量显著负相关;不同叶龄叶片δ~(13)C与Fe、Mn、Zn的关系在各海拔表现形式复杂,综合所有海拔,δ~(13)C与Fe、Zn相关性不显著,与1年生叶片Mn含量表现为负相关,与2年生叶片Mn含量表现为正相关。不同海拔梯度上以及不同叶龄δ~(13)C值与各叶属性指标的回归关系不相同,说明δ~(13)C值与叶属性的关系随物种、环境以及叶片年龄的变化而有所不同。
6.对影响叶片δ~(13)C值的13个叶属性因子进行主成分分析表明,不同年龄叶片均提取了前五个主分量,综合了80%之多的叶属性因子的信息,且每个主分量的特征根大于1。不同叶龄叶属性指标的每个主分量所综合的主要生理因子不同,主成分回归分析表明,1年生叶片各叶属性指标与δ~(13)C值的相关性由大到小依次为:LDMC>Na含量>P含量>Ca含量>Mn含量>SLA>Fe含量>灰分含量>叶绿素含量>Zn含量>K含量>Mg含量>N含量,其中,Na含量、SLA、灰分含量、Mg含量与δ~(13)C值呈负相关;2年生叶片各叶属性指标与δ~(13)C值的相关性由大到小依次为:Ca含量>SLA>N含量>P含量>叶绿素含量>LDMC>K含量>Mg含量>Mn含量>Zn含量>Fe含量>灰分含量,其中,SLA、N含量和Mg含量与δ~(13)C值呈负相关;这说明植物在生长过程中,各生长因子对植物所起的作用处于动态变化中。SLA、LDMC、P含量、Ca含量在植物的生长过程中均起着重要的作用,相对于2年生叶片,Fe含量、Mn含量和灰分对于1年生叶片对于δ~(13)C值的影响更大,K含量和Mg含量对各叶龄的δ~(13)C影响程度相当,而2年生叶片叶绿素含量对于δ~(13)C值的影响更大。
7.对主要优势种米槠、九节木、鹅掌柴、微毛柃不同叶龄δ~(13)C值与不同土壤深度的各种养分含量以及坡度、坡向等地形因子进行Pearson相关性分析,表明每一土壤深度的养分含量对叶片δ~(13)C值都有一定的影响。不同叶龄δ~(13)C值与土壤养分含量的相关性不一致,以不同年龄鹅掌柴叶片差异最明显,与2年生叶片相比,1年生鹅掌柴叶片δ~(13)C值与土壤养分含量的相关性更强。叶δ~(13)C值与土壤养分含量主要以正相关为主,说明不同的土壤养分能降低植物叶片的同位素判别,提高植物的水分利用效率。地形因子对叶δ~(13)C值也有一定的影响。
8.为探讨在不局限于某一物种某一特定环境时叶绿素计SPAD的适用性,采用便携式叶绿素仪SPAD-502对灵石山米槠林不同海拔优势种不同年龄叶片的SPAD值进行野外测定,分别对各海拔不同叶龄不同优势种以及各叶龄不同海拔不同优势种的叶片SPAD值双因素方差分析表明,灵石山米槠林优势种群叶片SPAD值存在叶龄、种间以及海拔梯度间的差异。对各海拔优势种不同年龄叶片SPAD值与室内分光光度法测定的叶绿素含量以及叶片N含量进行回归分析,表明,SPAD值与叶片单位面积叶绿素含量及N含量最适合线性关系。SPAD值是叶绿素含量的真实反映,利用叶绿素仪SPAD-502可以快速有效的诊断林木N营养状况。
9.对叶属性指标种间和叶龄间的差异性进行分析,SLA、LDMC以及叶片C、N、P、K、Ca、Mg、Fe、Mn、Zn含量和C/N、C/P、N/P均存在极显著的种间差异性;SLA、LDMC、叶P、K、Ca、Fe、Zn含量以及C/P存在显著的叶龄间差异。1年生叶片的SLA大于2年生叶片,而2年生LDMC大于1年生叶片。对SLA与其它叶属性指标进行回归分析,SLA与不同年龄叶片LDMC、Ca、C/N、C/P成负线性回归关系,与不同年龄叶片N、P、K、Na、Mg、Zn、Ash含量以及2年生叶片Fe、Mn、N/P成正线性回归关系。对不同养分含量及营养结构的关系进行回归分析,不同叶龄N、P、K、Ca、Mg含量与C含量回归关系不显著,不同叶龄N含量与P、K含量以及P含量与K含量之间存在极显著的正线性回归关系,C/N与C/P显著相关,不同叶龄N、P以及1年生叶片K与Ca含量均呈显著负线性回归关系,不同叶龄N、K与Mg元素均呈显著或极显著性正线性回归关系,2年生叶片Ca与Mg含量正线性回归关系,而不同叶龄P与Mg含量回归关系不显著,1年生叶片Na含量、不同叶龄K、Ca、Mg、Fe、Mn、Zn含量与Ash含量均呈显著正回归关系。
10.分别对米槠、九节木、鹅掌柴、微毛柃不同叶龄养分含量与环境的关系进行排序分析,结果表明,土壤养分、土壤PH、海拔、坡度、坡向等环境因子对优势种叶养分含量都有不同程度的影响,但是环境因子对不同优势种养分的影响程度不同,相同优势种不同叶龄养分对环境的响应也不同。通过对4种优势种养分含量与环境进行排序分析不能够分离出哪些环境因子的影响更强。由不同海拔梯度上样地在排序图上的位置可以看出,各样地间存在一定的环境变化梯度。
Castanopsis carlessi forests are important constructive species for the evergreen broad leaves forests. The characteristics and adaptation to environment of leaf have been one of the key and hot points of Ecology. For the first time, foliar stable carbon isotope ratio(δ~(13)C) SLA, LDMC and nutrient content of different age leaves were investigated along the altitudinal distribution gradient of dominant species in Castanopsis carlessi forests in Lingshi shan National Forest Park. The variation ofδ~(13)C and the leaf traits such as concentration of mineral nutrients, specific leaf area(SLA), leaf dry matter content(LDMC) between dominant species, leaf age and the different altitude gradient were analyzed. The relationship between foliarδ~(13)C and leaf traits was studied by regression analysis and principal component. Also the influence of environmental factors on the value of foliarδ~(13)C was analyzed. Meanwhile the correlation between leaf traits and its relation to environmental factors such as soil nutrient status, elevation and slope was studied. The main content of this paper were concluded as follows:
