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本文以形态特征为主,并参考其它证据,将中国Sect. Metapolystichum分为2个亚组,8个系,共记载45种2变种,其中新分类群21种1变种,新分布1种,新组合1个,并有11个新异名。对每种载有文献和标本引证及简要特征,新种则记有拉丁描述,并对某些疑难种进行了讨论。本文对叶表皮细胞的解剖研究表明,中国Sect.Metapolystichum植物气孔基本类型有两种,即十字型四细胞型和轴极型。各种植物的叶表皮细胞形态特征都不尽相同,有一定的分类学意义。作者对孢子形态的研究表明,中国Sect. Metapolystichum植物孢子周壁纹饰可分为八类,即波状、皱波状、条纹状、脑纹状、假网状、穴状、网状和复网状;在44种2变种中,孢子周壁纹饰表现出43种不同的形态。这一研究为分类上解决一些疑难问题、组下分类及系统发育研究提供了重要依据。本文中至少23种植物的孢子形态属首次报道。本文还根据中国Sect. Metapolystichum植物形态、解剖和孢子特征,遵循分支系统学的理论,借以确定了本组植物27个性状的极性;应用最大同步法(略有改进),通过简约性分析,构建了中国Sect. Metapolystichum植物有根的Wagnet树。作者对中国Sect. Metapolystichum植物的植物地理学特征也进行了分析,讨论了本组植物地理分布特征、间断分布现象、特有现象、替代现象、植物区系成分等,并就中国Sect. Metapolystichum植物区系进行了初步分区。
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群落结构的确定是群落研究的基础。本文立足于个体、种群、群落三级水平,从空间和时间两种概念出发,对贡嘎山麦吊杉(Picea brachytyla)群落的结构进行了研究。经初步统计,在麦吊杉林分布的主要地段共有维管束植物93科、263属、约560种,其中种子植物71科、220属、479种。通过种子植物区系地理成份的分析表明,麦吊杉群落的区系组成具有区系成分复杂、特有现象明显和成分古老的特点。在生活型组成上,麦吊杉群落的高位芽植物所占比例最大,其中小高位芽植物最多,而在叶级谱中小叶类型占有较明显的优势。通过种—面积曲线和重要值—面积曲线两种方法求得的麦吊杉群落的最小面积在800-900平方米之间,实际应用时可选用1000平方米的保守面积。在物种的综合特征分析中,群落表现出优势种明显的特征,而存在度II占有最大的比例。物种多样性的研究表明,贡嘎山麦吊杉群落的Shannon-Wiener指数在2.33-3.26之间,Simpson指数在0.43-0.70之间,种间相遇(PIE)在0.42-0.79之间,而以Shannon-Wiener指数为基础的均匀度在34.45%-62.90%之间。麦吊杉群落的垂直结构较为复杂,分层明显。一般可划分出乔木层、灌木层、草本层和活地被层四个基本层次,乔木层通常还可划分为2-3个亚层,由高海拔至低海拔成层现象有逐渐复杂的趋势。构成麦吊杉群落的层片类型较多,其中以常绿针叶大高位芽植物为其建群层片。麦吊杉林中的层外植物较丰富,种子植物、蕨类植物、以及苔藓植物和地衣植物皆可构成其空中层片,其中苔藓植物生长繁茂,反映出群落结构的完整性。麦吊杉群落结构的水平分化较明显,经研究,其密度值在0.32-0.41之间,平均距离在1.56-1.41之间。应用Raunkiaer的频度指数方法对整个麦吊杉群落的种群频度进行调查后表明,其频度分布情况与Raunkiaer的频度定律基本上是符合的,即A>B> C
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岷江上游半干旱河谷地区地处横断山区东北边缘,在31°15'N-32°32'和103°10'E-103°54'E之间,气候干暖,焚风和山谷风强烈,半干旱河谷灌丛分布在河面以上300-400(500)m的海拔幅度内。植物区系以温带成分占优势,但热带成分也相当多,且在灌丛的灌木层和草本层中占有较显著的地位,显示出植物区系由热带、亚热带向温带植物区系过渡到特点;另外植物区系还具有地区特有种多,与其它地区植物区系,尤其是川西其它几条大河的干旱河谷植物区系关系密切的特点,区系中出现了一定数量的中亚荒漠及半荒漠成分,其中刺旋花在群落中成为建群种之一,值得重视。半干旱河谷植物表现出一系列适应干旱环境的生态特征;在生活型谱中,高位芽植物和地面芽植物占优势。灌丛可依据建群种(或指示种)分为2个植被亚型,2个群系组(暖温性半干旱河谷落叶阔叶灌丛和温性半干旱河谷落叶阔叶灌丛),8个群系和11个群丛。植被的分布表现出比较明显的水平和垂直分布规律。半干旱河谷的治理应遵循工程措施与生物措施相结合的原则。并非所有地段都可直接造林。在造林中应注意:总体规划、先易后难、集中成片、 适宜的乔灌草种和造林技术的选择。
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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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近年来,随着对作物重茬(连年种植)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。而作为重要调料和药用植物的生姜,其连作障碍也备受关注,系统地研究生姜化感作用将有助于理解和最终解决生姜连作障碍问题。本文通过研究生姜不同部位、不同浓度的水浸液对与其间作的两个物种(大豆和四季葱)种子的萌发及幼苗生长的影响,从而证明生姜化感作用的存在;并通过温室盆栽实验研究了生姜的自毒作用(即研究生姜不同部位、不同浓度的水浸液对其幼苗的形态、生理生化、光合作用、土壤酶、土壤微生物多样性及土壤养分的影响),从而揭示生姜退化和衰老的机制,并为生姜筛选出合适的间作物种提供科学依据,对生姜连作障碍提出科学的解决方法。主要研究结果如下: 1. 与对照相比,生姜所有部位(根茎、茎、叶)、所有浓度(10、20、40、 80 g l-1)的水浸液均抑制了大豆种子和葱籽的萌发率、幼苗生长、水分吸收和脂肪酶活性,并且其抑制程度随着水浸液浓度的增加而增强,其生姜各部位水浸液抑制效应的强弱顺序为茎>叶>根茎。这一结果表明生姜根茎、茎、叶含有能够抑制大豆种子和葱籽种子萌发和幼苗生长的水溶性化感物质。根茎是生姜的主要收获部位,而生姜的残株(主要是茎和叶)应该从大田中处理掉以减轻其抑制效应。生姜水浸液中主要化感成分包括:根茎水浸液中主要是丁香酸和伞花内脂;茎水浸液中主要是阿魏酸,且其含量最高为73.4 ug/g;叶水浸液中除了阿魏酸,其他六种物质均检测出来,但含量较高的主要有丁香酸、伞花内脂和香豆酸。 2. 生姜茎和叶不同浓度的水浸液均显著抑制了生姜幼苗的株高、每株叶片数和叶面积,其抑制程度随着水浸液浓度的增加而有所增强,而生姜幼苗每株分枝数差异不显著;同时生姜水浸液也极大程度地影响了生姜幼苗的生物量(包括地下生物量、地上生物量和总生物量,均为鲜重)。在同一浓度下,茎水浸液对生姜幼苗形态指标及生物量指标均显示出最强的抑制作用,叶水浸液次之,根茎水浸液最弱。