56 resultados para Faba bean leaves and seeds


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以冬小麦品种长武134(抗旱性强)和陕253(抗旱性弱)为材料,研究干旱对旗叶净光合速率和叶绿素含量、主茎及其组成节间花前积累干物质的转运及其对穗粒重贡献的影响。结果表明,干旱条件下长武134主茎穗粒重降幅小于陕253;干旱缩短了小麦花后旗叶光合速率高值持续期(PAD)和叶绿素含量缓降期(RSP),长武134受影响程度较小。长武134除穗下节外,其余各节间及茎杆干物质转运量及对籽粒贡献率降幅均大于陕253;干旱提高了陕253穗下节和倒二节干物质转运率,降低了倒三节和下部节干物质转运率,茎杆干物质转运率无明显变化;长武134除穗下节外其余各节间及茎杆干物质转运率均明显低于对照,而且距离穗部越远的节位降低幅度越大。上述结果表明,干旱条件下不同节位干物质转运能力变化与距离穗远近有关,花前茎杆干物质转运并不能补偿籽粒产量的损失,花后旗叶光合功能期延长是小麦抗旱高产的主要原因。

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大田条件下设置不同的施氮和灌水水平,研究施氮量和灌水量对冬小麦(Triticum aestivum L.)叶片光合作用生理及籽粒形成的影响。结果表明,施氮和灌水提高叶片Fv/Fm和Pn,并显著提高冬小麦籽粒产量、单位面积穗数和每穗粒数。施氮明显降低千粒重,而适量灌水明显提高千粒重。

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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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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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岷江上游地区高山/亚高山植被分布的坡向性分异显著,阴阳坡高山林线不仅物种组成差异明显,并且分布海拔呈现出阴坡高阳坡低的格局.阳坡林线树种主要是圆柏属乔木,林线类型多为渐变型,海拔高度大约在3 400m~3 800m;阴坡林线树种主要是冷杉,林线类型多为骤变型,海拔高度约在3 800m~4 400m.本研究采用土壤种子库物理筛选、室内萌发实验及野外群落调查等方法,对岷江上游地区阴坡岷江冷杉和阳坡祁连圆柏两类林线树种不同海拔梯度上土壤种子库以及幼苗库特征进行了调查,从土壤种子库和幼苗更新特征的角度对林线乔木树种种群更新特征进行了分析,进而对该地区高山林线在阴阳坡分布差异的原因进行了探讨,结果显示: 1.土壤种子库 阴坡:阴坡高山林线附近岷江冷杉土壤种子的平均密度大约为50.96粒/m2,其中树线以上10m处土壤种子密度为1.00粒/m2,树线处大约19.33粒/m2,林线交错带内土壤种子密度最高为136.83粒/m2,郁闭林内种子密度小于林线交错带,只有30.50粒/m2,种子平均空壳率为52%,霉变率达34%,完好种子只有6%.土壤种子库垂直分布特征为地被物层含种子比重最大,大约在67.50%左右;其次为0~2cm层,约18.84%左右;2~5cm层所占种子比例最小,约13.66%左右.霉变种子数量与土壤深度呈负相关. 阳坡:阳坡祁连圆柏土壤种子的平均密度为60.16粒/m2.树线以上10m处密度为1.92粒/m2,树线位置大约108.16粒/m2,林线交错带内平均为75.80粒/m2,郁闭林内种子密度小于林线交错带,只有20.00粒/m2.种子平均空壳率为36%,完好种子占49%,霉变率较低,大约为10%.阴阳坡林线树种土壤种子库垂直分布特征为:地被物层含种子最多,其次为0~4cm层,4~10cm层所占种子比例最小,霉变种子数量与土壤深度也呈负相关. 2. 幼苗库调查 阳坡:在树线以上区域没有发现幼苗,林线交错带内幼苗密度平均达3 250株/hm2,郁闭林内仅2 750株/ hm2.整个样地内1~2a幼苗很少甚至没有出现,3~10a的幼苗相对较多.空间分布上,祁连圆柏幼苗在林线交错带内接近随机分布,郁闭林内则介于随机分布和均匀分布之间. 阴坡:在树线以上幼苗密度为1 250株/ hm2,全部为1~2a幼苗,林线交错带内幼苗密度平均达7 000株/ hm2,郁闭林内达6 250株/ hm2.林线附近岷江冷杉幼苗丰富度以及幼苗的出现频率明显高于祁连圆柏,年龄结构也较祁连圆柏完整.岷江冷杉幼苗空间分布除了树线处幼苗的分布为随机分布,其他海拔则为集群分布. 3.从不同土壤深度的种子总量和幼苗数量的相关性检验发现,当年生幼苗数量跟表层种子总量相关性极显著, 但是两年生幼苗的数量与底层种子数量相关性显著.土壤种子在土壤中的垂直分布格局从一定程度上可以反映种子库的年际特征.岷江冷杉土壤种子库较丰富,种子散布后的存活力随着时间的变化逐渐下降,属于季节性瞬时种子库;祁连圆柏土壤种子散布格局为集群型分布,成熟种子大部分散布在母株冠幅内,属于永久性土壤种子库. 4.在阴坡林线交错带及以上区域还存在较为丰富的乔木土壤种子,并且在树线以上区域还发现了少量的岷江冷杉幼苗.从样地乔木的年龄结构发现,在林线交错带内上部到树线位置主要以幼龄林为主,且年龄结构完整,基本符合入侵性林线特征;阳坡林线交错带内幼苗出现频率很低,树线以上区域虽然存在种子库,但是没有幼苗出现,在林线交错带内乔木径级差距很大,年龄结构异常不完整,这种特征的林线将会面临两个可能结果:一种是维持现有状态,保持平衡;另外一种就是退化,但阳坡林线的实际动态趋势还有待长期定点研究. Treelines on the upper region of Minjiang River differ between the north aspect and the south aspect in their appearances, altitudes and tree species. On the north aspect, trees of Abies form a sharp and abrupt treeline ranging from 3800m to 4400m, while on the south the treeline is generally lower(3 400~3 800m), more open and gradual and mostly composed of Sabina. In this study, we examined the altitudinal gradients of soil seed banks and seedling recruitments at the treeline ecotones of a N-aspect and a S-aspect by using soil sieving, germination experiment and field investigations, analyzed the characteristics of population regeneration of tree species at the transitional zone and presented a analysis of the causes to the aspect-related difference in treeline patterns in the study area. Major results of our study include: 1. Soil seed bank N-aspect: Of the 50 plots investigated, the average density of soil seeds is 50.96/m2, in which well-formed seeds account for 6%, empty seeds 52%, parasitized seeds34%, and seeds damaged by animals 8%. The size of soil seed bank varies along altitude, being 1.00 seeds /m2 at the 10m above the treeline and ca.19.33 seeds/m2 at the upper limit of treeline. The highest density (136.83 seeds/m2) occurs at the treeline ecotone. By contrast, the density of soil seed for the closed forest is only 30.50 seeds/m2. In terms of vertical strata, 67.50% of the total seeds are at the surface layer, 18.84% at the middle layer (0~2cm) and 13.66% at deeper layer (2~5cm). The number of parasitized seeds is negatively correlated to soil depth. S-aspect: Of the 50 plots investigated, the average density of soil seeds is 60.16 seeds/m2, and the well-formed seeds account for 49%, empty seeds 36%, parasitized seeds10%, and seeds damaged by animals 1%. The size of soil seed bank varies along altitude, with 1.92 seeds/m2 recorded at the10m above the treeline,108.16 seeds/m2 at the upper limit of treeline, and 75.80 seeds/m2 at the treeline ecotone, while that for the closed forest is 20.00 seeds/m2. The number of seeds decreases with the depth of soil. As is on the N-aspect, the size of soil bank, from large to small, follows the order of the surface layer, the middle layer (0~4cm) and the bottom layer (4~10cm). The number of parasitized seeds is also negatively correlated to the depth of the soil. 2. Seedling bank N-aspect: A mean maximum seedling abundance of 31 000 seedlings/hm2 was recorded near alpine treeline at growing season. The density of seedlings is 1 250 seedlings/ha (all being 1 or 2 years old) at the alpine meadow 10m away above treeline, 7 000 seedlings/ha at treeline ecotone and 6 250 seedlings/ha for closed forest.The spatial distribution of Abies faxoniana seedlings is random at the upper limit of the treeline but clumped at other altitudes. S-aspect: No seedlings were found at the alpine meadow 10m away from the treeline. The density of seedlings was 3 250 seedlings/ha at treeline ecotone and 2 750 seedlings/ha for the closed forest.Hardly any 1 year current and 2 year-old seedlings appeared at the plots. The spatial distribution of Sabina przewalskii seedlings is random at treeline ecotone and between “random” and “even” forest closed forest. 3.Correlation tests of seedling population and seed bank at different soil layers indicated that the emergents were strongly correlated to seed bank at surface layer while the number of two-year seedlings was significantly correlated to the seed bank at the bottom of soil layer, indicating that germination mainly occurs at the soil surface while the middle or bottom layer was the reserve for non-germination or dead seeds. It can thus be postulated that Abies faxoniana soil seed bank is of seasonal transient type. By contrast, the soil seed bank of Sabina przewalskii is of persistent type and the soil seeds and seedlings of this species occurred more frequently near the islands of adult trees. 4.A good many soil seeds of both tree species were found near the treeline ecotone and above at N- and S-aspects. A few young seedlings were found above the Abies treeline. Investigation of five altitudinal transects respectively on N- and S-aspects indicated that Abies faxoniana has a more complete age structure than the stands of Sabina przewalskii. The age of firs decreased from closed forest to the upper limit of treeline, which suggests that the Abies treeline is advancing to higher altitude. While on the south aspect, only few Sabina przewalskii soil seeds and nearly no seedlings were found above the treeline ecotone. The stands exhibit extremely great difference in diameter classes with significantly incomplete age structure. This would lead to two possible results for the treelines: maintaining an equilibrium state at the current position or degenerating. But more studies should be carried out at longer time scales or larger spatial scales to understand whether the Sabina treeline is degenerating.