The range of foliarδ~(13)C value was between -34.52‰and -28.377‰, with an average of -30.885‰for Castanopsis carlessi forest in Lingshishan National Forest Park. The variance analysis showed that there was significant difference between dominant species at each altitude gradient except for A7 and A8. And there no significant difference for different age leaves. The frequency distribution of foliarδ~(13)C become single along altitude gradient. Regression analysis showed the value of foliarδ~(13)C was more positive with the increasing altitude, with an average change of 3.2‰? km-1.
Closely related to photosynthesis, chloroplast pigments are vital factors to measure the intensity of photosynthesis. Quantification of chloroplast pigment content could supply the important information about the correlation between plants and environment. The variation of chloroplast pigments between different age leaves and the change patterns of different age leaf chloroplast pigments along elevation gradients were analyzed. The results showed that there were significant or very significant differences of chlorophyll a, chlorophyll b, and chlorophyll(a+b) between different age leaves. Generally, the content of chlorophyll a, chlorophyll b, and chlorophyll (a+b) in two–year-old leaves was higher than in one-year-old leaves and the variations are more substantial at middle and high elevation gradients than at low elevation gradients. Yet the variation of chlorophyll a/b and carotenoid between different age leaves was complicated, and it indicated the adaptation of plants to environment. There were significant variations of the chlorophyll a, chlorophyll b, and chlorophyll (a+b) between different elevation gradients and some certain change patterns were presented. With the increase of elevation gradients, the chlorophyll a, chlorophyll b, and chlorophyll (a+b) content of seven dominant species among the dominant populations become higher, reaching a maximum at the elevation of A5 or A6 and then become lower. And the change patterns of chlorophyll a/b were more diversified with the increase of elevation gradients. As a type of functional pigment, the function of carotenoid was complicated, the response patterns to elevation were diversified too. The results of this research indicated that as an important environment factors elevation had vital and significant impacts on plants and that the dominant species had different adaptation strategies.
The data analysis of leaf nutrient concentration showed generally wide concentration ranges for most mineral nutrients except P and Na. And most tree species were clustered at the lower or middle end of the concentration ranges indicating they have low nutrient status. Among the macronutrients, P and Na had the lowest and narrowest foliar concentration, with the range of 0.259～2.026 g·kg-1 for P, 0.150～1.804 g·kg-1 for Na. Concentration of Ca had widest range. The nutrient concentration of different age leaves was different. The concentration of N, P, K for one-year-old leaves was higher than two-year-old leaves and the concentration of Na, Ca, Fe, Mn, Zn was opposite. A few tree species were observed to accumulate (called‘‘accumulator species’’) some mineral nutrients like N, P, K, Na.