与对照相比,低浓度的生姜根茎水浸液提高了生姜幼苗叶片内四种抗氧化酶(SOD、POD、CAT、APX)活性,高浓度的根茎水浸液抑制了四种抗氧化酶活性,而茎和叶水浸液均随着浓度的增加而抑制了四种抗氧化酶活性,三种水浸液均随着浓度的增加降低了生姜幼苗叶片内叶绿素的含量,而增加了生姜幼苗叶片的相对电导率和丙二醛含量。同时,三种水浸液均随着浓度的增加降低了生姜幼苗的光合参数(包括胞间CO2浓度、气孔导度、蒸腾速率及净光合速率)。 3. 三种生姜水浸液对所测六种土壤酶活性均产生了不同程度的影响,其中影响最大的是酸性磷酸酶和蔗糖酶,在10 g l-1 时就达到了显著水平,并且所有酶均有随着水浸液浓度增加而增大的趋势;相同部位的水浸液随着浓度的增加,细菌和真菌的数量呈增加趋势,而放线菌的数量呈减少趋势;三种生姜水浸液均随着浓度的增加降低了土壤中有机质的含量,加剧了土壤中硝态氮含量的积累,根茎水浸液对土壤有效磷、速效钾和铵态氮均显示出低浓度提高其含量而高浓度降低其含量的趋势,而茎和叶水浸液则随着浓度的增加均降低了其含量。 4. 与生姜单作相比,所有间作系统均在旺盛生长期和收获期不同程度地提高了土壤酶活性,同时也增加了土壤细菌数量及土壤微生物总数但不显著;所有间作系统在旺盛生长期和收获期均不同程度地影响了土壤真菌及放线菌数量(增加或减少),所有间作系统间的多样性指数差异不显著,除了旺盛生长期四种作物(生姜-大豆-四季葱-大蒜)的间作模式显著降低了多样性指数,其值仅为生姜单作的33.18%;生姜与大豆间作不仅提高了19.6%的生姜产量而且获得了较好的经济效益,并且,所有间作系统均显著抑制了生姜姜瘟病的发生。 5. 不同栽培模式不同程度地影响了收获期生姜的株高、分枝数、根茎产量及内在品质。其中处理2显著地促进了生姜的分枝(10.5%),同时处理2、3和4也促进了生姜的生长(株高分别增加了15.0%、11.4%和14.0%),并且这三个处理提高了生姜的产量;处理2和3能有效提高生姜块茎中维生素C(分别较单作生姜显著提高了3.29%和4.05%)、处理3显著提高了可溶性糖(8.2%)、姜辣素(4.6%)和蛋白质等有益物质的含量,降低硝酸盐有害物质的含量(处理2显著降低了14.0%),改善了姜块的外观和内在品质。并且,生姜与大豆间作具有最高的纯收入和产投比,分别较生姜单作提高了24.80%和8.8%。Recently, allelopathy has been more and more paid attentions by national and foreign scholars with profound research on reasons of crop replanted (continuous planted) obstacle. Ginger rhizome is valuable all over the world either as a spice or herbal medicine and ginger replanted obstacle is also paid attentions. Systematic research on ginger allelopathy will contribute to understanding and ultimate solving problem of ginger replanted obstacle. The effects of ginger aqueous extracts with different parts and concentrations on seed germination and early seedling growth of soybean and chive were studied in this article to testify that ginger existed allelopathy. Furthermore, ginger autotoxicity was also studied by pot experiment in greenhouse (namely research on effects of ginger aqueous extracts with different parts and concentrations on morphological indexes, physiological and biochemical indexes, photosynthesis, soil enzymes, soil microbial diversity and soil nutrients) to reveal mechanism of ginger degeneration and senescence, provide scientific basis for selecting appropriate intercropping species and put forward scientific resolvent for ginger replanted obstacle. The main results were as follows: 1. All aqueous extracts at all concentrations inhibited seed germination, seedling growth, water uptake and lipase activity of soybean and chive compared with the control, and the degree of inhibition increased with the incremental extracts concentration. The degree of toxicity of different ginger plant parts can be classified in order of decreasing inhibition as stem>leaf>rhizome. The results of this study suggested that rhizome, stem and leaf of ginger contained water soluble allelochemicals which could inhibit seed germination and seedling growth of soybean and chive. The rhizome is the main harvested part of ginger. The residue (mainly stems and leaves) of the ginger plant should be removed from the field so as to diminish its inhibitory effect. The main allelopathic components of three kind of aqueous extracts were as follows: Rhizome extract chiefly contained syringic acid and vmbelliferone and stem extract mainly contained frulic acid whose content was the highest (73.4 ug/g). The other six substances were detected except of frulic acid, but only contents of syringic acid, vmbelliferone and p-coumaric acid were higher. 