974 resultados para Leaves.


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An LCAO-scheme taking into account 10 atomic orbitals (s-, p-, and d-type) is used to calculate the electronic structure of the reconstructed 90-degrees partial dislocation in Si. Two different valence force fields producing deviating results are used for modelling the core structure. Geometrical data published by another group is also used. The aim is to explore the influence of geometry on energy levels. We find that the band structure depends sensitively on bond angles. Using data determined by the Tersoff potential we obtain two bands of which the upper one penetrates deeply into the indirect band gap while the geometry minimizing the simple Keating potential leaves the gap completely clear of dislocation states. Thus, from a theoretical point of view, the chief difficulty in calculating the electronic structure of the reconstructed 90-degrees partial is the lack of accurate structural information.

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We have studied the spontaneous emission of polarized excitons in the GaInP/AlGaInP VCSEL from 30K to room temperature. It is observed that the spontaneous emission peak enters and leaves the resonant regime. At the resonant regime, the emission intensities of the perpendicular and horizontal polarized exciton are enhanced at different ratio to those in non-resonant regime. These experiment results are explained through the dressed exciton theory of the semiconductor microcavity device. From this theory, the intensity enhancement and the polarization dependence are understood as cooperative emission and the microcavity anisotropy.

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文中提供了一种对植物叶子造型的新方法,可以非常真实地展现叶子在干枯、老化过程中几何形态的变化.基于植物学和物理原理,文中引入了双层结构模型表达叶子的力学结构,很好地模拟了不同种类植物叶子形状的多样性.文中首先分析了叶子形状变化的原因,即在叶子枯萎过程中,叶肉和叶脉由于各自不同的组织结构,导致了收缩比例不同,从而在物理模拟中,通过建立关于叶子基本结构的双层质点-弹簧模型,并对上下两层的不同参数的合理设置,很好地表现了叶肉和叶脉的不同力学特性.双层模型的相互作用,决定了叶子最终变形的方式和效果,由此可以得到非常接近真实树叶的各种叶子形态.

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探讨全球气候变化的生物学和生态学效应是当今生态学中的热点,研究大气CO2浓度升高对植物-昆虫相互作用关系的影响具有重要的理论和实践意义。本文使用开顶式气室(Open-top chamber,OTC)在野外条件下研究了CO2浓度升高对三种树木(小青杨、白桦和蒙古栎)叶片化学成分含量的影响,以及树木叶片品质变化对一种广食性森林昆虫(舞毒蛾)幼虫取食、生长发育和取食偏嗜性的影响。得出如下结果:(1)CO2浓度升高对3个受试树种叶片中的营养成分及次生代谢物含量均有显著影响,总体表现为氮含量降低,而碳氮比、非结构性碳水化合物、总酚和缩合丹宁含量增加。叶片中的化学成分含量可随时间发生显著变化,不同树种、甚至同一树种不同冠层高度的叶片对CO2浓度升高的响应强度也是不同的。叶片的干物质含量和比叶重对CO2浓度升高的响应不显著。(2)室内非选择性取食实验、室内选择性取食实验以及上树取食饲养方式下的多龄期取食实验,均发现高浓度CO2处理组内舞毒蛾幼虫的生长发育受到显著抑制。但对四龄舞毒蛾幼虫所进行的短期生物测定并未发现不同CO2浓度处理下幼虫的生长发育速率、对食物的取食率和转化率等昆虫营养指标存在显著差异。(3)叶片品质的降低是导致舞毒蛾幼虫生长发育受抑制的主要原因。但是总体上,CO2浓度升高导致的叶片品质变化并未显著影响幼虫的取食率和取食量。(4)舞毒蛾幼虫对不同叶片种类表现出清晰的取食选择性,这种选择性在其幼龄期就可表现出来。幼虫对小青杨上层叶片有最显著的偏嗜性,对蒙古栎下层叶片有最明显的拒食性。但是CO2浓度升高导致的叶片品质变化对舞毒蛾幼虫的取食选择性和寄主偏嗜行为并未产生显著影响。(5)检测出高浓度CO2处理组内舞毒蛾幼虫虫粪中含有浓度更高的植物次生代谢物质(总酚和缩合单宁),这很可能是昆虫整体生长发育受抑制的重要原因之一。

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采用开顶箱法模拟大气O3和CO2浓度升高(80 nmol•mol-1,700 μmol•mol-1),对银杏(Ginkgo biloba)和油松(Pinus tabulaeformis)幼树进行了连续两个生长季的熏蒸试验,研究了银杏叶片和油松针叶活性氧产生清除系统的生理变化过程,从活性氧代谢生理机制上揭示其抗性变化规律,为研究城市森林对全球变化的响应与反馈机制提供重要的理论基础。 得出如下结论: 1. 高浓度O3显著增加了银杏叶片ASA含量和抗氧化酶(SOD、APX)活性,增强了活性氧清除能力;但这种抗氧化能力的提高不足以消除高O3带来的氧化逆境,H2O2和MDA含量显著高于对照处理。与银杏相比,高浓度O3在试验初期诱导油松针叶SOD和APX酶活性升高,增强了活性氧清除能力;随着处理时间的延长ASA被耗竭,H2O2过量积累显著地抑制了抗氧化酶活性,加剧了氧化伤害。银杏和油松结果的差异表明,银杏对80 nmol•mol-1浓度的O3具有更高的抗性,而油松针叶活性氧代谢等生理过程受其影响更大。 2. 高浓度CO2对银杏叶片活性氧代谢无显著性影响。然而,高浓度CO2减缓了油松针叶活性氧的产生,抗氧化酶活性呈降低的趋势;但针叶内ASA含量显著高于对照处理,这说明高浓度CO2促进了油松针叶ASA合成,或者是减少了ASA的消耗。在试验处理后期,针叶内H2O2含量和MDA含量显著低于对照处理。 3. 高浓度O3和高浓度CO2复合处理中,高浓度CO2缓解了高浓度O3对银杏叶片和油松针叶造成的氧化胁迫,但其生理机制不同:银杏叶片ASA含量以及抗氧化酶活性普遍低于高浓度O3单因素处理而高于对照处理,并且叶片H2O2含量和膜质过氧化产物MDA含量显著低于高浓度O3单因素处理,这说明高浓度CO2通过减少活性氧的产生来缓解高浓度O3所致的活性氧积累以及膜质过氧化伤害。与银杏不同的是,高浓度CO2使油松针叶内ASA含量显著高于高浓度O3处理,增强了活性氧清除能力,减轻了活性氧对抗氧化酶活性的抑制作用,有效的控制了ROS的产生与清除之间的平衡,缓解了高O3带来的氧化伤害。

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文冠果(Xanthoceras sorbifolia Bunge)是我国北方特有的木本油料树种,已被列为生物质能源树种之一。现阶段产量低是限制其发展的重要因素。本文以内蒙古赤峰市翁牛特旗文冠果经济林场内50年生文冠果树为研究对象,对文冠果生物学、生态学特性进行定位观测和试验,分析了文冠果发育节律与积温的关系。野外布置水肥措施试验,测定了文冠果产量和叶片、果实养分元素(N、P、K、Mg、Ca、Fe、Zn、Cu、Mn)的含量,分析了水肥措施对叶片、果实养分分配的影响。 主要结论: 1.文冠果物候期与积温呈显著的直线函数关系。能够用来预测文冠果年生长发育期有效积温的理论值,为经营管理提供依据。 2.文冠果叶片中主要元素含量随物候期的变化呈下降趋势,但微量元素呈增加趋势。养分重吸收水平由大到小的顺序是:Mg>N>K>P,一定的水肥措施提高了文冠果养分再吸收水平,并降低了功能叶中的N/P、C/N。 3.文冠果果实中Ca和Fe随物候期的变化呈下降趋势,其他元素呈先减后增趋势。水肥措施降低了果实对养分元素的分配和收获系数;提高了养分利用效率。 4.合理的水肥措施能显著提高种子产量,增加含油率。最佳水肥措施为:5月初灌水50mm,施肥量为N 63 g•株-1、P2O5 83.3 g•株-1,于5、7月初分2次等量施入。 5.水分、P是文冠果生长的主要限制性因子,限制性因子具有阶段性。水分对肥效有“激发效应”。