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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.

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土壤是人类赖以生存的自然环境和农业生产的重要资源,世界面临的粮食、资源和环境问题与土壤密切相关,目前危害土壤的主要因素是干旱和重金属污染。杨树具有适应性强、生长快和丰产等特性,本论文以青杨组杨树为模式植物,采用植物生态、生理及生物化学等方法,研究杨树对土壤干旱和锰胁迫的生态生理反应以及种群间差异,研究成果可为我国干旱半干旱地区营造人工林、防止沙漠化提供理论依据,也为恢复与重建重金属污染地区退化生态系统提供科学指导。主要研究结果如下: 1. 青海杨不同种群对干旱胁迫的响应差异 干旱胁迫显著降低了两个青海杨种群(干旱种群和湿润种群)生物量积累,包括株高、基径、干物质积累等,通过植物结构的调整,有更多的生物量向根部分配。干旱胁迫还显著降低了叶绿素和类胡萝卜素含量,增加了游离脯氨酸和总氨基酸含量。另一方面,干旱胁迫诱导了活性氧的积累,作为第二信使,激活了抗氧化系统,包括抗坏血酸(ASA)含量和酶系统如超氧化物歧化酶(SOD),愈创木酚过氧化物酶(GPX),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)。这样,杨树既有避旱机制又有耐旱机制,使其在干旱胁迫下有相当程度的可塑性。与湿润种群相比,干旱种群杨树有更多的生物量分配到根部,积累了更多的游离脯氨酸和总氨基酸来进行渗透调节,并且有更有效的抗氧化系统,包括更高含量的ASA 和更高活性的APX 和GR,这些使得干旱种群杨树比湿润种群杨树对干旱有更好的耐性。 2. 喷施硝普钠(SNP)对青海杨阿坝种群干旱胁迫耐性的影响 干旱胁迫显著的降低了青海杨阿坝种群的生长和生物量积累以及叶片相对含水量,还诱导了脯氨酸的合成以进行渗透调节。干旱胁迫下过氧化氢(H2O2)显著累积从而造成对膜脂和蛋白的伤害,使得丙二醛和蛋白羰基含量升高。干旱胁迫下喷施SNP可以减轻干旱胁迫造成的伤害,包括增加叶片的相对含水量,增加脯氨酸和总氨基酸的积累,并激活抗氧化酶系统如SOD,GPX和APX,从而减少丙二醛(MDA)和蛋白羰基(C=O)的积累,但是在水分良好情况下SNP的效果不显著。 3. 青杨不同种群对锰胁迫的生长与形态响应差异 在同一锰浓度下,干旱种群的耐性指数都要高于湿润种群,这表明青杨对干旱和高锰胁迫具有交叉耐性。两个种群的株高,生物量和叶绿素含量都随锰浓度的升高而逐渐下降。就累积浓度而言,0 和0.1 mM 锰胁迫下,干旱种群积累的锰浓度要高于湿润种群,而在高浓度锰胁迫下(0.5 和1 mM),湿润种群要高于干旱种群。在0,0.1 和0.5 mM下,锰大多积累在根中,叶片次之,茎中最少。而在1 mM,锰更多的积累在叶片中。就累积总量而言,在各个锰浓度胁迫下,根,茎和叶相比,两个种群青杨都是叶片累积的锰总量要高于根和茎。两个种群间比较,对照中没有显著区别,0.1 mM 锰胁迫下,湿润种群根中累积的锰要高于干旱种群,而在地上部中,干旱种群要高于湿润种群。而0.5 和1 mM 锰胁迫下,根、叶、茎+叶、根+茎+叶中,锰累积总量都是湿润种群高于干旱种群。锰胁迫下,青杨叶片数和叶面积包括总叶面积和平均叶面积都显著降低。叶片横切面的光学显微观察结果表明未经锰胁迫的栅栏组织的细胞饱满,海绵组织发达、清晰;胁迫后杨树叶片栅栏组织细胞出现不同程度的皱缩,海绵组织几乎不可见,此外还发现输导组织在胁迫下密度变小和分生组织严重割裂等现象。 4. 青杨不同种群对锰胁迫的生理与生化响应差异 青杨两个种群脱落酸(ABA)含量在锰胁迫下都显著增加,干旱种群的增幅更大。三种多胺含量在锰胁迫下显示了不同的响应趋势:腐胺在两个种群的各个锰处理下都增加,亚精胺只在干旱种群中显著增加,而精胺除了干旱种群在1 mM 下有所增加外,在锰胁迫下变化很小。谷胱甘肽含量随锰浓度升高而增加,在0.5 mM 锰时达到最高值,1mM 时有所下降。植物络合素(PCs)含量与非蛋白巯基(NP-SH)趋势相似,随锰浓度的升高而增加,且干旱种群中含量要高于湿润种群。锰处理还引起氧化胁迫,表现为过氧化氢和丙二醛含量增加。SOD 活性在湿润种群中,在0 到0.5 mM 锰胁迫下活性升高,但在1 mM 锰胁迫时,其活性有所下降。而在干旱种群中,SOD 活性变化较小,并始终维持在一个较高的水平。APX 活性在两个种群中都随锰浓度的升高而增加,干旱种群活性要高于湿润种群。锰胁迫还显著增加了酚类物质的含量,同时GPX 和多酚氧化酶(PPO)活性也随锰浓度的升高而增加。干旱种群的酚类含量和GPX 与PPO 活性都要高于湿润种群。锰胁迫还改变了氨基酸的含量和构成,根据锰胁迫下浓度变化的不同,可以将游离氨基酸分为三组:第一组包括,谷氨酸,丙氨酸和天门冬氨酸,这一组氨基酸含量在锰胁迫下有所下降。第二组包括缬氨酸,亮氨酸和苏氨酸含量在锰胁迫下基本不变化或变化很小。剩下的氨基酸为第三组,这组氨基酸含量在锰胁迫下显著增加,而根据增加的幅度又可以将它们分为两个亚组,丝氨酸,酪氨酸,苯丙氨酸,组氨酸和脯氨酸,在1 mM 下的含量是对照的4 倍以上。异亮氨酸,赖氨酸,精氨酸和甘氨酸含量在1 mM 下是对照含量的2 倍以下。同时,同一锰浓度下,干旱种群比湿润种群积累的氨基酸含量要高。 Soil is the indispensable environment for human survival and important resource foragriculture development. Food and environmental problems facing the world are all closelyrelated to soil and nowadays it is threatened by many factors, among which drought stress andheavy metal pollution are the most serious ones. Poplars (Populus spp.) are importantcomponents of ecosystem and suitable as a source of fuel, fiber and lumber due to their fastgrowth. In this study, different populations of Section Tacamahaca spach were used as modelplants to investigate the adaptability to drought stress and manganese toxicity and differencesbetween populations from dry and wet climate regions. Our results can provide theoreticalevidence for the afforestation and prevention of desertification in the arid and semi-arid areas,and also can supply scientific direction for the reconstruction and rehalibitation of ecosystemscontaminated by heavy metals. The results are as follows: 1. Differences in ecophysiological responses to drought stress in two contrastingpopulations of Populus przewalskii Drought stress not only significantly affected dry mass accumulation and allocation, butalso significantly decreased chlorophyll pigment contents and accumulated free proline andtotal amino acids. On the other hand, drought also significantly increased the levels ofabscisic acid and reactive oxygen species, as secondary messengers, to induce the entire set ofantioxidative systems including the increase of reduced ascorbic acid content and the activities of superoxide dismutase, guaiacol peroxidase, ascorbate peroxidase and glutathioneredutase. Thus the combination of drought avoidance and tolerance mechanisms conferredpoplar a high degree of plasticity in response to drought stress. Compared with the wetclimate population, the dry climate population showed lower dry matter accumulation andallocated more biomass to root systems, and accumulated more free proline and total aminoacids for osmotic adjustment. The dry climate population also showed more efficientantioxidant systems with higher content of ascorbic acid and higher activities of ascorbateperoxidase and glutathione redutase than the wet climate population. All these made the dryclimate population superior in adaptation to drought stress than the wet climate population. 2. Effect of exogenous applied SNP on drought tolerance in Populus przewalskii Drought stress significantly increased hydrogen peroxide content and caused oxidativestress to lipids and proteins assessed by the increase in malondialdehyde and total carbonylcontents, respectively. The cuttings of P. przewalskii accumulated proline and other aminoacids for osmotic adjustment to lower water potential, and activated the antioxidant enzymes such as superoxide dismutase, guaiacol peroxidase and ascorbate peroxidase to maintain thebalance of generation and quenching of reactive oxygen species. Moreover, exogenous SNPapplication significantly heightened the growth performance of P. przewalskii cuttings underdrought treatment by promotion of proline accumulation and activation of antioxidant enzymeactivities, while under well-watered treatment the effect of SNP application was very little. 