2. Stem and leaf aqueous extracts of ginger with different concentrations significantly inhibited plant height, leaf numbers per plant and leaf area, and the degree of inhibition increased with the incremental extracts concentration. However, tiller number per plant of ginger seedling showed no significant difference. At the same time, ginger aqueous extracts also influenced biomass including under-ground biomass, above-ground biomass and total biomass (fresh weight) to a large extent. Under the same concentration, stem aqueous extract showed the mostly inhibitory effect on morphological indexes and biomass indexes of ginger seedling. Rhizome aqueous extract showed the leastly inhibitory effect and leaf aqueous extract was intervenient. Enhanced concentration of ginger aqueous extracts significantly reduced total chlorophyll content, accompanying with increases in memberane permeability (REL) and lipid peroxidation (MDA). Compared with the control, rhizome ginger aqueous extract of lower concentration (10 g l-1) increased the activities of major antioxidant enzymes (superoxide dismutase, SOD; peroxidase, POD; catalase, CAT; ascorbate peroxidase, APX) of ginger leaf tissue and higher concentration inhibited the activities of four antioxidant enzymes. However, stem and leaf aqueous extract inhibited the activities of four antioxidant enzymes with increase in concentration. Meanwhile, enhanced concentration of ginger aqueous extracts significantly reduced photo-parameters of ginger seedling (including CO2 concentration, stoma conductivity, net photosynthesis rate and transpiration rate). 3. Rhizome, stem and leaf ginger aqueous extract showed different effect on six soil enzyme activities, and acid phosphatase and invertase showed significant effect when aqueous extract concentration got 10 g l-1. Furthermore, six soil enzyme activities increased with increase in aqueous extract concentration. Bcterial and fungi number tended to increase while antinomyces tented to decrease with the increase in aqueous extract concentration of identical part. Ginger aqueous extracts reduced soil organic matter content with increased concentration, accompanying with NO3-—N accumulation in soil. Rhizome aqueous extract showed the same tendency for available P, available K and NH4+—N, namely lower concentration increased their contents in soil and higher concentration reduced their contents. While stem and leaf aqueous extracts reduced their contents with the increamental concentration. 4. All intercropping systems increased soil enzyme activities to different extent both at VGS and at HS compared to solo ginger. All intercropping systems increased the colony numbers of soil bacteria and total of soil microbe but not significantly either at VGS or at HS. All intercropping systems increased the colony numbers of soil fungi and actinomytes to a different extent (increase or decrease) both at VGS and at HS. For DI, difference between all cultivation patterns and S-G was not significant either at VGS or at HS except that G-S-C-G whose value was only 33.18% of S-G at VGS significantly decreased. G-S not only increased ginger yield by 19.6% but also obtained better economic benefit. Furthermore, all intercropping systems significantly inhibited occurrence of bacterial wilt of ginger. 