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本论文由四章组成。第一、二和三章分别报道了双花千里光、川芎和宽叶羌活的化学成分研究。从三种药用植物中共分离和鉴定了40 个化学成分,其中8个为新化合物。第四章概述了藳本属植物及日本川芎的化学成分研究进展。 第一章包括三个部分。第一部分报道双花千里光(Senecio dianthus Franch.)地上部分乙醇提取物的化学成分。采用正、反相硅胶柱层析等各种分离方法,从中共分离出8 个艾里莫酚型倍半萜内酯,其中5 个是新化合物,并且有1 个为首次发现的连接了含氮原子取代基的艾里莫酚型倍半萜内酯。它们的结构经MS、IR、NMR及X-单晶衍射等解析方法确定为2b-angeloyloxy-10b-hydroxyeremophil-7(11)-en-8a,12-olide (1)、6b-angeloyloxy-10b-hydroxyeremophil-7(11)-en-8a,12-olide (2)、2b-angeloyloxy-8b,10b- dihydroxyeremophil-7(11)-en-8a,12-olide (3)、2b-angeloyloxy-8a-hydroxyeremophil-7(11),9(10)-dien-8b,12-olide (4)和8b-amino-10b- hydroxyleremophil-7(11)-en-8a,12-olide (5)。这8 个倍半萜内酯经体外生物活性测试表明均具有通过抑制巨噬细胞增殖抵制破骨细胞增生的活性。第二部分对艾里莫酚型倍半萜内酯的质谱裂解规律进行了初步探讨。第三部分报道双花千里光茎、和叶花的挥发油成分分析。采用传统水蒸气蒸馏法分别提取了双花千里光茎、叶和花的挥发油,用气相色谱-质谱联用(GC-MS)技术分别分离鉴定了其化学成分,从茎、叶和花挥发油中各分离和鉴定出70、80 和73 种化学成分,分别占挥发油总量的91.2%、85.7%及93.4%。 第二章包括两个部分。第一部分报道川芎(Ligusticum chuanxiong Hort.)根茎乙醇提取物的化学成分。通过正、反相硅胶柱层析等分离纯化和MS、NMR及X-单晶衍射等解析方法,共分离鉴定了21 个化合物,结构类型分属于苯酞、二聚苯酞、香豆素和脂肪酸类。其中2 个为结构比较新颖的二聚苯酞类化合物:chuanxiongnolide A (19)和chuanxiongnolide B (20),化合物19 的结构经X-单晶衍射得到确证。第二部分报道川芎挥发油的化学成分。采用不同的提取方法(溶剂萃取法、水蒸气蒸馏法、CO2 超临界流体萃取法)提取川芎挥发油,同时采集不同产地(四川彭县、四川郫县、云南鹤庆)及不同品质(川芎、奶芎、苓子)的川芎产品,利用GC-MS 技术分离鉴定其挥发油的化学成分,计算各成分的相对含量,并对比分析其中的异同。 第三章报道宽叶羌活(Notopterygium forbesii Boiss.)根茎化学成分的分离纯化和结构鉴定。通过正、反相硅胶柱层析等分离纯化和MS、NMR 等解析方法,共分离鉴定了13 个化合物,结构类型分属于香豆素、二氢异香豆素、甾体和羧酸类。其中1 个新二氢异香豆素类成分鉴定为6-methoxy-hydrangenol (37)。 第四章概述了藳本属植物及日本川芎化学成分的研究进展。 This dissertation consisted of four chapters. The former three chaptersrespectively elaborated the phytochemical investigation of three herbal medicines:Senecio dianthus Franch., Ligusticum chuanxiong Hort. and Notopterygium forbesiiBoiss.. Forty compounds, including eight new ones, were isolated and identified byspectral and chemical evidence. The fourth chapter elaborated the study progress ofchemical constituents of Ligusticum genus and Cnidium offcinale. The first chapter consisted of three parts. The first part is about the chemicalconstituents of ethanol extraction and essential oils from the aerial parts of S. dianthu.Eight eremophilenolides were isolated and identified. Among them, five ones are newcompounds and one of them is a novel eremophilenolide attched with an amino group.The structures of the new compounds were identified as 2b-angeloyloxy-10b-hydroxyeremophil-7(11)-en-8a,12-olide (1),6b-angeloyloxy-10b-hydroxyeremophil-7(11)-en-8a,12-olide (2),2b-angeloyloxy-8b,10b-dihydroxyeremophil-7(11)-en-8a,12-olide (3),2b-angeloyloxy-8a-hydroxyeremophil-7(11),9(10)-dien-8b,12-olide (4) and8b-amino-10b-hydroxyeremophil-7(11)-en-8a,12-olide (5) by spectral evidence andX-ray crystallography analysis. All the compounds were evaluated for theiranti-osteoclstogenesis activity using a proliferation inhibit assay with microphagecells. The second part elementarily discussed the characteristic fragmentation oferemophilenolides isolated from S. dianthus in ESI-MS.The latter part is about thechemical constituents of essential oil extracted from stems, leaves and flowers of S.dianthus with steam distillation. By the GC-MS analysis, 70, 80 and 73 compoundswere respectively isolated and identified which accounted for more than 91.2%, 85.7% and 93.4% of total essential oil. The second chapter, including two parts, is about the the chemical constituents ofethanol extraction and essential oils from rhizomes of L. chuanxion. In the first part, twenty-one compounds were isolated and iedntified. Two ones are novel dimericphthalides and the structures were suggested as chuanxiongnolide A (19) andchuanxiongnolide B (20) by spectral evidence and confirmed by X-raycrystallography analysis. In the second part, the samples were collected from differentextract techniques (solvent extraction, steam distillation and supercriticalfluid extraction), different habitats (Peng and Pi counties, Sichuan province; Heqing,Yunnan province) and different qualities (Chuanxiong, Naixiong and Lingzi). Thechemical constituents of essential oil from L. chuanxiong were analyzed by GC-MS and were compared each other. The third chapter is about the chemical constituents of rhizomas of N. forbesii,which belongs to a endemic genus of China. Thirteen compounds were isolated andidentified. One of them is a new dihydroisocoumarin and the structure was identifiedas 6-methoxy-hydrangenol (37) by spectral evidence. The fourth chapter is a review on study progress of chemical constituents ofLigusticum species and Cnidium offcinale.

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近年来,随着对作物重茬障碍原因的深入研究,植物的化感作用越来越受到国内外众多学者的重视。花椒(Zanthoxy piperitum.)为芸香科植物,是一种收益早、用途广、价值高的经济树种,是川西干旱河谷地区的重要经济作物,其连作障碍也倍受关注,系统研究花椒化感作用将有助于理解和最终解决花椒连作障碍问题。本文首先通过萃取、层析等方法分离花椒主效化感成分;通过外加不同浓度的花椒叶水浸液研究了对土壤氮素养分循环的影响;研究了花椒叶水浸液对苜蓿生理生化、光合作用、氮素养分吸收的影响,并对外施氮肥对这种化感影响的缓解作用做了研究;研究了花椒化感潜力对全球变化——UV-B增强辐射的响应。主要研究结果如下: 1.用不同极性的有机溶剂对花椒叶水浸液浓缩浸膏萃取、柱层析,结合生物活性检测,分离得到主效化感作用组分的一种化感物质——对甲氧基苯酚。采用该物质纯品进行生物活性检测,证明其具有化感作用。 2.花椒叶水浸液处理土壤30天后,土壤硝态氮、铵态氮、无机氮(硝态氮+铵态氮)与对照相比,随着花椒叶水浸液浓度的增加呈现降低的趋势,其中土壤铵态氮含量显著降低,而硝态氮含量的变化则不显著,无机氮含量也显著降低。土壤脲酶和蛋白酶的活性与无机氮含量的变化趋势相同。随着花椒叶水浸液浓度的增加,氨化细菌数量显著降低,固氮菌的数量变化不显著,硝化细菌和反硝化细菌数量有减少的趋势。60天后,硝态氮含量、铵态氮含量、无机氮随水浸液浓度增加的变化趋势与30天时相似;随着花椒叶水浸液浓度的增加,氨化细菌、固氮菌的数量显著减少,硝化细菌数量、反硝化细菌数量仍呈减少趋势;土壤脲酶、蛋白酶活性与第30天的变化趋势相同。第60天与第30天的结果相比,相同水浸液浓度处理的硝态氮、铵态氮、无机氮均有下降的趋势,但除了25g.L-1水浸液处理的外,其它相同浓度的处理间差异均不显著;除了12.5 g.L的处理外土壤脲酶活性均呈增强的趋势;蛋白酶活性都有不同程度的增加;花椒叶水浸液处理的土壤硝化细菌和反硝化细菌数量呈增加趋势。 3.随着花椒叶水浸液浓度的增加,显著抑制了苜蓿根长、地上地下生物量、叶绿素含量、叶片中可溶性蛋白的含量,净光合速率。苜蓿体内四种抗氧化酶(POD、SOD、CAT、APX) 活性随着水浸液浓度的增加而降低,而丙二醛含量则增加。苜蓿氮初级同化相关酶硝酸还原酶(NR)、谷氨酰合成酶(GS)、谷氨酸脱氢酶(GDH)的活性随着水浸液浓度的增加受到不同程度的影响。总的来说,苜蓿硝酸还原酶、谷氨酰合成酶的活性受到抑制,而谷氨酸脱氢酶活性的变化则比较复杂,根呈先降低后增加的趋势,叶片则无显著变化。外施两种不同浓度的硝酸铵氮肥后,对12.5、25 g.L-1花椒叶水浸液处理的苜蓿化感作用有显著的缓解作用,表现在株高、生物量、光合作用等方面,大多达到与对照(0 g.L-1)未施氮肥无显著差异的水平,而对50 g.L-1水浸液处理的苜蓿幼苗,虽有一定的缓解作用,但这种作用均未达到与对照(0 g.L-1)未施氮肥时无显著差异的水平。 4. UV-B增强辐射处理花椒后,花椒的化感潜力显著增强。花椒叶片内UV-B吸收物质的含量和总酚含量均显著增加。 In recent years, with profound research on the reasons of continuous cropping obstacles, allelopathy received increasing attention to many scholars at home and abroad. Zanthoxy bungeanum as a Rutaceae plant is a high economic value species which gains early and uses widely. Zanthoxylum is an important economic crop in the arid valley of western Sichuan region, and its not even has received much concern for the continuous cropping obstacles. The systematic study of allelopathy of Zanthoxylum will contribute to the understanding and final settlement of this issue. The major allelopathic composition was separated through the extraction, chromatography combined with other methods. The impact on soil nutrient cycling was also studied through the addition of different concentrations of water extracts of Zanthoxylum. Furthermore, the effects of water extracts of Zanthoxylum leaves on alfalfa leaf physiological and biochemical indexes, photosynthesis, soil enzymes and nutrient uptake of nitrogen and the mitigation of allelopathy through using external fertilizer were studied to put forward scientific resolvent for Zanthoxylum continuous cropping obstacles .The response of allelopathic potential of Zanthoxylum to global change - UV-B enhanced radiation was studied . The main findings are as follows: 1. Through extraction with different polar organic solvents on concentrated water extract of Zanthoxylum leaf and then using column chromatography combined with detection of biological activity, one of the main allelopathic components- methoxy-phenol was isolated. The biological activity testing of the pure material of methoxy-phenol proved that it does have allelopathic potential. 2. Thirty days after treating soil with water extract of Zanthoxylum leaf, as compared with the control, the contents of soil nitrate, ammonium, nitrate plus ammonium nitrogen showed a trend of decrease with the increase of the concentration of water extract whereas the content of ammonium nitrogen showed a significant reduction, and the content of nitrate did not change significantly, the content of nitrate plus ammonium nitrogen also showed a significant (P <0.05) redction. The activity of soil urease and protease showed the same trend as the content of nitrate nitrogen plus ammonium nitrogen. With the increase in the concentration of water extract, the number of ammonification bacteria significantly reduced but nitrogen-fixing bacteria did not change significantly and there was a decreasing trend in the number of nitrifying bacteria and denitrifying bacteria. Sixty days after the treatment, with the increase in solution concentration of water extract of Zanthoxylum leaf, the content of nitrate、 ammonium nitrogen, nitrate plus ammonium nitrogen showed a similar change trend to 30 days’; the number of ammonification bacteria, nitrogen-fixing bacteria significantly reduced ; the number of nitrifying bacteria, denitrifying bacteria was still an downward trend; the activity of soil urease and protease showed the same trend as the 30th days’. Compared to the results of the 30th days’, the content of nitrate, ammonium, nitrate plus ammonium nitrogen showed a decrease trend between the treatment of same concentration, but there was no significant difference except the treatment of 25g.L-1 between the same concentration; the activity of soil urease showed enhanced trend except the treatment of 12.5 g.L-1; the activity of protease increased to varying degrees; the number of ammonification bacteria、 nitrifying bacteria and denitrifying bacteria were growing while nitrogen-fixing bacteria reduced.. 3. With the increase of the concentration of water extract of Zanthoxylum leaf, the water extract significantly inhibited the root length, aboveground biomass, content of chlorophyll and soluble protein in leaf and net photosynthetic rate. The activity of four antioxidant enzymes (POD, SOD, CAT, APX) reduced with the increase in concentration of the water extract but the content of MDA increased. The activity of enzymes related to primary nitrogen assimilation such nitrate reductase (NR), glutamyl synthetase (GS), glutamate dehydrogenase (GDH) were subject to different degrees with an increase in the concentration of water extracts. In general, the activity of nitrate reductase, glutamyl synthetase were inhibited, while change in the activity of glutamate dehydrogenase was more complex. The activity of glutamate dehydrogenase in leaf was first reduced and then increase,but did not change significantly in root. After using two external different concentrations of nitrogen fertilizer, there was a significant mitigation in inhibiton in plant height, biomass, photosynthesis, etc. in the treatment of 12.5,25 gL-1 of water extract of Zanthoxylum leaf, and most of these indexes showed no significant difference with the control (0 g.L-1, no external fertilizer was added) .Although there showed a certain degree of ease in the treatment of 50 g.L-1 , there was still a significant difference compared with the control (0 gL-1) in which no external fertilizer was used. 4.The allelopathic potential of Zanthoxylum positively responded to enhanced UV-B significantly. The content of UV-B absorbing compounds and the total phenol also significant increased.