3. Morphological responses to manganese toxicity in the two contrasting populations ofPopulus cathayana High concentration of manganese caused significant decrease in shoot height andbiomass accumulation. The tolerance index of the dry climate population was significantlyhigher than that of the wet climate population, suggesting the superior Mn tolerance in theformer and the existence of cross-tolerance of drought stress and high Mn toxicity. Injuries tothe leaf anatomical features were also found as the reduced thickness in palisade and spongyparenchyma, the decreased density in the conducting tissue and the collapse and split in themeristematic tissue in the central vein. As for the Mn concentrations in the plant tissues, under0, 0.1 and 0.5 mM, most of the Mn accumulated in the roots, then leaves, and stem the least, while under 1 mM, most of the Mn accumulated in the leaves. As far as the total amounts ofMn extraction are concerned, the leaf extracted more Mn than the root and stem in the twopopulations under various Mn concentrations. There is no difference between the twopopulations under control. Under 0.1 mM, the wet climate population extracted higher Mn inthe root than the dry climate population, while in the shoot, the dry climate populationextracted much more Mn. Under 0.5 and 1 mM, the wet climate population translocated moreMn both in the root and the shoot than the dry climate population. 4. Physiological and biochemical responses to manganese toxicity in the two contrastingpopulations of Populus cathayana Mn treatment resulted in oxidative stress indicated by the oxidation to lipids, proteinsand DNA. A regulated network of defence strategies was employed for the chelation,detoxification and tolerance of Mn including the enhanced synthesis of ABA and polyamines,the accumulation of free amino acids, especially His and Pro, and the activation of theenzymes superoxide dismutase and guaiacol peroxidase. Contents of non-protein thiol,reduced glutathione, phytochelatins and phenolics compounds and activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase also increased significantly forantioxidant or chelation functions. The wet climate population not only accumulated lessabscisic acid, free amino acids, phytochelatins and phenolics compounds, but also exhibitedlower activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase thusresulting in more serious oxidative damage and more curtained growth.

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干旱胁迫是全球范围内影响植物生存、生长和分布的重要环境因子。岷江上游干旱河谷区,由于生态环境的脆弱性和长期人类活动的干扰和过度利用,导致植被严重退化,水土流失加剧,山地灾害频繁,干旱化和荒漠化趋势明显。这种趋势若不能遏制,将严重阻碍区域社会经济的快速协调发展,并且威胁成都平原地区的发展和长江中下游地区的生态安全。因而开展干旱河谷生态恢复研究成为解决这些问题的关键。水分匮乏是限制干旱河谷生态恢复的关键因子,在全球气候变化的背景下,干旱胁迫在区域尺度上可能会更加严重,并使干旱河谷的生态环境更加恶化。因此,深入研究干旱河谷乡土植物对干旱胁迫的响应和适应机理,具有非常重要的理论和实践意义。 本论文以岷江上游干旱河谷的三种乡土豆科灌木,白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha)理论和实践意义。为研究对象,在人工控制条件下设计了4-5个连续性干旱胁迫处理,系统地研究了灌木幼苗的生长、生物量积累和水分利用效率(WUE)、形态结构和生理过程等对干旱胁迫的反应,揭示了幼苗的干旱适应能力及种间差异。主要研究结论如下: 1) 灌木生长和繁殖对干旱胁迫的反应 在干旱胁迫下,幼苗生长速率显著减小,叶片衰老和脱落比率增大,这些变化随着胁迫强度的增加具有累积效应。叶片比茎对干旱胁迫的反应更敏感。在严重干旱胁迫下,幼苗的有性繁殖被限制,但在中等程度干旱胁迫下,幼苗的有性繁殖能力被提高。 2) 灌木生物量积累及其分配和WUE对干旱胁迫的反应 在干旱胁迫下,灌木各器官的生物量都显著减小,但是生物量的分配侧重于地下部分,使得根茎比在干旱条件下增大。幼苗的耗水量(WU)随着干旱胁迫的增加而显著减少。白刺花和小马鞍羊蹄甲WUE在干旱胁迫下降低;小雀花的WUE在中等干旱胁迫下升高。 3) 灌木叶片结构特征对干旱胁迫反应 白刺花叶片具有较为典型的旱生型结构,而小马鞍羊蹄甲和小雀花则为中生型结构。在1至2年的干旱胁迫下,灌木叶片结构组成未发生本质性的改变,主要是细胞大小的变化。在中等和严重干旱胁迫下,叶肉组织厚度明显减小;并且气孔和表皮细胞面积也显著减小,气孔和表皮细胞密度显著增大;叶肉细胞层数、P/S值、表皮厚度等无显著变化。 4) 灌木对干旱胁迫的生理响应 气体交换参数和叶片相对含水量(RWC)在中等干旱胁迫下发生了明显的改变,而叶绿素荧光参数和光合色素含量在严重干旱胁迫下才发生显著变化。在干旱胁迫下,净光合作用速率(Pn)、气孔导度(gs)和RWC呈下降趋势,而叶片温度(Tl)呈增加趋势,蒸腾速率(Tr)的变化不明显。除了日最大Pn减小以外,干旱胁迫对气体交换参数的日变化无显著影响,但是对光合-光响应曲线有显著的影响,使有效光合时间缩短。在严重干旱胁迫下光系统受到损害而代谢减弱,PSⅡ中心的内禀光能转换效率(Fv/Fm)、量子产量(Yield)、光化学淬灭参数(qP)显著降低,而非光化学淬灭参数(NPQ)明显增加。气孔限制和非气孔限制对Pn的影响与干旱胁迫强度有关。在中度胁迫下,气孔限制起主导作用,在严重胁迫下非气孔限制起主导作用,40% FC水分条件可能是灌木由气孔限制向非气孔限制的转折点。 5) 灌木对干旱胁迫的适应能力及其种间差异 三种灌木对干旱胁迫具有较好的适应能力,即使在20% FC,幼苗未因干旱胁迫III而死亡;80% FC适宜于幼苗生长。白刺花生长速率慢,耗水量较少,具有较强的耐旱和耐贫瘠能力,并具有干旱忍受机制,能够在较干旱的环境中定居和生长。小马鞍羊蹄甲和小雀花,生长快,水分消耗量较大,尤其是小雀花,对干旱胁迫的忍受能力较弱,具有干旱回避机制,因而适宜于在较为湿润的生境中生长。综合分析表明,生长速率较慢的物种抗旱能力较强,其更适宜于作为干旱地区植被恢复物种。 