5. Different cultivated pattern influenced plant height, tiller numbers, rhizome yields and intrinsic quality of ginger. Treatment 2 significantly facilitated tiller occurring (10.5%). Treatment 2, 3 and 4 promoted ginger growth (plant height respectively increased 15.0%、11.4% and 14.0%) and enhanced rhizome yields. Treatment 2 and 3 effectively increased vitamin C content (significantly increased 3.29% and 4.05% compared to solo ginger). Treatment 3 significantly increased contents of beneficial substances such as soluble sugar (8.2%), gingerols (4.6%) and protein. Treatment 2 significantly decreased contents of deleterious substance namely nitrate (14.0%) and improved appearance and intrinsic quality of ginger rhizome. Furthermore, treatment 2 (ginger/soybean intercropping) could obtain better economic benefit and showed the highest net income and ratio of benefit and cost whose values respectively increased by 24.80% and 8.8% compared to solo ginger.
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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,目前土壤受到干旱和盐胁迫的危害越来越严重。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,研究杨树对土壤干旱和盐胁迫的生态生理及蛋白质组学反应,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,也为恢复与重建盐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 青杨不同种对逐步干旱胁迫的响应差异 将来自喜马拉雅山东缘高海拔的康定杨和低海拔的青杨枝条扦插在温室中,用来检测它们对逐步干旱胁迫的响应。研究结果表明来自不同海拔的杨树对逐步干旱胁迫的适应性反应是不一样的。株高、叶片发育、叶片相对含水量、丙二醛、过氧化氢等指标的显著性变化在青杨中比在康定杨中来得早些,而且随着干旱胁迫程度的增加,这些参数的变化越来越明显,尤其是当青杨受到严重干旱胁迫的时候;而可溶性蛋白、可溶性糖、游离脯氨酸、抗氧化酶活力变化在康定杨中来得早一些。与青杨相比,在干旱胁迫下,康定杨仍能保持较好的植株生长和叶片发育;康定杨也能在逐步干旱条件下积累更多的可溶性蛋白、可溶性糖、游离脯氨酸及抗氧化酶活力,但是在丙二醛和过氧化氢含量方面增加的更少些。而且,我们的研究结果表明高海拔的康定杨有更强的耐干旱能力,杨树对干旱胁迫的适应能力与干旱发生的速度、强度、持续时间及两种杨树的海拔有关。 2 干旱胁迫下青杨不同种的蛋白质组学分析 来自青杨和康定杨雌株的枝条扦插在温室中,用来研究它们对干旱胁迫的蛋白质组学反应。采用TCA-丙酮/酚提取法提取总蛋白,并进行双向电泳分析。在每个处理的重复图像中都能检测到1,000 个以上的蛋白点。在青杨中有58 个蛋白在干旱处理后发生显著变化,其中22 个蛋白通过肽指纹图谱成功鉴定。康定杨中有69 个蛋白的表达量发生了显著变化,其中有25 个蛋白通过肽指纹图谱成功鉴定。这些被鉴定的蛋白主要参与了光合作用、氧化还原平衡、信号传导、能量代谢、蛋白质合成等过程。尽管被鉴定的蛋白只占叶片总蛋白的很少一部分,但这些被鉴定的干旱响应蛋白可能对维持植株内部平衡方面有重要作用。 3 青杨的盐胁迫响应 青杨植株分别用 0、50 和100 mM NaCl 溶液进行处理。叶片相对含水量、叶绿素a、b 含量、CO2 同化速率和气孔导度的降低表明叶绿体受到了盐胁迫的影响。过氧化氢、丙二醛含量及电导率的升高表明细胞受到了伤害。可溶性糖、游离脯氨酸含量及抗氧化酶含量的上升增加了植株耐盐胁迫的能力。在每个处理的重复图像中都能检测到1,000 个以上的蛋白点。其中有38 个盐响应蛋白被成功鉴定,有16 个蛋白(点4、10、11、14、15、21、24、26、27、28、33、34、35、36、37 和38)出现在盐胁迫的植株中;3 个蛋白(点10、11 和35)只出现在重度盐胁迫处理中;而1 个蛋白(点1)只出现在对照处理中。2 个蛋白(点1 和2)表达量下降,其余蛋白点表达量都增加。被鉴定的蛋白一部分参与了生理生化反应,而另一部分则在信号传导、蛋白质合成等方面有重要作用。盐胁迫下的生理生化变化及蛋白质组学的联合研究有利于青杨对盐胁迫的适应性分析。 