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土壤微生物量、可溶性有机碳与氮虽然只占土壤有机碳、氮总量的较小部分,但可以在土壤全碳、氮变化之前反映土壤微小的变化,又直接参与土壤生物化学转化过程,因而在植被恢复过程中,较其它土壤理化性质等能够更好地指示土壤恢复情况。在青藏高原东缘存在大面积的次生人工林替代灌丛或采伐迹地,而关于这些人工林替代后的生态效果和生态过程的评估却十分缺乏,本研究通过评估岷江上游植被恢复重建过程中典型人工替代次生植被凋落物层与土壤碳、氮等养分大小,动态监测土壤微生物生物量、水溶性碳、氮等指标,结合温度与凋落物输入等影响土壤活性有机碳、氮因子的控制试验,系统分析不同人工替代次生植被土壤碳、氮等养分的差异原因,试图寻找低效人工林优化调控与持续管理技术,为区域生态公益林持续管理提供理论和技术依据。主要结论如下: 1. 通过对不同人工替代次生植被凋落物层和土壤碳、氮分析发现,油松和华山松人工林替代次生灌丛后土壤碳、氮含量较灌丛和阔叶人工林低,主要原因可能为凋落物质量(C/N)较差,而引起碳、氮等元素难以归还土壤。进而通过对不同人工替代次生植被凋落物层和土壤微生物生物量、水溶性有机碳、氮等指标的季节性动态模式的分析,发现各次生植被土壤微生物生物量C、N,P以及土壤水溶性碳、氮含量均呈明显季节性动态,呈现秋季明显大于其它季节,冬季最低,在表层土壤最为明显。 2. 油松、华山松人工林凋落物层和土壤水溶性有机碳(WDOC)、土壤水溶性有机氮(WDON)明显低于灌丛和连香树,土壤微生物生物量C、N也以油松和华山松人工林最低,而落叶类植被,如灌丛、连香树和落叶松之间没有明显差异,说明可利用底物的数量和质量差异是影响各次生植被凋落物分解和土壤微生物活性的主要原因。MBC/OC和MBN/ON能较好地指示土壤微生物活性的变化,MBC/OC凋落层总体以灌丛和连香树人工林最高,油松和华山松人工林最低;而土壤中MBC/OC连香树人工最高,华山松人工林最低。说明以油松和华山松为主的人工造林替代乡土阔叶灌丛造成土壤C、N等养分严重匮乏,微生物活性低下是影响其养分周转的主要原因。 3. 从各次生植被凋落物产生看,凋落物年归还量最大的为华山松人工林(5.1×103 kg ha-1),其次为落叶松人工林(4.8×103 kg ha-1),阔叶灌丛林地凋落物产生总量(4.4×103 kg ha-1)略大于油松人工林(4.2×103 kg ha-1),最小的为连香树人工林(3.6×103 kg ha-1);叶是凋落物的主体,落叶类树种月动态表现为单峰型,高峰主要在10-11月,如落叶松、连香树和灌丛林;常绿的松类月动态不明显,各月基本相同,最为明显地为油松林,华山松人工林略有二个小峰,分别出现在11月和5月。落叶阔叶灌丛的凋落物分解速率大于常绿针叶林,如油松和华山松。结合凋落物的产生量和分解速率,不同树种人工林替代次生阔叶灌丛后,人工油松和华山松林枯落物总贮量和厚度明显大于落叶松人工林、灌丛林和连香树人工林,说明以油松和华山松为主的人工造林替代乡土阔叶灌丛延缓了有机物向土壤的顺利归还,不利于土壤C、N等养分循环。 4. 通过控制地面凋落物和地下根系输入有机物对土壤碳、氮的影响研究发现,(1) 单独去除根系以及根系与地面凋落物同时去除处理1年后对表层(0-10cm)土壤WDOC均没有显著影响,而土壤WDON显著增加,油松人工林土壤微生物生物量C、N显著降低,人工落叶松林没有显著差异,说明油松人工林土壤微生物活性对地下碳输入的依赖大于其它次生植被,而落叶松土壤微生物活性对地下碳输入依赖性较小;去除地面凋落物,明显降低了落叶松人工林土壤WDOC,华山松和连香树土壤WDON均较对照显著减少,油松人工林土壤微生物量C较对照显著减少;双倍增加地面凋落物处理对土壤微生物生物量、WDOC和WDON没有明显地增加,相反,连香树、华山松和油松人工林土壤WDON较对照减少。说明油松人工林微生物活性不仅依赖于地下碳输入,而且对地上有机物输入的依赖性也较大;连香树、落叶松和华山松人工林土壤微生物生物量并没有因地面凋落物的去除减少可能与土壤总有机碳含量及活性均较高有关,而双倍增加地面凋落物反而降低了土壤微生物生物量,说明凋落物覆盖后改变了土壤微气候。 5. 碳矿化累积量与有机碳含量和活性有机碳含量之间存在显著地正相关关系。凋落物碳累积矿化量、矿化速率以连香树最高,油松和华山松人工林次之,落叶阔叶灌丛低于常绿针叶纯林,导致其差异的主要原因可能为凋落物产生的时间动态模式不一样,致使凋落物起始分解时间不一致。而土壤层有机碳矿化速率和矿化量以阔叶落叶灌丛和连香树最高,油松和华山松人工土壤最低,再次证实利用针叶纯林恢复植被阻碍了有机质周转与循环。 6. 凋落物累积矿化量与C/N值呈显著地相关关系,并随着温度的升高而明显增加,而土壤累积矿化量与C/N值没有显著相关关系,说明土壤有机碳质量(C/N)对温度的响应不十分明显。通过双指数模型对不同温度下碳矿化过程进行模拟和计算出活性有机碳与惰性有机碳比例,发现温度升高促进了惰性有机碳向活性有机碳的转化,增加了活性有机碳含量,说明温度升高可促进次生植被凋落物与土壤有机质的分解,进而可影响到林地碳源/汇关系的变化。 综上,通过对不同人工替代次生植被凋落物与土壤C、N大小、以及土壤微生物生物量、水溶性C、N等指标动态变化模式研究,结合温度与凋落物数量输入等影响土壤活性C、N因子的综合分析,以油松和华山松人工纯林对山地植被恢复,延缓或阻碍了有机质周转与循环,造成了土壤肥力退化。对现有低效人工纯林改造,应为地面大量有机物分解创造条件。 Although soil microbial biomass, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) are a small part of total soil organic carbon and nitrogen, they can directly participate in the process of soil biochemical translation and indicate the fine changes before changes of soil total organic carbon and nitrogen occur. So, they are good indexes to indicate soil restoration condition during the process of vegetation rehabilitation. There are large areas of secondary vegetations which substitute for indigenous shrubs in the eastern fringe of Qinghai-Tibet Plateau. However, it is not well known that the ecological effect and process after substitution by different secondary plantations. Based on comparison of soil organic and nitrogen contents in litter layer and soil under different secondary vegetations in upper reaches of Minjiang River, soil microbial biomass, DOC and DON in litter layer and soil were investigated in order to analyze the seasonal dynamic. Combining the effects of temperature, litter input and root exclusion on soil microbial biomass, DOC and DON, we also aim to understand the reason and mechanism of difference in soil carbon and nitrogen contents among different secondary vegetations. The study would contribute to comprehensively understanding C and N cycling processes and provide optimal control and sustainable technology of low-effect plantations in these regions. The results are as follows: (1) Organic carbon and nitrogen in litter layers and soil under different substitution plantations were investigated. The results showed that contents of soil organic carbon and nitrogen were lower in P. tabulaeformis (PT) and P. armandi Franch(PA) than those in native broad-leaf shrub and broad-leaf plantation. The low quality (C/N) of litter in PT and PA plantations caused carbon and nitrogen returning to soil difficultly. Seasonal dynamic of soil microbial carbon (MBC),-nitrogen (MBN),-phosphor (MBP), and WDOC and WDON showed similar pattern, which had the highest values in autumn and the lowest values in winter. (2) WDOC and WDON in litter layers and soil under PT and PA plantations were significantly lower than those in native broad-leaf shrub and Cercidiphyllum japonicum Sieb. et Zucc.(CJ). Soil MBC and MBN were also the lowest, while there were no significant differences among deciduous vegetations, i.e. native broad-leaf shrub, CJ and Larix kaempferi Lamb.(LK) plantation. The results suggested that difference in quantity and quality of available substance was main reason that affected the activity of microbe in soil and litter layer. MBC/OC and MBN/ON were good indexes to indicate the change of soil microbial activity. MBC/OC of litter had the highest value under native broad-leaf shrub and CJ plantation, and had the lowest value in PT and PA plantations, while MBC/OC of soil was the highest under CJ plantation, and was the lowest in PT and PA plantations. These results indicated that PT and PA plantations substituting for native broad-leaf shrub caused deficit of carbon and nitrogen in soil, low microbial activity was a main reason influencing the cycling and turnover of carbon and nitrogen in soil. (3) The annual litter fall production, composition, seasonal dynamic and decomposition of five typical secondary stands in upper reaches of Minjiang River were studied in this paper. The annual litter productions were: PA (5.1×103 kg ha-1), LK(4.8×103 kg ha-1), native broad-leaf shrub (4.4×103 kg ha-1), PT(4.2×103 kg ha-1),CJ(3.6×103 kg ha-1). The litter production of leaves in five secondary vegetations occupied a higher percentage in the annual total litter production than those of other components. The litterfall was mostly producted in the cool and dry period (October-November) for deciduous vegetations and relatively equably producted in every season for evergreen coniferous vegetations. The decomposition rate of leaf litter in the broad-leaf stand was higher than those in evergreen coniferous stand. Combined with annual litter fall production and decomposition rate of leaf litter, we found that stock and depth of litter layer were significantly larger in PT and PA plantations than those in native broad-leaf shrub, LK and CJ plantations. The results confirmed that PT and PA plantations substituting for native broad-leaf shrub delayed organic matter returning to soil and hindered cycling of carbon and nitrogen again. (4) We explored plant litter removal, double litter addition, root trenching, and combining root trenching and litter removal treatments to examine the effects of above- and belowground carbon inputs on soil microbial biomass, WDOC and WDON in four secondary plantations. During the experimental period from June 2007 to July 2008, 1 year after initiation of the treatments, WDOC in soil did not vary in root trenching, and combining root trenching and litter removal treatments, while WDON in soil significantly increased compared with CK treatment. Root trenching reduced soil MBC and MBN in PT plantation, while MBC and MBN in soil did not vary in LK plantation. The rasults implied that soil microbial activity was more dependent on belowground carbon input in PT plantation than those in other secondary plantations, on the contrary, soil microbial activity in LK plantation was not dependent on belowground carbon input. Plant litter removal significantly decreased soil WDOC in LK plantation, decreased WDON in PA and CJ plantations, and also significantly reduced soil MBC in PT plantation. However, double litter addition did not increase soil microbial biomass, WDOC and WDON, on the contrary, soil WDON in CJ, PA and PT plantations were decreased. These suggested that soil microbial activity was not only dependent on belowground carbon input, but also on aboveground organic material input. Double litter addition could change the microclimate and result in the decrease of soil microbial activity in CJ, PA and PT plantations. (5) We measured carbon mineralization in a 107 days incubation experiment in 5℃,15℃ and 25℃. Carbon cumulative mineralization was positively correlated with organic matter and labile organic carbon in litter layer and soil. Cumulative carbon mineralization and mineralization rate of litter layers in PT and PA plantations were higher than that in native broad-leaf shrub. This difference between native broad-leaf shrub and coniferous plantations in cumulative carbon mineralization and mineralization rate of litter layers could be attributed to the initiating time of decomposition due to the difference in seasonal dynamic of litter fall production between two types of secondary plantations. However, cumulative carbon mineralization and mineralization rate in soil were the highest in native broad-leaf shrub and CJ plantation, and were the lowest in PT and PA plantations. This also confirmed that PT and PA plantations substituting for native broad-leaf shrub hindered the cycling and turnover of organic matter again. (6) Carbon cumulative mineralization was positively correlated with C/N in litter layer and increased with temperature increasing, while carbon cumulative mineralization was not correlated with C/N in soil. This indicated that soil organic matter quality (C/N) was insensitive to temperature. Applying bi-exponential model, we computed the percent of labile and stable carbon in different temperature incubation and found that temperature increasing would accelerate the transform from stable carbon to labile carbon and increase the percentage of labile organic carbon. This illuminated that temperature incraesing could facilitate the decomposition of litter and soil organic matter in secondary vegetations and hence affect the relationship between carbon source and sink.