Drought is often a key factor limiting plant establishment, growth and distribution inmany regions of the world. The harsh environmental conditions and long-termanthropogenic disturbance had resulted in habitat destruction in the dry valley ofMinjiang river, southwest China. Recently, it tended to be more severe on the vegetationdegradation, soil erosion and water loss, natural disaster, as well as desertification, whichimpact on regional booming economy and harmonious development, and would be verydangerous to the environmental security in the middle and lower reaches of Yangzi River.Therefore, ecological restoration in the dry valley is one of the vital tasks in China. Waterdeficit is known to affect adversely vegetation restoration in this place. Moreover, in thecontext of climate change, an increased frequency of drought stress might occur at aregional scale in the dry valleys of Minjiang River. The selection of appropriate plantingspecies for vegetation restoration in regard to regional conditions is an important issue atpresent and in further. The research on responses of indigenous species to drought stresscould provide insights into the improvement of the vegetation restoration in the dry valleys of Minjiang River. In this paper, the responses of three indigenous leguminous shrubs, Sophora davidii,Bauhinia faberi var. microphylla and Campylotropics polyantha, to various soil watersupplies were studied in order to assess drought tolerance of seedlings, and to compare interspecific differences in seedlings’ responses to drought stress. The results were as follows: 1 Growth and reproduction of shrubs in response to drought stress Seedling growth reduced significantly while leaf senescence accelerated underdrought stress, the cumulative responses to prolonged drought were found. The capacityfor reproduction was limited by severe drought stress, and improved by moderate droughtstress. Leaf responses were more sensitive than shoot to various water supplies. 2 WUE, biomass production and its partitioning of shrubs in response to drought stress Drought stress reduced significantly the total dry mass and their components ofseedlings, and altered more biomass allocation to root system, showing higher R/S ratiounder drought. Water use (WU) and water-use efficiency (WUE) of both S. davidii and B.faberi var. microphylla declined strongly with drought stress. The WU C. polyantha ofalso declined with drought stress, but WUE improved under moderate drought stress. 3 Anatomical characteristics and ultrastructures of leaves in response to drought stress There were xeromorphic for S. davidii leaves and mesomorphic for B. faberi var.microphylla and C. polyantha at the all water supplies. The foundational changes in leafstructures were not found with drought stress. However, mesophyll thickness, the areas ofstomatal and epidermis reduced slightly while the densities of stomatal and epidermisincreased under severe drought stress. Variations in these parameters could mainly be duoto cell size. Other structures did not displayed significant changes with drought stress. 4 Physiological responses of shrubs to drought stress The gas exchange parameters and leaf relative water content (RWC) were affectedby moderate stress, while chlorophyll fluorescence and chlorophyll content were onlyaffected by severe stress. Drought stress decreased net photosynthesis rate (Pn), stomatalconductance, light-use efficiency and RWC, and increased leaf temperature. Therespiration rates (Tr) were kept within a narrower range than Pn, resulting in aprogressively increased instantaneous water use effiecency (WUEi) under drought stress.Moreover, drought stress also affected the response curve of Pn to RAR, there was adepression light saturation point (Lsat) and maximum Pn (Pnmax) for moderate andsevere stressed seedling. However, diurnal changes of gas exchange parameters did notdiffer among water supplies although maximum daily Pn declined under severe stress.VISevere stress reduced Fv/Fm, Yield and qP while increased NPQ and chlorophyll content.Photosynthetic activity decreased during drought stress period due to stomatal andnon-stomatal limitations. The relative contribution of these limitations was associatedwith the severity of stress. The limitation to Pn was caused mainly by stomatal limitationunder moderate drought stress, and by the predominance of non-stomatal limitation undersevere stress. In this case, 40% FC water supply may be a non-stomatal limitation 5 Interspecific differences in drought tolerance of shrubs Three shrubs exhibited good performance throughout the experiment process, evenif at 20% FC treatment there were no any seedlings died, 80% FC water supply wassuitable for their establishment and growth. S. davidii minimized their water loss byreducing total leaf area and growth rate, as well as maintained higher RWC and Pncompared to the other two species under drought stress, thus they might be more tolerantto the drought stress than the other two species. On the contrary, it was found that C.polyantha and B. faberi var. microphylla had higher water loss because of their stomatalconductance and higher leaf area ratios. They reduced water loss with shedding theirleaves and changing leaf orientation under drought stress. Based on their responses, thestudied species could be categorized into two: (1) S. davidii with a tolerance mechanismin response to drought stress; (2) C. polyantha and B. faberi var. microphylla withdrought avoidance mechanism. These results indicated that slow-growing shrub speciesare better adapted to drought stress than intermediate or fast-growing species in present orpredicted drought conditions. Therefore, selecting rapid-growing species might leavethese seedlings relatively at a risk of extreme drought.