The variation of leaf traits along altitude gradient showed that leaf traits of SLA,C, K, Mg content were lower with the increasing altitude and LDMC was higher. It indicated the adaptation strategies to habitat of the dominant species. The variation of SLA and LDMC demonstrated the adaptation to nutrient inadequate.
The results of regression analysis showed that leaf traits except for leaf organic carbon content had significant linear relationship with the value of foliarδ~(13)C. Theδ~(13)C value of different age leaves was positively correlated with LDMC and the concentration of P, Ca, and negatively correlated with the concentration of N, K, Na, Mg, Ash and chlorophyll. The correlation ofδ~(13)C value and the concentration of Fe, Mn, Zn were multiform at different altitude gradients. For the data all the altitude gradients, theδ~(13)C value wasn’t correlated with the concentration of Fe and Zn and positively correlated with the concentration of Mn of one-year-old leaves, negatively correlated with the concentration of Mn of two-year-old leaves. The results indicated that species, leaf age and environmental factors could influence the relationship ofδ~(13)C and leaf traits.
Results of PCA(Principal Component Analysis) for 13 leaf trait factors effecting foliarδ~(13)C showed that the first five principal components composited 80 percent leaf traits and the eigenvalue of each primary component was greater than 1. Principal component regression analysis showed that the correlation ofδ~(13)C and leaf traits was different for different age leaves. The strength sequence of relevance ofδ~(13)C with leaf traits factors for one-year-old leaves was LDMC, Na, P, Ca, Mn, SLA, Fe, Ash, chlorophyll, Zn, K, Mg and N declining. And the strength sequence for two-year-old leaves was Ca, SLA, N, P, chlorophyll, LDMC, K, Mg, Mn, Zn, Fe, Ash.This showed that the role of factors playing on plant growth was in dynamic change. SLA, LDMC, concentration of P and Ca played important roles on plant growth. Compared with two-year-old leaves, concentration of Fe, Mn and Ash played more important roles on two-year-old leaves. And concentration of chlorophyll was vital forδ~(13)C value of two-year-old leaves.
The Pearson correlation was analyzed between foliarδ~(13)C and environmental factors such as soil nutrient status at different depth, elevation and slope was studied. Soil nutrient status at each depth could influence value of foliarδ~(13)C.The correlation between foliarδ~(13)C and soil nutrient content was mainly on positive. We could conclude that soil nutrients can reduce the isotope discrimination of plant leaves and improve water use efficiency of plants. Topographical factors could influence value of foliarδ~(13)C too.
Portable chlorophyll meter SPAD-502 was used to measure the greenness (SPAD) of different age leaves of dominant species at different altitude gradients. There was different variation of SPAD value between the different age leaves, different dominant species and different altitude gradients. And the correlation between SPAD value and chlorophyll and foliar N concentration showed that significant correlations between SPAD value and concentration of chlorophyll and foliar N were observed,indicating that SPAD value could not only well reflect the chlorophyll concentration measured by spectrophotometry method, but also a useful tool for nondestructively assessing foliar N status. In the case of comparison in a relative value instead of real magnitude, SPAD value could be indicators for foliar chlorophyll and nitrogen, which may have efficient application for forest assessments in decision-making and operational nutrient management programs.
Variance analysis showed that there was significant variation between dominant species for leaf traits such as SLA, LDMC, the concentration of C, N, P, K, Ca, Mg, Fe, Mn, Zn and the ratio of C/N, C/P, N/P. And there was significant variation between different age leaves for SLA,LDMC the concentration of P, K, Ca, Fe, Zn and C/P. The correlation between SLA, LDMC and leaf nutrient content were analyzed. SLA was positively correlated with concentration of N, P, K, Na, Mg, Zn, Ash for different age leaves and positively correlated with concentration of Ca, Fe, Mn and ratio of N/P for two-year-age leaves. And it was negatively correlated with LDMC, Ca, C/N and C/P. There was significant regression relation between leaf nutrient concentrations and nutrient structures too. We can conclude that the leaf traits were the representation of adaptation to environment long-term.
Redundancy analysis (RDA) of environmental variables and foliar nutrient concentration showed that foliar mineral nutrient concentration of the trees was generally correlated with the environmental factors. However, the effects were interrelated and difficult to isolate from each other. The influence of environmental factors on different dominant species and was different. The strategy of resources use was different for dominant species, which was benefit to coexistence of species long-term.
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