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随着工业化的发展,大气中二氧化碳的浓度(CO2)预测从现在的平均350μmol·mol-1升高到2030年的570μmol·mol-1,其增温作用将持续多个世纪。植被在大气二氧化碳减排以及调控区域水热状况过程中起重要作用,而其机理过程目前十分不清楚。本实验应用自控、封闭、独立生长室,研究了CO2浓度和温度升高对红桦根、茎、叶和枝可溶性蛋白含量和分配的影响,从蛋白水平上来解释川西亚地区的建群种-红桦对CO2升高和温度升高及其交互作用的响应规律,为全球气候变暖川西亚高山的植被保护和恢复提供理论依据。研究结果表明: 1. CO2浓度升高增加了可溶性蛋白的总量,改变了可溶性蛋白分配模式,即,可溶性蛋白分配到根的比例增加,分配到茎、枝、叶的比例减少。可能意味:在CO2浓度升高条件下,红桦根系的生长和营养物质吸收功能将会增强。 2. CO2浓度升高增加了根和茎的清蛋白含量,降低了叶片的清蛋白含量,叶片的球蛋白含量、醇溶蛋白含量和谷蛋白含量均增加。表明CO2浓度升高增加了清蛋白在根中积累,球蛋白、醇溶蛋白和谷蛋白大量在叶片中积累;前人研究所指出的CO2浓度升高使植物叶片可溶性蛋白的含量降低可能仅仅是由于清蛋白含量的降低造成的。 3. 温度升高使红桦幼苗整株所含可溶性蛋白总量增加,但可溶性蛋白总量的分配因红桦幼苗器官的不同而异。温度升高下根、茎、叶和枝的分配量分别占总可溶性蛋白的27.74%、35.57%、23.00%、13.68%,即茎>根>叶>枝。对照的根茎叶枝的分配量分别占总可溶性蛋白的21.01%、41.41%、23.08%、14.50%,即茎>叶>根>枝。表明温度升高使可溶性蛋白分配到根的比例增加,有利于根的可溶性蛋白的积累,增强了根吸收水分和矿质营养的能力,从而有利于根系的生长。 4. 温度升高处理下清蛋白和球蛋白在根中含量升高,在茎、叶和枝中含量下降,但没有达到显著水平;醇溶蛋白在根和叶中含量显著增加;谷蛋白在茎中的含量显著降低。表明温度升高增加清蛋白和球蛋白在红桦幼苗根部的积累,也有利于根和叶醇溶蛋白的积累,但不利于谷蛋白在茎的积累;温度升高条件下叶片可溶性蛋白升高是醇溶蛋白在叶片中积累的结果。 5. CO2浓度和温度同时升高条件下红桦幼苗的可溶性蛋白总量增加很少,只有分配到茎的可溶性蛋白比例增加,并且对可溶性蛋白分配规律没有影响。CO2和温度同时升高下红桦幼苗枝的可溶性蛋白含量的降低是可溶性蛋白总量的降低而不是碳水化合物稀释的结果,并且CO2和温度同时升高对红桦幼苗的生长没有明显的促进作用。 6. CO2和温度同时升高处理对可溶性蛋白含量有显著影响。清蛋白含量在根、茎、叶和枝中均降低,球蛋白含量在根中显著降低,醇溶蛋白含量在根、茎、叶和枝中均降低,谷蛋白含量在根中显著降低。表明CO2浓度和温度同时升高对根的影响显著,即降低了根的可溶性蛋白含量,可能导致根的吸收能力下降。 7. 因此,CO2和温度同时升高对可溶性蛋白影响不能简单地通过CO2和温度单因子影响机理来解释。 It is well known that atmospheric CO2 concentration and temperature are increasing as a consequence of human activities. Atmospheric CO2 concentration are predicted to increase from 350μmol·mol-1 now to 570μmol·mol-1 2030. And temperature will continue to increase for several centuries as a result of CO2 enrichment. Vegetation play a key role in reducing atmospheric CO2 and adapting and controlling warter and energy process in a certain region, while the underlying mechanism are not clear, yet. Betula albo-sinensis, as the dominating tree species of subalpine dark coniferous forest in west Sichuan province, play an important role in determing structure and function of forest ecosystem. In our study, effects of elevated atmospheric CO2 concentration (ambient±350±25μmol·mol-1), increased temperature (ambient±2.0±0.5℃) and their combination on contents and allocation of soluble protein were studied in independent and enclosed-top chamber system under high-frigid conditions. Chambers with ambient CO2 concentration and temperature are taken as control. The results are as the following, 1) Elevated atmospheric CO2 increased the accumulation of total weight of soluble protein in whole plant and changed allocation of soluble protein in red birch by increasing its allocation to roots and reducing its allocation to stem. This caused much more accumulation of soluble protein in roots which might help to prompt growth, development and nutrient absorption ability of roots. 2) Treatment EC increased content of albumin in roots and stems, reduced the content of albumin in leaves, and increased the content of globulin, promalin and glutenin in leaves. That is to say EC increased the accumulation of albumin in roots and accumulation of globulin, promalin and glutenin in leaves. The reduced soluble protein contents in plant leaves by EC, as reported by former researchers, are mainly resulted from the reduced content of albumin in leaves. 3) Elevated temperature increased the total of soluble proteins, but its allocation was dependent on organs. In treatment ET, roots, stems, leaves and branches take 27.74%, 35.57%, 23.00% and 13.68% of total weight of soluble protein. In treatment CK, roots, stems, leaves and branches take 21.01%, 41.41%, 23.08% and 14.50%. Elevated temperature changed allocation of soluble proteins in that it stimulated soluble proteins accumulation in roots and improved the uptake of water in roots. 4) Treatment ET increased the content of albumin and globulin in roots, and reduced the content of albumin and globulin in stems, leaves and branches. The content of promalin in roots and leaves was increased significantly, and the content of glutenin in stems was reduced significant. This suggested that ET stimulated the accumulation of albumin and globulin in roots and accumulation of promalin in leaves and roots; that treatment ET increased content of soluble protein in leaves was mainly resulted from the increased promalin content in leaves. 5) Regarding treatment ETC, the total of weight of soluble proteins increased, but not significantly; but increased in stems. So the combination of elevated atmospheric CO2 and temperature had not changed the allocation of soluble proteins in red birch seedling and reduced soluble proteins in branches were not the result of increased carbohydrate. 6) Treatment ETC reduced the content of albumin and promalin in roots, stems, leaves and branches, reduced the content of globulin and glutenin in roots significantly. That is to say elevated atmospheric CO2 and temperature reduced the content of soluble proteins in roots significantly which might help to prompt growth, development and nutrient absorption ability of roots. 7) The effects of elevated atmospheric CO2 and temperature on soluble protein cannot be simply interpreted through their mechanism that obtained when they were imposed on plant separately.