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常绿阔叶林以其富饶的生物资源、丰富的生物多样性和巨大的生态与环境效益引起了人们越来越大的重视,它的研究已成为国际植被科学界关注的主题之一。我国分布着世界上面积最大的亚热带常绿阔叶林,在世界植被中具有重要地位,它的分布表现出明显的地带性差异,存在着多样的植物群系及其对应的气候特征。但是在植物功能性状领域,与全球范围其它生物群系相比,常绿阔叶林物种的研究较少,其功能性状间、功能性状与环境间的关系尚不清晰。 本研究以常绿阔叶林木本植物的当年生小枝为对象,试图从小枝水平上的生物量分配格局、叶片大小与数量的权衡关系、小枝茎的构型效应、叶片元素化学计量学,以及小枝大小的成本与效益分析等方面,较为系统地揭示小枝水平上的植物功能性状间及其与气候间的关系。因此,在华西雨屏带内部的不同纬度设置峨眉-青城-雷波-平武的温度梯度进行比较,并对有降水差异的川西南偏湿性(雷波)与偏干性常绿阔叶林(西昌)进行对比研究,同时在不同山体进行不同海拔梯度的比较研究。 本文主要研究结果如下: (1)小枝生物量分配格局叶水平上,叶片重-叶柄重(Y轴vs.X轴,下同)呈斜率小于1的异速生长关系,表明叶柄对叶内部的生物量分配影响显著。小枝水平上,叶和茎的生物量以及它们与小枝总生物量间基本呈等速生长关系,表明大的小枝或大叶物种不一定在叶生物量的分配上占优势。不同生活型间,在小枝或者茎的生物量一定时,常绿物种叶片的生物量比例较落叶物种稍高。与温度和水分较优越(峨眉及其低海拔)的生境相比,在相对低湿(螺髻)与低温(平武)的生境中的植物会减少对叶的投入而增加对支撑部分的投资比例。 (2)小枝叶片大小与数量的权衡无论是不同气候带还是不同生活型以及不同海拔梯度,叶片大小与出叶强度基本都是呈负的等速生长关系,表明了叶片大小-数量在小枝水平上的权衡。在不同气候梯度的对比中,叶片数量(出叶强度)一定时,高温和高水分生境(峨眉)比低温(平武)和低湿(螺髻山)生境中的物种的叶片大小(质量和面积)更大,表明不同生境的比较中,小的叶片可能具有较高的出叶强度和更高的适合度收益。“出叶强度优势”(Leafingintensitypremium)假说可能不适宜解释不同生境物种叶片大小差异。 (3)小枝茎的构型效应虽然茎长和茎径与叶片大小都呈正相关关系,与出叶强度都呈负相关关系,但茎长/茎径比与叶/茎生物量之比呈负相关关系;与叶片的大小呈负相关关系,与出叶强度呈正相关关系。这说明小枝构型能影响小枝叶/茎生物量分配和叶大小-数量的权衡关系。其影响机制可能是小枝内部的顶端优势。另外,茎长/茎径比在低湿和低温等不利生境中的植物中较高,而在降水和温度较适宜环境中较低。 (4)叶片C、N、P化学计量学N含量和P含量,C/N比和比叶重(LMA,leafmassperarea)呈正的等速生长关系,而N和LMA,P和LMA呈负的等速生长关系。在LMA一定时,C/N比随着生境胁迫压力的增加而降低,N、P含量随着生境压力的增加而增加。在P含量一定时,N含量随着生境压力的增加而降低,即N/P比在生境条件较优(峨眉及其低海拔)时较高。常绿和落叶植物叶片的N/P比没有差异,在LMA一定时,常绿植物的N、P含量较高、C/N比较低。总之,植物的C、N、P化学计量学特征受叶片属性如LMA与气候,及其相互作用的影响。 (5)小枝大小的代价与效益分析、TLA与小枝总重总叶面积(TLA,totalleafarea,Y轴,下同)与总叶重(X轴)均呈斜率小于1的异速生长关系,TLA与小枝横切面积呈斜率为1的等速生长关系。表明叶片面积的增加总是小于叶重和小枝总重的增加,随着小枝的增大,它的叶面积支撑效率下降。在热量和降水优越的生境(峨眉及其低海拔)中,相同小枝重或者相同茎横切面积的小枝,其叶面积支撑效率较低湿与低温环境下(螺髻山、平武及高海拔)的高。 总体上,本文初步研究了小枝水平上可能存在的以下三种权衡关系:叶-茎生物量分配权衡;叶片大小-数量的权衡;小枝茎长-茎径的权衡关系,以及气候要素等对这三种权衡关系的影响。在此基础上,我们还讨论了这些权衡关系的可能形成机制,及其与物种生态适应的联系。本研究丰富了生活史对策中关于权衡关系的研究内容,为我国常绿阔叶林功能生态学研究积累了材料。 Evergreen broad-leaved forests are attracting much more attention from vegetation ecologists than ever before because of their abundant nature resource and biological diversity, and also great ecological benefits. China has the largest distribution of subtropical evergreen broad-leaved forests (temperate rainforests) that are typical and representative in the world. The forests span over more than ten degrees in latitude and more than 30 degrees in longitude, providing an ideal place to study plant functional ecology, i.e., the climatic effect on plant functional traits and the relationship between the traits. However, relative to the other biomes, there are few studies addressing functional ecology of the plant species from subtropical evergreen broad-leaved forests. In this study, I focused on the leaf size-twig size spectrum of the woody species of subtropical evergreen broad-leaved forests in southwestern china. I collected data on leaf size and number, twig size in terms of both mass and volume, and stem architecture from five temperate mountains, and then I analyzed the relationships between leaf and stem biomass and between leaf size and number, the effect of stem length/diameter ratio on biomass allocation and on the relationship between leaf size and number, leaf C:N:P stoichiometry, and the twig efficiency of supporting leaf area in relation to twig size. I also addressed the climate effect on the spectrum. The temperature gradient from warm to cool sites was represented by Emei Mountain, Qingchengshan, Leibo, and Pingwu, and the rainfall gradient was assumed to emerge from the comparison between Leibo (High) and Luojishan (Low). In addition, altitudinal effects were analyzed with comparisons between low and high altitudes for each mountains. My main results are as follows. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to leaves or laminas was independent of twig mass. Petiole mass disproportionably increase with respect to lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. In addition, the investigated species tended to have a larger leaf and lamina mass, but a smaller stem mass at a given twig mass at favorable environments including warm and humid sites or at low altitude than unfavorable habitats, which might be due to the large requirements in physical support and transporting safety for the species living at unfavorable conditions. Moreover, the evergreen species invested more in leaves and laminas than the deciduous at given stem or twig biomass within any specified habitats. Negative, isometric scaling relationships between leaf number and size broadly existed in the species regardless of climate, altitude, and life forms, suggesting a leaf size/number trade-off within twigs. Along the climatic gradients, at given leaf number or leafing intensity, the leaves were larger in the favorable environments than the poor habitats. This suggested that the fitness benefit gained by small leaves could be larger than that with high leafing intensity in the stressful sites. I concluded that the “leafing intensity premium” hypothesis was not appropriate to interpreting between-habitat variation in leaf size. Both stem length and diameter were positively correlated to leaf size but negatively correlated to leafing intensity. The ratio of stem length to diameter was negatively correlated to leaf mass fraction, and it was negatively correlated to leaf size but positively correlated to leafing intensity. This suggested that the stem architecture influenced twig biomass allocation and the relationship between leaf size and number. The mechanism underlying the architectural effect might lie in the apical dominance within twig. Moreover, the ratio was greater in unfavorable habitats but smaller in favorable environments. Positive, isometric relationships were found between N and P contents per leaf mass, and between C/N ratio and leaf mass per area (LMA), but N and P contents scaled negatively to LMA. C/N ratio decreased but N and P increased with increasing habitat stress at a given LMA. N content declined with increasing habitat stress at given P content. These indicated that N/P and C/N were higher but LMA was lower in favorable habitats than in the other circumstances. The evergreen and deciduous species were non-heterogeneous in N/P, but the evergreen species have higher N and P contents and lower C/N than the deciduous ones. In general, C:N:P stoichiometry were related to both climatic conditions and other important functional traits like LMA. Total leaf area (TLA) allometricly scaled to leaf mass with a slope shallower than 1, similar to the relationship between TLA and total twig mass (leaf mass plus stem mass), suggesting that TLA failed to keep pace with the increase of leaf mass and twig size. However, TLA scaled isometricly to twig cross-sectional area. Thus, it could be inferred that the twig efficiency of displaying leaf area decreased with increasing twig size. In addition, the efficiency at a given twig size was large in favorable than unfavorable habitats. In general, in this preliminary study, I studied three tradeoff relationships within twigs, i.e., between leaf and stem biomass, between leaf number and size, and between stem length and diameter, as well as the climatic effect on the relationships. I discussed the mechanisms underlying the tradeoff relationships in view of biophysics and eco-physiology of plants. I believe that this study can serve as important materials advancing plant functional ecology of subtropical forest and that it will improve the understanding of life history strategies of plants from this particular biome.

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岷江柏(Cupressus chenggiana S. Y. Hu)是我国川甘地区特有的珍稀濒危乔木,一般生长在干旱的河谷区,在涵养水源和保护水 土等方面起着重要的作用。本文选择4个岷江柏种群,采用了野外调查和室内实验相结合的研究方法,调查岷江柏种群结实状况, 分析种子和球果形态特征,阐明种子发芽的基本特征,研究岷江柏种子贮藏过程中几个生理指标的动态变化特点,目的是为岷江柏 种苗繁育、自然更新能力评估以及珍稀濒危机制分析提供理论依据。研究得出如下结论:1.岷江柏球果呈椭球形,长为1.5~ 2.2cm,宽为1.5~1.9cm,质量为1.7~4.2g,球果鳞片数量为8~11片,球果内种子数量一般在40~70粒。岷江柏种子为椭圆形,长 为3.58~4.02mm,宽为3.10~3.15mm,厚为0.96~1.11mm,千粒重为3.1~3.5g。岷江柏的结实率很低,并且有显著的地理差异和 大小年差异。2. 岷江柏种子发芽温度范围是5℃~30℃,其中种子的适宜发芽温度范围是10℃~25℃。种子最适发芽温度随着贮藏 时间的增加而变化。在适宜温度范围内,种子发芽周期为20d。温度对种子的发芽势和T50有显著影响,对种子发芽率没有显著影响 ;光照有利于种子发芽;岷江柏种子的发芽特征是岷江柏保护种子资源、防止物种濒危的一种环境适应,有助于岷江柏种子提高发 芽率和幼苗的存活率。岷江柏种子是一种耐贮藏的正常性种子,在短期贮藏过程中,贮藏温度和种子含水量对于种子生理指标和种 子发芽没有显著影响。3. 岷江柏种子在短期贮藏过程中,千粒重没有显著变化;含水量都经历了先下降,再稳定的过程;粗脂肪 含量和可溶性糖含量逐渐降低;可溶性蛋白含量和丙二醛含量逐渐增加;脯氨酸含量在贮藏1~7个月时变化差异不明显,但是贮藏 7~10个月后显著增加。岷江柏种子的各个生理指标之间的相关性差异不显著。4. 岷江柏球果和种子的形态特征存在显著的地理差 异。岷江柏种子的发芽能力的地理性差异不大,种群间差异不大。岷江柏种群的地理差异由种群特征、生境特征和气候特征共同决 定。5. 在岷江柏的人工繁育中,对于刚刚采集的种子,发芽温度在15℃~25℃比较适合,其中以25℃最佳;而对于短期贮藏(4~ 10个月)后的种子,发芽温度在10℃~25℃均可,以15℃~20℃为最佳。野外播种的最适时间为4~6月,6~9月的间歇性干旱和降 水波动可能是限制岷江柏自然更新的因素之一。在短期贮藏过程中,种子可以采用常规室温贮藏,可以节约成本。Cupressus chenggiana is a specific and endangered plant in Sichuan and Gansu provinces of China, and it usually grows in dry valley and plays an important role in water supply and soil and water conservation in the dry valley of alpine and canyon region of southwest China. The research selected four Cupressus chenggiana populations and used the methods of the field investigation and the lab experiments. The fruiting characters of Cupressus chenggiana populations, the morphological characters of seeds and cones, the germination characters of seeds and the store physiological dymatics of several factors of seeds have been studied in order to give some theoretical advices on the artificial propagation and the ability of natural regeneration and the endangered principle of Cupressus chenggiana in the paper. The main results may be clarified as follows: 1. The cones of Cupressus chenggiana are ellipsoidal, length ranged from 1.5 to 2.2cm, with ranged from 1.5 to 1.9 cm, weight ranged from 1.7 to 4.2g, the number of cone squama ranged from 8 to 11, and the seed number of per cone ranged from 40 to 70. The seeds of Cupressus chenggiana are elliptical, length ranged from 3.58 to 4.02 mm, width ranged from 3.10 to 3.15 mm, thickness ranged from 0.96 to 1.11 mm, and the weight of 1000 seeds ranged from 3.1 to 3.5g. The fruiting rate of Cupressus chenggiana is very low, and the fruiting period of Cupressus chenggiana has the geographical differences and the big or small year differences. 2. Seed germination temperature is between 5℃ and 30℃, while the suited temperature is between 10℃ and 25℃. The optimum temperature of seed germination will change as the store time of seeds changes logner. The cycle of seed germination can persist 20 days in the range of the suited temperature. The germination temperatures have significant influences on the germination potential and T50, but have no significant infuluences on the germination rate. The photoperiod is in favor of seed germination. The characters of Cupressus chenggiana seed germination represent a kind of environmental adaptability to protect the seed sources and endangered species, and it can give help to increase the germination rate of seeds and the livability of seedings. The seeds of Cupressus chenggiana are a kind of orthodox seeds that can endure the long time storage. In the short time storage, the store temperatures and the moisture contents of seeds have no significant infuluences on the physiological factors and the germination of seeds, but the store time has significant influences on the physiological factors of seeds. 3. In the short store course of Cupressus chenggianna seeds, the 1000 seed weight has no significant variation; The moisture content descends at the beginning of the storage, but has no significant variation later; The crude fat content and the soluble sugar content descend gradually; The soluble protein content and MDA content increase gradually; The praline content has no significant variation after 1~7 months storage, but increase significantly after 7~10 months storage. The correlations of different physiological factors are not significant. 4. The morphological characters of cones and seeds of four populations exist significant differences. The germination of Cupressus chenggiana seeds has no significant geographical variation. The geographical variation of Cupressus chenggiana populations can be ascribed to the population characters, climate and environment. 5. In the course of artificial propagation of Cupressus chenggiana, it is favored that the germination temperature of newly collected seeds is between 15℃ and 25℃, while the optimum temperature is 25℃. After the short storage ranged from 4 months to 10 months, it is favored that the germination temperature is between 10℃ and 25℃, while the optimum temperature is ranged from 15℃ to 20℃. The field sowing optimum time is between April and June, and the interval drought and fallrain fluctuation between July and September may be one of the reasons that restrict natural regeneration of Cupressus chenggiana. In the short storage, seeds can be stored in the condition of room temperature.