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大气臭氧的损耗导致了地球表面具有生物学效应的紫外线-B(UV-B)辐射增强。同时,大气成分变化中除了UV-B辐射增强外,氮沉降是一个新近出现而又令人担忧的环境问题,其来源和分布正在迅速扩展到全球范围,并不断向陆地和水生生态系统沉降。本试验在四川省境内的中国科学院茂县生态站内进行,以云山、冷杉、色木槭和红椋子幼苗为模式植物,从生长形态、光合作用、抗氧化能力和矿质营养等方面研究了青藏高原东缘4种树苗对全球变化-增强UV-B辐射和氮供应(氮沉降)的响应。该试验为室外盆栽试验,包括四个处理:(1)大气UV-B辐射+无额外的氮供应(C);(2)大气UV-B辐射+额外的氮供应(N);(3)增强UV-B辐射+无额外的氮供应(UV-B);(4)增强UV-B辐射+额外的氮供应(UV-B+N)。其目的:一方面有助于丰富我国对全球变化及区域响应研究的全面认识,进一步完善在全球气候变化条件下臭氧层削减和氮沉降对陆地生态系统影响的内容;另一方面,在一定程度上有助于我们更好的理解在全球变化下森林更新的早期过程。具体结果如下: 增强的UV-B辐射在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有显著的影响。UV-B辐射增强对幼苗的影响不仅与物种有关,而且,还与氮营养水平相关。总体表现为,高的UV-B辐射导致了色木槭和红椋子幼苗叶片的皱缩和卷曲,并降低了色木槭幼苗的叶片数和叶重,在额外的氮供应下,云杉、冷杉和红椋子的叶重也显著地降低了;色木槭和红椋子幼苗叶片的解剖结构受到了增强的UV-B辐射的影响,增强的UV-B辐射显著地降低了色木槭叶片的栅栏组织厚度,提高了红椋子叶片的厚度;增强的UV-B辐射显著地降低了4种幼苗的单株总生物量、植物地下部分的生长、总叶绿素含量 [Chl (a + b)]、净光合速率和最大量子产量(Fv/Fm),提高了4种幼苗叶片的膜脂过氧化(MDA含量),改变了植物体不同器官中的矿质元素含量;增强的UV-B辐射提高了冷杉、色木槭和红椋子叶片中的过氧化氢含量(H2O2)、超氧负离子(O2-)生成速率,在额外的氮供应下,云杉叶片中的活性氧含量也显著地提高了;在无额外的氮供应条件下,增强的UV-B辐射显著地提高了4种幼苗叶片中的UV-B吸收物质、脯氨酸含量和抗氧化酶的活性(SOD、POD、CAT、GR和APX)。在额外的氮供应条件下,UV-B辐射的增强却显著地降低了冷杉叶片中脯氨酸含量和红椋子叶片中UV-B吸收物质含量,但是,在4种幼苗叶片中,5种抗氧化酶的活性对UV-B辐射的增强没有明显的规律性,增强的UV-B辐射显著地提高了云杉叶片中的POD、SOD和GR的活性,提高了冷杉叶片中的POD和GR活性,提高了色木槭叶片中的POD、SOD和CAT活性和红椋子幼苗叶片中的POD和SOD活性。从这些结果可知,植物在遭受高的UV-B辐射导致的过氧化胁迫时,植物体内形成了一定的保护机制,但是,这种保护不能抵抗高的UV-B辐射对植物的伤害。 额外的氮供应在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有一定的影响,不同幼苗对额外的氮供应响应不同,并且受到UV-B辐射水平的影响。在当地现有的UV-B辐射水平下,额外的氮供应显著地提高了幼苗的单株总生物量、植物地下部分的生长、Chl (a + b)、净光合速率(红椋子除外)、UV-B吸收物质(冷杉除外)、脯氨酸含量(红椋子除外)和部分抗氧化酶的活性,降低了H2O2的含量、O2-的生成速率和MDA含量(红椋子除外),改变了植物体内部分矿质元素含量,显著地提高了云杉和冷杉叶片中的Fv/Fm。这些指标总体表明,在当地现有大气UV-B辐射水平下,额外的氮供应对植物的生长和发育是有利的。在增强的UV-B辐射水平下,4种幼苗的生长形态和光合大部分指标都没有受到额外氮供应的影响,额外的氮供应提高了红椋子幼苗的单株总生物量和Chl (a + b)含量,提高了冷杉和色木槭叶片中的活性氧含量和MDA含量,却降低了红椋子叶片中的活性氧含量;额外的氮供应也提高了云杉、色木槭和红椋子叶片中UV-B吸收物质和脯氨酸含量,降低了冷杉叶片中UV-B吸收物质和脯氨酸含量;在抗氧化酶活性方面,额外的氮供应降低了云杉、冷杉叶片中5种抗氧化酶的活性和红椋子叶片中POD和GR的活性,提高了色木槭叶片中的POD和SOD的活性;4种幼苗植物体内的矿质元素含量对额外的氮供应没有显著的规律性。从这些结果可知,在高的UV-B辐射下,额外的氮供应提高了云杉、冷杉和色木槭幼苗对高的UV-B辐射的敏感性,然而,额外的氮供应却促进了红椋子幼苗的生长,原因可能是,在高的UV-B辐射下,额外的氮供应增加了红椋子叶片的厚度、叶重和叶片数,降低了叶片中活性氧含量的结果。表明在高的UV-B辐射水平下,额外的氮供应降低了红椋子幼苗对高的UV-B辐射的敏感性。 在全球变化的趋势下,UV-B辐射增强和氮沉降可能同时存在,我们的研究表明,与大气UV-B辐射+无额外的氮供应处理相比,增强UV-B辐射+额外的氮供应处理显著地降低了幼苗的单株总生物量(红椋子除外)、Chl (a + b)、净光合速率、Fv/Fm(冷杉除外)和MDA含量(红椋子除外),提高了活性氧含量 (云杉除外)、UV-B紫外吸收物质含量(冷杉除外)、脯氨酸含量和部分抗氧化酶的活性,改变了植物体不同器官中的矿质元素含量。结果表明,在当地现有条件下,全球变化(UV-B辐射增强和氮沉降)对云杉、冷杉和色木槭幼苗的生长是不利,尽管植物体内一些抗氧化性指标提高了,然而,却对红椋子幼苗的单株总生物量的累积没有显著的影响。 The depletion of the ozone led to the increase of ultraviolet-B (UV-B) with biological effects in the earth’s surface. At the same time, except for enhanced UV-B radiation, nitrogen deposition was an anxious environmental problem at present, rapidly expanding to the global scope and continuously depositing to land and aquatic ecosystem. The experiment was conducted in Maoxian Ecological Station of Chinese Academy of Sciences, Sichuan province, China. Picea asperata, Abies faxoniana, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings were selected as model plants to assess the effects of enhanced UV-B radiation and supplemental nitrogen supply on growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings in east Qinghai-Tibetan Plateau. The experiment was potted outdoor, including 4 treatments: (1) ambient UV-B without supplemental nitrogen (control, C); (2) ambient UV-B with supplemental nitrogen (N); (3) enhanced UV-B without supplemental nitrogen (UV-B); (4) enhanced UV-B with supplemental nitrogen (UV-B+N). One hand, it was helpful for enriching our country to comprehensive understanding of the researches in the global change and the region response, further perfecting the effects of the depleted ozone layer and nitrogen deposition on land ecosystem under the global change; the other hand, it was favorable for us to better understanding of the early process of forest renews under the global change. The results were as follows: Enhanced UV-B radiation had significant effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings. The effects of enhanced UV-B on plants were not only related with species, but also related with nitrogen nutrient level. Generally, the increase of UV-B radiation led to the shrinkage and curl of leaves in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, and reduced the number of leaf and leaf weight of Acer mono Maxim seedlings, under supplemental nitrogen supply, leaf weight of Picea asperata, Abies faxoniana and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings significantly also reduced; the anatomical features of leaf in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings affected by enhanced UV-B radiation, the increase of UV-B radiation markedly reduced the palisade tissue thickness of Acer mono Maxim leaf and enhanced the leaf thickness of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings; the enhanced UV-B radiation significantly reduced total biomass per plant of 4 species seedlings, the growth of the underground parts, Chl (a + b), net photosynthetic rate and maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA content) and changed the content of mineral elements in different parts of plants; the enhanced UV-B radiation also increased the rate of superoxide radical (O2-) production and hydrogen peroxide (H2O2) content in leaves of Abies faxoniana, Acer mono Maxim, Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, under supplemental nitrogen supply, the reactive oxygen species in leaves of Picea asperata seedlings also significantly increased by enhanced UV-B radiation; under without supplemental nitrogen supply, enhanced UV-B radiation evidently induced an increase in UV-B absorbing compounds, proline content and the activities of antioxidant enzymes (SOD, POD, CAT, GR and APX) of leaves in 4 species seedlings. Under supplemental nitrogen supply, enhanced UV-B radiation induced a decrease in proline content of leaves in Abies faxoniana seedlings and UV-B absorbing compounds of leaves in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, but, there were no obvious rules in the activities of five antioxidant enzymes of 4 species seedling leaves to enhanced UV-B radiation, enhanced UV-B radiation significantly increased the activities of POD, SOD and GR in Picea asperata leaves, the activities of POD and GR in Abies faxoniana leaves and the activities of POD, SOD and CAT in Acer mono Maxim leaves. The results indicated that some protective mechanism was formed when plants were exposed to enhanced UV-B radiation, but the protection could not counteract the harm of high UV-B radiation on plants. Supplemental nitrogen supply had some effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits. The response of 4 species seedlings was different to supplemental nitrogen supply, and was affected by UV-B levels. Under local ambient UV-B radiation, supplemental nitrogen supply significantly increased the total biomass per plant, the growth of underground parts, Chl (a + b), net photosynthetic rate (except for Acer mono Maxim seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and the activities of some antioxidant enzymes, and reduced H2O2 content, the rate of O2- production and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and changed the content of mineral elemental in different parts; supplemental nitrogen supply also evidently increased Fv/Fm in Picea asperata and Abies faxoniana seedlings. These results indicated that supplemental nitrogen supply was favorable for the growth of plants under local ambient UV-B radiation. Under enhanced UV-B radiation, mostly parameters in growth and morphology of 4 species seedlings were not affected by supplemental nitrogen supply. Supplemental nitrogen supply increased the total biomass per plant and Chl (a + b) of Swida hemsleyi (Schneid. et Wanger.) Sojak seedling, increased the reactive oxygen species and MDA content in Abies faxoniana and Acer mono Maxim leaves, and reduced the reactive oxygen species in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves; supplemental nitrogen supply also increased UV-B absorbing compounds and proline content in Picea asperata, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, decreased UV-B absorbing compounds and proline content in Abies faxoniana leaves; in the activities of antioxidant enzymes, supplemental nitrogen supply significantly reduced the activities of antioxidant enzymes in Picea asperata and Abies faxoniana leaves and the activities of POD and GR in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, and increased the activities of POD and SOD in Acer mono Maxim leaves; the content of mineral elements in 4 species seedlings was no significantly rule to supplemental nitrogen supply. We knew from the results, under enhanced UV-B radiation, supplemental nitrogen supply made Picea asperata, Acer faxoniana and Acer mono Maxim seedlings more sensitivity to enhanced UV-B radiation, however, accelerated the growth of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings. The reason was probably that supplemental nitrogen supply increased the leaf thickness, leaf weight and leaf number, reduced the reactive oxygen content of leaf in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings grown under high UV-B radiation. This showed that supplemental nitrogen supply reduced the sensitivity of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings to high UV-B radiation. Under the tendency of the global change, enhanced UV-B radiation and nitrogen deposition may probably coexist. The results showed, compared with the treatment of ambient UV-B radiation without supplemental nitrogen supply, the treatment of enhanced UV-B radiation with supplemental nitrogen supply significantly reduced the total biomass per plants (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), Chl (a + b), net photosynthetic rate, Fv/Fm and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), and increased reactive oxygen content (except for Picea asperata seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content and part antioxidant enzymes, and changed the content of mineral elements of different parts. The results indicated that the global change (enhanced UV-B and nitrogen deposition) were not favorable for the growth of plants under local ambient UV-B radiation and nitrogen nutrient level,, though increased some antioxidant indexes, however, the treatment of enhanced UV-B with supplement nitrogen supply did not significantly affect on the biomass accumulation of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings.