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光是植物赖以生存的重要环境因子,但是植物在获得光的同时不可避免的会受到紫外辐射的伤害。尤其是近年来,人类向大气中排放的大量氮氧化合物和氟氯烃类化合物(CFC’s)引起臭氧分子的分解,导致到达地球表面的紫外辐射增加,特别是UV-B辐射增强。而另一方面,植物对UV-B辐射反应的敏感性在种间和品种间存在差异,主要受植物基因型,生态型和生活型的控制。本项目分别以粗枝云杉和青杨组杨树为模式植物,从形态和生理生化方面分别研究了来自不同水分背景下的粗枝云杉种群和来自不同UV-B背景下的青杨种群在增强UV-B下的反应及其反应差异,并探讨了干旱、喷施外源脱落酸(ABA)对它们抗UV-B能力的影响。研究成果可为生态系统的恢复与重建提供理论依据和科学指导。主要研究结果如下: 1. 粗枝云杉的两个种群,湿润种群(来自四川黑水)和干旱种群(来自甘肃迭部)在水分良好和干旱状况下表现出对增强UV-B的不同响应。同时,干旱对粗枝云杉抗UV-B能力的影响也得到研究:两种胁迫共同作用时,干旱表现出在一定程度上减弱了增强UV-B对粗枝云杉的生理特性的影响。 干旱胁迫显著降低了两个粗枝云杉种群的光合同化速率(A), 气孔导度(gs)和PSII的有效光量子产量(Y), 同时,提高了非光化学猝灭效率(qN)和超氧化物歧化酶(SOD)的活性。与湿润种群相比,干旱种群抗旱性更强,表现为干旱种群拥有更高的SOD和干旱进一步加剧了UV-B的胁迫效应。 本研究中,干旱胁迫单独作用时,显著降低了青杨两个种群的生物量积累和气体交换,具体包括A、gs、蒸腾速率(E)和光合氮利用效率(PNUE),提高了两个种群的瞬时水分利用效率(WUEi)、长期水分利用效率(WUET)、碳同位素组分(δ13C)和氮含量(N)。同时,UV吸收物质和ABA含量也得到积累。另一方面,增强UV-B对青杨两个种群各个指标的影响,同干旱所引起的效应有着相似的趋势。同低海拔种群相比,高海拔种群有着更强的抗旱和抗UV-B能力,具体表现在高海拔种群有着更多的生物量积累,更强的气体交换和水分利用效率及更高水平的ABA和UV吸收物质含量。相比干旱诱导的生物量积累和气体交换的降低,在干旱和增强UV-B两个胁迫同时作用于青杨时,这种降低表现的更为明显。显著的干旱和UV-B的交互作用还表现在WUEi, WUET, δ13C, 可溶性蛋白含量, UV吸收物质含量, ABA, 叶片和茎中的N含量以及C/N比中。 3. 经过一个生长季的试验观察,增强UV-B、外源ABA及两因子共同作用对青杨的生物量积累、气体交换、内源ABA和UV吸收物质含量、抗氧化系统以及碳、氮含量和碳/氮比均产生显著影响。本试验中,青杨的两个种群分别来自中国西南部的不同海拔地区,高海拔种群来自青海大通而低海拔种群来自四川九寨。外源ABA的胁迫为直接喷施ABA到青杨叶片,而增强UV-B胁迫是利用平方波系统分别保证青杨苗暴露于外界UV-B强度和两倍于外界UV-B强度下。 研究结果显示,增强UV-B显著的降低了两个青杨种群的株高、基茎、总叶面积和总生物量等生长指标,同时也导致其A、gs、E和叶片中碳含量的减少。而显著增加了SOD和过氧化物酶(GPx)活性水平,诱导了过氧化氢(H2O2)和MDA的显著增加,促进了UV吸收物质和不同器官中内源ABA含量的显著积累。另一方面,外源ABA引起了青杨光合同化速率的下降,SOD和GPx酶活性的增强,H2O2 和 MDA含量也表现出显著增加,同时,内源ABA含量得到显著累积。同低海拔种群相比,高海拔种群具有更加抗UV-B和外源ABA的特性。显著的UV-B和ABA的交互作用表现在A, E, SOD和GPx活性,以及叶片和根部的内源ABA等一系列指标中。在所有胁迫下,叶片中的碳和氮含量同其在茎和根中的含量显著相关,另外,叶片和茎中的氮含量同茎中的碳含量显著相关。 Sunlight is an indispensable environment factor for plants survival and development. Meanwhile, photosynthetic organisms need sunlight and are thus, inevitably, exposed to UV radiation. Especially for recent years, ultraviolet radiation, especially UV-B reaching the Earth’s surface increased because of depletion of ozone layer resulted from emission of NxO and CFC’s from human activities. On the other hand, the sensitivity of plants to UV-B radiation depends on the species, developmental stage and experimental conditions. In this experiment, two populations of Picea asperata Mast from different water background and two populations of Populus cathayana Rehder from different altitude background were selected as model plants to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B in each plant species were observed and the different responses were discussed, furthermore the influences of drought and exogenous ABA on responses induced by enhanced UV-B were studied. The study could provide a strong theoretical evidence and scientific direction for the afforestation and rehabilitation of ecosystem. The results are as follows: 1. Different responses of two contrasting Picea asperata Mast. populations to enhanced ultraviolet-B (UV-B) radiation under well-watered and drought conditions were investigated. And the effects of enhanced UV-B on tolerance of drought were also observed in our study that the UV-B exposure may have alleviated some of the damage induced by drought. Two contrasting populations, originating from a wet and dry climate region in China, respectively, were employed in our study. Drought significantly decreased CO2 assimilation rate (A), stomatal conductance (gs) and effective PSII quantum yield (Y), while it significantly increased non-photochemical quenching (qN) and the activity of superoxide dismutase (SOD) in both populations. Compared with the wet climate population, the dry climate population was more acclimated to drought stress and showed much higher activities of SOD and ascorbate peroxidase (APX), and much lower levels of malondialdehyde (MDA) and electrolyte leakage. On the other hand, enhanced UV-B radiation also induced a significant decrease in the chlorophyll (Chl) content in both populations under well-watered conditions, and a significant increase in UV-absorbing compounds in the wet climate population. After one growing season of exposure to different UV-B levels and watering regimes, the increases in MDA and electrolyte leakage, as induced by drought, were less pronounced under the combination of UV-B and drought. In addition, an additive effect of drought and UV-B on A and gs was observed in the wet climate population, and on the activity of APX and qN in the dry climate population. 2. The significant effects of drought, enhanced UV-B radiation and their combination on Populus cathayana Rehd. growth and physiological traits were investigated in two populations, originating from high and low altitudes in south-west China. Our results showed that UV-B acts as an important signal allowing P. cathayana seedlings to respond to drought and that the combination of drought and UV-B may cause synergistically detrimental effects on plant growth in both populations. In both populations, drought significantly decreased biomass accumulation and gas exchange parameters, including A, gs, E and photosynthetic nitrogen use efficiency (PNUE). However, instantaneous water use efficiency (WUEi), transpiration efficiency (WUET), carbon isotope composition (δ13C) and nitrogen (N) content, as well as the accumulation of soluble protein, UV-absorbing compounds and abscisic acid (ABA) were significantly increased by drought. On the other hand, cuttings from both populations, when kept under enhanced UV-B radiation conditions, showed very similar changes in all above-mentioned parameters, as induced by drought. Compared with the low altitude population, the high altitude population was more tolerant to drought and enhanced UV-B, as indicated by the higher level of biomass accumulation, gas exchange, water-use efficiency, ABA concentration and UV-absorbing compounds. After one growing season of exposure to different UV-B levels and watering regimes, the decrease in biomass accumulation and gas exchange, induced by drought, was more pronounced under the combination of UV-B and drought. Significant interactions between drought and UV-B were observed in WUEi, WUET, δ13C, soluble protein, UV-absorbing compounds, ABA and in the leaf and stem N, as well as in the leaf and stem C/N ratio. 3. During one growing season, significant effects induced by enhanced UV-B radiation, exogenous ABA and their combination on biomass accumulation, gas exchange, endogenous ABA and UV-absorbing compounds concentrations, antioxidant system as well as carbon (C) content, nitrogen (N) content and C/N ratio were investigated in two contrasting Populus cathayana populations, originating from high and low altitudes in south-west China. Exogenous ABA was sprayed to the leaves and enhanced UV-B treatment was using a square-wave system to make the seedlings under ambient (1×) or twice ambient (2×) doses of biologically effective UV-B radiation (UV-BBE). Enhanced UV-B radiation significantly decreased height, basal diameter, total leaf area, total biomass, A, gs, E and carbon (C) content in leaves, and significantly increased activities of SOD and guaiacol peroxidase (GPx), hydrogen peroxide (H2O2) and malonaldehyde (MDA) content as well as the accumulation of UV-absorbing compounds and endogenous ABA concentrations among different organs in both populations. In contrast, exogenous ABA showed significant decrease in A and significant increases in activities of SOD and GPx, H2O2, MDA content and the endogenous ABA concentrations. Compared with the low altitude population, the high altitude population was more tolerant to enhanced UV-B and exogenous ABA. Significant interactions between UV-B and ABA were observed in A, E, activities of SOD and GPx, as well as in endogenous ABA in leaves and roots of both populations. Across all treatments, C and N content in leaves was strongly correlated with those were in stems and roots, respectively. Additionally, leaf and stem N content were significant correlated with stem C content.