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大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库,约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦(Betula albosinensis Burk.)是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物,采用人工模拟的方法,研究CO2浓度升高对不同种内竞争强度(种群水平)下红桦幼苗的生理特征、生长、干物质积累及其分配的影响,探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征,为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下: 1)在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后,叶片出现“光合适应”现象。与对照相比,低种植密度(28株m-2)和高种植密度(84株m-2)条件下的红桦幼苗净光合速率(A)、气孔导度(gs)、蒸腾速率(E)、表观量子产量(AQY)和羧化速率(CE)显著降低,而水分利用效率(WUE)则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物(TNC)浓度,结果是红桦幼苗的比叶面积(SLA,cm2 g-1)显著降低。 2)与对照相比,CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率(R GR)显著提高,尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积,结果是单株红桦幼苗的总叶面积比对照显著增加。 3)CO2浓度升高处理显著增加红桦幼苗干物质积累(尤其是细根生物量),改变了红桦幼苗生物量的分配格局。与对照相比,CO2浓度升高处理的红桦幼苗叶重比(LWR)、叶面积比(LAR)、叶根重比(Wl/Wr)和源汇重比(leaf weight to non-leaf weight ratio, Wsource/Wsink)显著下降(高种植密度的LWR除外),而根冠比(R/S)则显著增加。在两种种植密度条件下,CO2浓度升高显著增加红桦幼苗根生物量的分配比率,显著降低叶片的生物量分配比率,对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之,长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降,并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照,而后期迅速下降,说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比,表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration([CO2])has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.

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当前大气CO2浓度升高是全球变化的主要趋势之一,CO2浓度升高还会引起全球变暖等其它环境问题,因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种,在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验,研究了CO2浓度倍增环境下,不同密度水平红桦碳氮固定、分配可能发生的改变,并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应,为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下: 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长,同时也改变生物量在体内的分配格局,主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异,它们对CO2浓度升高的反应程度不同,但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应,也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度,显著增加红桦幼苗的群体生物量,从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度,对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加,但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理,这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明,红桦在升高CO2浓度的条件下,会作出积极的响应,从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高,植株各器官N、P含量降低,但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高,N、P向主茎和根的分配增加,向叶片的分配减少,主要是由于前者在总生物量中的比重增加,而后者减少了。(3) CO2浓度升高,氮磷利用效率(NUE和PUE)提高,氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后,亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加,后期积累速率下降。(3) 升高CO2浓度以后,土壤呼吸显著增强;土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的;所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期,随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势,同时MDA(丙二醛)含量下降,保护酶SOD(超氧化物岐化酶)、CAT(过氧化氢酶)活性升高。由此说明,升高CO2浓度有利于减缓生长季节后期叶片的衰老,使叶片维持较高的光合速率,也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.

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海拔梯度造成的环境异质性,如崎岖的地形、复杂的植被结构以及花期延迟等可能会极大地影响到物种的形态和遗传变异格局。理解物种形态和遗传变异的海拔格局对于物种多样性的管理和保护是非常重要的。尽管植物群体遗传学是一个飞速发展的研究领域,然而与海拔相关的形态变异、遗传变异及群体间遗传差异的研究却很少。到目前为止,还不清楚遗传变异与海拔之间是否必然的相关性。 川滇高山栎是一种重要的生态和经济型树种,广泛分布于中国西南的四川、西藏、贵州和云南省的高海拔地区,在保持水土、调节气候方面起着十分重要的作用。尽管主要受阳光限制而仅分布于阳坡,但其海拔梯度范围较大,表明川滇高山栎对不同的环境具有很强的适应性。本文通过叶型及生理响应、微卫星分子标记和扩增性片段长度多态性方法,试图探索川滇高山栎叶沿海拔梯度的形态和生理响应及其沿海拔梯度的遗传变异格局,为川滇高山栎的保护和利用提供进一步的遗传学理论依据和技术指导。 对叶形、含氮量及碳同位素的试验结果表明,平均比叶面积、气孔密度、气孔长度和气孔指数等气孔参数随海拔的升高呈非线性变化。在海拔大于2800 m时,川滇高山栎的比叶面积、气孔长度和气孔指数都随海拔升高而降低,但是在海拔小于2800 m时,这些指标都随海拔的升高而增大。相对而言,单位叶面积的含氮量和碳同位素则表现出相反的变化模式。另外,比叶面积是决定碳同位素沿海拔梯度变化的最重要参数。本研究结果表明,海拔2800 m附近是川滇高山栎生长和发育的最适地带,在这里生长的植物叶片厚度更薄、气孔更大、叶碳同位素值更小。 利用六对微卫星引物对五个不同海拔川滇高山栎群体遗传多样性进行研究,结果表明,群体内表现出较高的遗传多样性,平均每位点等位基因数11.33个,平均期望杂合度达0.820。群体间差异较小,分化仅为6.6%。聚类分析也并没有显示出明显的海拔格局。然而低频率等位基因却与海拔呈显著性正相关(R2=0.97, P < 0.01),表明在高海拔处,川滇高山栎以更多的稀有基因来适应恶劣的环境条件。本试验结果表明由海拔梯度形成的选择性压力对川滇高山栎群体的遗传变异影响并不明显。 为了进一步探讨川滇高山栎群体遗传变异与海拔之间的相互关系,我们还对其进行了扩增性片段长度多态性分析。结果表明:(1)随海拔的升高(从群体WL2到群体WL5),群体内遗传变异降低,而群体间遗传差异增加;(2)低海拔群体WL1表现出最低的遗传变异性(HE = 0.181),同时与其余四个群体间呈现出最大的遗传差异性(平均FST = 0.0596);(3)在除去低海拔群体WL1后,Mantel检测表明群体间遗传距离与海拔距离之间表现出正相关性。另外,研究结果还表明,遗传变异受生境条件(过度的湿热环境)及人为干扰(火烧、砍伐和放牧)的影响,这一点至少在低海拔群体WL1上发生了作用。 通过叶形态、生理及DNA分子水平的研究,结果表明叶形态特征和碳同位素与海拔紧密相关,与海拔之间呈非线性变化,海拔2,800 m附近是川滇高山栎生长和发育的最适地带。海拔梯度在一定程度上会影响到川滇高山栎群体的遗传变异结构,但在这样一个狭窄的地理分布区域里,这种影响并不足以导致群体间较大的遗传分化。同时生境条件及人为干扰也是影响遗传变异的限制性因子,不容忽视。 Altitudinal gradients impose heterogeneous environmental conditions, such as rugged topography, a complex pattern of vegetation and flowering delay, and they likely furthermore markedly affect the morphological and genetic variation pattern of a species. Understanding altitudinal pattern of morphological and genetic variation at a species is important for the management and conservation of species diversity. Although plant population genetics is a fast growing field of research, there are only few recent investigations, which analyzed the genetic differentiation and changes of intra-population variation along altitudinal gradients. At present, it is still unclear whether there are some common patterns of morphological and genetic variation with altitude. Quercus aquifolioides Rehder & E.H. Wilson, which is an important ecological and economical endemic woody plant species, is widely distributed in the Yunnan and Sichuan provinces, Southwest China. Its large range of habitat across different altitudes implies strong adaptation to different environments, although it is mainly restricted to sunny, south facing slopes. It plays a very important role in preventing soil erosion, soil water loss and regulating climate, as well as in retaining ecological stability. In this paper, we tried to understand the altitudinal pattern of morphological and genetic variation along altitudinal gradients through the experiments of leaf morphological and physiological responses, microsatellite analysis and AFLP markers. In leaf morphological and physiological responses experiment, we measured leaf morphology, nitrogen content and carbon isotope composition (as an indicator of water use efficiency) of Q. aquifolioides along an altitudinal gradient. We found that these leaf morphological and physiological responses to altitudinal gradients were non-linear with increasing altitude. Specific leaf area, stomatal length and index increased with increasing altitude below 2,800 m, but decreased with increasing altitude above 2,800 m. In contrast, leaf nitrogen content per unit area and carbon isotope composition showed opposite change patterns. Specific leaf area seemed to be the most important parameter that determined the carbon isotope composition along the altitudinal gradient. Our results suggest that near 2,800 m in altitude could be the optimum zone for growth and development of Q. aquifolioides, and highlight the importance of the influence of altitude in research on plant physiological ecology. Genetic variation and differentiation were investigated among five natural populations of Q. aquifolioides occurring along an altitudinal gradient that varied from 2,000 to 3,600 m above sea level in the Wolong Natural Reserve of China, by analyzing variation at six microsatellite loci. The results showed that the populations were characterized by relatively high intra-population variation with the average number of alleles equaling 11.33 per locus and the average expected heterozygosity (HE) being 0.779. The amount of genetic variation varied only little among populations, which suggests that the influence of altitude factors on microsatellite variation is limited. However, there is a significantly positive correlation between altitude and the number of low-frequency alleles (R2=0.97, P < 0.01), which indicates that Q. aquifolioides from high altitudes has more unique variation, possibly enabling adaptation to severe conditions. F statistics showed the presence of a slight deficiency of heterozygosity (FIS=0.136) and a low level of differentiation among populations (FST=0.066). The result of the cluster analysis demonstrates that the grouping of populations does not correspond to the altitude of the populations. Based on the available data, it is likely that the selective forces related to altitude are not strong enough to significantly differentiate the populations of Q. aquifolioides in terms of microsatellite variation. To further elucidate genetic variation pattern of Q. aquifolioides populations under sub-alpine environments, genetic variation and differentiation were investigated along altitudinal gradients using AFLP markers. The altitudinal populations with an average altitude interval of 400 m, i.e. WL1, WL2, WL3, WL4 and WL5, correspond to the altitudes 2,000, 2,400, 2,800, 3,200 and 3,600 m, respectively. Our results were as follows: (i) decreasing genetic variation (ranging from 0.253 to 0.210) and increasing genetic differentiation with altitude were obtained from the WL2 to the WL5 population; (ii) the WL1 population showed the lowest genetic variation (HE = 0.181) and the highest genetic differentiation (average FST = 0.0596) with the other four populations; (iii) the positive correlation was obtained using Mantel tests between genetic and altitude distances except for the WL1 population. Our results suggest that altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides populations to some extent. In addition, habitat environments (unfavorable wet and hot conditions) and human disturbances (burning, grazing and felling) were possible influencing factors, especially to the low-altitude WL1 population. The present study shows that there were close correlations between morphological features and carbon isotope composition in our data. This indicates that a coordinated plant response modified these parameters simultaneously across different altitudes. Around 2,800 m altitude there seems to be an optimum zone for growth and development of Q. aquifolioides, as indicated by thinner leaves, larger stomata and more negative d13C values. All available evidence indicates altitudinal gradients may have influenced the genetic variation pattern of Q. aquifolioides to some extent. Decreasing genetic variation and increasing genetic differentiation with altitude was obtained except for the WL1 population. And the environment of habitats and human disturbances were also contributing factors, which impact genetic variation pattern, especially to the low-altitude WL1 population.