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杨树具有分布广、适应性强,在生态环境治理和解决木材短缺方面均占有重要位置。青杨(Populus cathayana Rehd.)是青杨派树种的重要成员之一,也是我国的特有种。本研究通过对不同水分梯度的干旱胁迫下青杨形态和生理生化的反应,不同pH值盐碱胁迫下不同海拔和不同气候地区的四个青杨种群在生理生态上的反应差异,以及在干旱和低温胁迫下青杨lea2, lea3组基因表达差异的研究,从形态、生理、生化和分子生物学水平系统地研究了青杨在不同逆境胁迫下的反应和青杨不同种群在盐碱胁迫下的反应差异。主要研究结果如下: 1. 青杨在干旱胁迫下的反应机制:中度和重度干旱胁迫下植株的生长受到明显抑制。表现在光合系统上青杨的净光合同化速率(A)下降,主要原因是气孔导度(gs),胞间二氧化碳浓度(Ci)下降。另外最大量子产量(Fv/Fm)、光化学猝灭效率(qP)降低反应了干旱胁迫下光合系统II(PSII)受到严重损伤, 而且非光化学猝灭效率(qN)上升,导致可利用化学能产量下降,叶绿体产生淀粉的量减少。qP降低qN上升导致产生的过量电子对光合系统的伤害造成活性氧以及丙二醛(MDA)的含量增加。超微解剖结构显示,干旱胁迫增强时,叶绿体内淀粉粒的数目减少,而且叶绿体、线粒体等细胞器中嗜锇颗粒的数目增加。为清除细胞内的活性氧,植物一般的反应是抗氧化系统酶活性增加,对青杨来讲超氧化物歧化酶(SOD), 抗坏血酸过氧化物酶(APx)活性的增加远大于过氧化物酶(POD),这显示了在青杨中SOD、APx酶在清除活性氧的作用上大于POD。另外同工酶研究结果显示这些酶活性的升高主要是由于各条同工酶带表达量的增加,而不是诱导新酶带的产生。另外,75% FC水分处理下有些指标非但没有下降,像A和有效光量子产量(Y)的值都略有增加,而且gs同时增加。另外,100% FC比75% FC细胞内淀粉粒的数目少一些,但有少量的嗜锇颗粒。这证明100% FC土壤水分也许并非最适合青杨生长。 2. 盐碱胁迫对不同海拔地区青杨种群的反应差异:青杨高海拔和低海拔种群的各种生理特性随着pH值上升都受到了很大的影响。两种群叶和根中Na+、K+ 含量, Na+/K+比率随着pH值的上升影响显著。在pH值高于10.4时高海拔种群叶和根中Na+/K+比率急剧下降但是低海拔种群中却一直维持在较高水平。两种群中MDA、脯氨酸(Proline)的含量,抗氧化系统酶的活性都受到了严重的影响,证明两个种群都属于盐碱胁迫敏感类型但是高海拔的种群对盐碱胁迫的耐性要高于低海拔。这主要是由于高海拔种群一般具有耐干旱、低温胁迫的能力,而植物的抗逆机制一般都有共通之处。 3. 盐碱胁迫对不同气候地区青杨种群的反应差异:盐碱胁迫下两种群的光合作用受到明显的抑制,具体表现在叶绿素的含量和A 显著下降。净光合速率的下降主要是由于叶片gs,Ci 值降低引起的。与湿润地区的种群相比盐碱胁迫增强时,干旱地区的种群叶绿素含量和光合能力的升高与K+离子含量增加有关。植物维持细胞质高K+/Na+值对植物的抗盐性有很重要的作用。为清除盐碱胁迫产生的活性氧,抗氧化系统酶活性增加。盐碱胁迫下干旱地区的种群在SOD、CAT 和谷胱甘肽还原酶(GR)等酶的活性均显著上升,而湿润地区种群只有谷胱甘肽氧化酶(GST)的活性明显增加,说明干旱种群的抗氧化酶系统在较高盐碱胁迫下的保护作用要强于湿润种群。这主要是由于植物抗盐碱胁迫与抗干旱胁迫在一些方面的机制是一致的,抗旱种群一般也能抵抗一定程度的盐碱胁迫。 4. 青杨lea2、lea3 基因在干旱和低温胁迫下的表达差异:通过荧光定量PCR 分析,lea2、lea3 组基因在干旱和低温胁迫下在mRNA 水平的瞬时表达量明显升高,说明了两基因在青杨耐干旱和低温胁迫上都起显著的作用。而且两基因在干旱胁迫下,表达量的升高和降低的时间近乎同步,表明两基因在干旱胁迫下对植物应急保护机制的启动都发挥着重要的作用。低温胁迫下lea3 基因在mRNA 水平上表达量显著上升的时间要早于lea2,而且lea3 基因的持续作用时间明显长于lea2 组基因,说明了低温胁迫开始时lea3基因在植物应对逆境的作用上要大于lea2 基因。 Poplars play an important role in lumber supply, and are important components of ecosystems due to their wide distribution and well adaptation. Populus cathayana Rehd., which belongs to Populus Sect. Tacamahaca Spach, is one of the most important resources of poplars and is specialist to china. In this study, different altitudes and climates populations of P. cathayana were used as experiment materials to investigate the adaptability to drought and salt-alkali stresses. And the cultures of P. cathayana were used to analyze the lea2 and 3 group genes expression when exposed to drought and low temperature stresses. The results are as follows: 1. A large set of parallel responses to drought stress: Drought stress caused pronounced growth inhibition. A decreased significantly and was mainly the result of gs and Ci down. Besides, Fv/Fm, qP decreased and that reflected the harmful effects to PSII of drought stress. In accordance with qN increasing, decreased useful energy production caused the starch numbers reduction in chloroplast. The qP up and qN down improved the levels of ROS and MDA. Starch numbers in chloroplast reduced and plastoglobuli numbers increased when soil water content decreased. To reduce ROS, the activities of SOD, APX, CAT and PPO were activated. The isozymes results show that the rising activities of the antioxidant enzymes resulted from certain isoform content increased, and not from the new band produced. Interestingly, morphological results show 100%FC maybe wasn’t the favorite water content for P. cathayana growth. 2. Effect of salt-alkali stress on morphological and physiological changes in two different altitudes populations of P. cathayana: We compared the physiological responses of two populations of Populus cathayana Rehder, originating from altitudes 2,840 m and 1,450 m. Our results demonstated that Na+ and K+ contents, and Na+/K+ ratios in leaves and roots are greatly affected by pH values. At pH 10.4, the Na+/K+ ratios in both leaves and roots sharply dropped in the higher altitude population but were always maintained at higher levels in the lower altitude population. The pH values causing maximum malondialdehyde (MDA) level, free proline content and antioxidant enzyme activities were significantly different in two populations. These results indicated that the higher altitude population exhibits greater tolerance to alkalinity stress than does the lower altitude population. 3. Morphological and physiological changes in two different climates populations of P. cathayana when exposed to salt-alkali stress. Salt-alkali stress caused pronounced inhibition of the growth and especially in photosystem. Pigments content and A decreased significantly and at the same time gs and Ci decreased too. Compared with wet climate population, the Chlorophyll content and A increased in drought climate population as pH value rising was related to the K+ content increasing. It is important to resist salt-alkali stress that the K+/Na+ ratio matained at high level in cytoplasm. To reduce ROS content, the SOD, CAT and GR activities rised significantly in drought population but only GST increased in wet population. The drought population showed higher salt-alkali tolerance than the wet population mainly resulted from the fact that drought tolerance was in accordance with salt-alkali tolerance to some extent. 4. The different expressional model of lea2 and lea3 gene when P. cathayana was exposed to drought and cold stress. RT-PCR results show both lea2 and lea3 suddenly expressed significantly in mRNA level under drought and cold stress. The expression level of two genes reached optimal level at the same time. But under cold stress, the earlier significantly rising expressional time and the longer maintained higher level time in lea3 than lea2 elucidated that lea3 may be more important than lea2 in resisting cold stress in short time in P. cathayana.