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在青藏高原东部的亚高山针叶林区,如何尽快恢复这一生态脆弱地区的植被,改变生态环境恶化的趋势,是一个十分重要的课题。光一直被认为是植物种间相互替代,尤其是森林演替过程中植物相互替代或植被恢复中的关键环境要素之一。植物能否适应林冠下或林窗中异质的、或多变的光照条件,对其在林中的生存、分布、更新以及森林动态都是非常重要的。 本文以青藏高原东部亚高山针叶林的主要森林类型——岷江冷杉林群落的几种树苗为研究对象,采用实验生态学、生理及生物化学等方法,通过模拟针叶林不同大小林窗内光照强度的变化,在中国科学院茂县生态站内采用遮荫处理设置6个光照梯度(100、55、40、25、15与7%全光照),来研究具有不同喜光特性的植物对光强的响应与适应机制,其研究结果可为揭示亚高山针叶林的演替规律、以及人工林下幼苗的存活与定居提供科学依据,也能为苗木的生产与管理提供科学指导,尤其是对针阔树种在不同光强下的响应与适应的比较研究,能为如何将阔叶树种整合到人工针叶林中提供新的思路。 光强对植物生长的影响 光强对植物的生长具有重要作用,不同植物在各自适宜的光强梯度下才能生长良好。通过一个野外盆栽实验,来研究不同光强对植物生长的影响(第三章)。主要研究结果如下,低光强下植物株高/茎生物量增加,说明植物会将生物量更多用于高生长,以便有效地拦截光资源;在强光下,植物将生物量更多地向根部分配,使得植物在强光下能够吸收更多的水分,而避免干旱胁迫。 在第一个生长季节,以相对生长速率(RGR)表示,红桦和青榨槭在100%全光照下RGR最大,粗枝云杉在55%最大,岷江冷杉在25-40%下较好;然而,在第二个生长季节,2种阔叶树的相对生长速率(RGR)的适宜光强则变为25-55%,云杉为55-100%,而冷杉为25-100%。可见,从第一年到第二年,2种阔叶树苗更适宜在部分荫蔽的条件下生长;而2种针叶树苗对光的需求则逐渐增加,这可能是增加对根生物量相对投资的结果,因为以这种方式,强光下生长的针叶树幼苗更能保持其内部水分平衡,其生长不会因干旱胁迫而受到严重影响。另外,严重遮荫会引起冷杉幼苗死亡。 植物对光强的生理适应 植物可以通过自身形态和生理特征的调整,来发展不同的光能利用策略从而能够在林中共存。通过一个野外盆栽实验,研究了不同光强下生长的几种树苗的生理特征(第四章)对不同光强的响应与适应。结果显示:强光下,粗枝云杉和红桦的光合能力增加,而岷江冷杉和青榨槭在中度遮荫(25-55%)的条件下光合能力最大。植物叶氮和叶绿素含量增高,而光补偿点和暗呼吸速率降低,这些都是植物对低光环境的适应性反应;而强光下植物叶片和栅栏组织变厚,是对强光的一种保护性反应。 植物对光的可塑性反应 不同植物会表现出对光适应有利的生理和形态可塑性反应。本文对第三章、第四章的实验数据进行可塑性指数分析,来研究植物对光强的表型可塑性反应(第五章)。结果显示,生理特征调整是植物对不同光环境的主要适应途径。红桦和青榨槭的可塑性指数平均值要大于粗枝云杉和岷江冷杉,充分表明这2种阔叶树在生理和形态上较强的可塑性更有利于对光环境的适应,而具有比耐荫树种更强的适应能力。另外,2种针叶树相比,云杉的适应性更强。本研究结果支持树种的生理生态特性决定了其演替状况和生境选择的假说。 植物的光抑制与防御 当植物叶片吸收了过多光能,会发生光抑制现象。植物对光抑制的敏感性及防御能力对其生长具有重要意义。本文通过两个野外盆栽实验,研究了生长在强光下(第六章)和变化光强下(第八章)植物的光抑制现象及其防御策略。结果表明,在强光下或从遮荫状态转入强光下,植物都会发生光抑制,其对光抑制的敏感性与植物的耐荫性(或喜光)和演替状态有密切联系。长期生长在强光下的植物受到光抑制是可恢复的,而当处于荫蔽环境的植物突然暴露于强光下时,受到的光抑制不能完全恢复,可能是(部分)光合机构受到破坏的缘故。粗枝云杉和青榨槭防御光抑制伤害的能力较强,热耗散是其防御光抑制的主要途径。长期的强光作用能使岷江冷杉和红桦发生严重光抑制,甚至光伤害,而红桦能够通过“凋落老叶,萌发新叶”的途径来适应新的强光环境。 How to restore the vegetation of subalpine coniferous forest in eastern Qinghai-Tibet Plateau, and change the trend of ecological deterioration is a very important issue. Acclimation of tree seedlings to different and varing light environment affects to a great extent the successful regeneration and establishment of subalpine coniferous forests in southwestern China’s montane forest areas, because the ability to respond to such changing resource are commonly assumed to be critical to plant success, and have a growth advantage than others. In this paper, several species seedlings in Abies faxoniana community were chosed to study the response and adaptation to light intensity and the interspecific differences of adaptability in six shaded sheds (100, 55, 40, 25, 15 and 7% of full sunlight) in the Maoxian Ecological Station of Chinese Academy of Sciences. Our results could provide a strong theoretical evidence for understanding the forest succession laws of subalpine coniferous forests, and the survival and settlement of seedlings under plantations, and provide scientific direction for the production and management of seedlings, especially the comparative studies of the acclimation to light between the conifer and broadleaf trees could provide new ideas for how to integrate the broad-leaved trees into the artificial coniferous forest. Growth under different light intensity Light intensity plays an important role on plant growth. One field experiments was conducted to study the growth of tree seedlings of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii under different light intensities. The results showed that plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources, while biomass greater allocation to the roots, could make plants under high light environment absorb more water, and avoid drought stress. During the first growing season, the relative growth rates (RGRs) of Betula albo-sinensis and Acer davidii had the greatest values under the 100% of full light, for 55% of Picea asperata, and for 25-40% of Abies faxoniana. However, in the second growing season the the relative growth rates of the two broad-leaved trees changed and were appropriate for 25-55% of full light, for 55-100% of spruce, and for 25-100% of fir. Thus, from the first year to the second year, two broad-leaved seedlings maybe more suitable to partly shading environment, and two coniferous seedlings would have an increase in light demand, which may be an increased root biomass investment. Because in this way, seedlings grown under high light could better maintain their internal water balance, and thus its growth would not be seriously affected by drought stress. In addition, serious shading would cause fir seedlings to die. Acclimation of physiology to light Plants could coexist in forest ecosystem by forming different strategies of light use. One field experiments was conducted to study the acclimation of tree seedlings to different light intensity of Picea asperata, Abies faxoniana, Betula albo-sinensis and Acer davidii. The results showed that the photosynthetic capacity of Picea asperata and Betula albo-sinensis exhibited a general tendency of increase with more light availability; but for Abies faxoniana and Acer davidii seedlings, their highest values of the same parameters were found under intermediate light regime (i.e. 25-55% of PFD relative to full sunlight). Plants under low light environment could increase the specific stem length (stem length/ stem dry mass), in order to effectively intercept light resources. Leaf nitrogen and chlorophyll content increased, while dark respiration rate and light compensation points decreased, all of which were adaptive response to the low light environment. On the contrary, plants under high light environment had the thicken leaves and palisade tissue, which was a protective response to high light. Phenotypic plasticity to light Phenotypic plasticity can be exhibited in morphological and physiological processes. Physiological characteristical adjustment is the main for plant adaptation to different light environment.The means of plasticity indexes for Betula albo-sinensis and Acer davidii seelings were greater than Picea asperata and Abies faxoniana, amplied that the two broad-leaved trees were much more adaptable to the environment. In addition, spruce had the higher adaptablity than fir. The findings supported the hypothesis that the ecological characteristics of the species determined the biological status and its biological habitat selection. Photoinhibition and photoprotection to light Compared with conifer, broad-leaved trees could better change leaf morphology and adjust biomass allocation to adapt to changing light environment. However, excess light can photoinhibit photosynthesis and may lead to photooxidative destruction of the photosynthetic appatus. Two field experiments were conducted to study the photoinhibition of photosynthesis. The results showed that when plants grown under high light environment or plants transferred from low to high irradiance, the four tree seedlings would undergo a period of photoinhibition. In four species, photoinhibited leaves could recover to initial photosynthetic rates when they were long-term planted under high light environment. However, when plants were suddenly exposed to high irradiance, this photoinhibition could not be reversible, may be the photosynthesis apparatus were (or partly) photooxidatively destructed.