Soil is the indispensable environment for human survival and important resource for agriculture development. Nowadays soil is threatened by drought stress and salt stress. Poplars (Populus spp.) possess some characters such as strong acclimilation, fast growth and great production of biomass. In this study, different species of Populus section Tacamahaca spach were used as model plants to investigate the ecophysiological and proteomic responses to drought stress and salt stress. Our results can provide theoretical evidence 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 ecosystems contaminated by salinity. The results are as follows: 1 Adaptive responses to progressive drought stress in two contrasting poplar species originating from different altitudes Cuttings of Populus kangdingensis C. Wang et Tung and Populus cathayana Rehd., originating from high and low altitudes in the eastern Himalaya, respectively, were examined during one growing season in a greenhouse to determine the effects of progressive drought stress. The results manifested that the adaptive responses to progressive drought stress were different in these two species from different altitudes. Significant changes in height increment, leaf development, relative water content (RWC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) appeared earlier in P. cathayana than in P. kangdingensis, whereas changes in soluble protein, soluble sugar, free proline and antioxidant enzymes appeared earlier in P. kangdingensis. In addition, changes in these parameters became more and more significant when the drought stress progressed, especially under severe drought stress in P. cathayana. Compared with P. cathayana, P. kangdingensis was able to maintain a superior height increase and leaf development under drought stress. Also, P. kangdingensis possessed greater increments in soluble protein, soluble sugar, free proline and antioxidant enzymes, but lower increments in MDA and H2O2 than did P. cathayana when the cuttings were exposed to progressive drought stress. Our results suggest that P. kangdingensis originating from the high altitude has a better drought tolerance than does P. cathayana originating from the low altitude. Furthermore, this study manifested that acclimation to drought stress are related the rapidity, severity, duration of the drought event and the altitude of two contrasting species. 2 Proteomic responses to drought stress in two contrasting poplar species originating from different altitudes The cuttings from a female clone of P. kangdingensis and P. cathayana were used to determine proteomic response to drought stress, respectively. Total proteins of the leaves were extracted by a combination of TCA-acetone and phenol, and separated by two-dimensional gel electrophoresis. More than 1,000 protein spots were reproducibly detected on each gel. 58 differentially expressed spots were detected under drought stress in P. cathayana and 22 drought-responsive proteins were identified by peptide mass fingerprint. 69 differentially expressed spots were detected under drought stress in P. kangdingensiss and 25 drought-responsive proteins were identified by peptide mass fingerprint. The identified proteins are involved in several processes, i.e., signal transduction, protein processing, redox homeostasis, CO2 fixation and energy metabolism. Although the proteins identified in this investigation represent only a very small part of the poplar leaf proteins, some of the novel drought-responsive proteins identified here may be involved in the establishment of homeostasis in response to drought stress in the woody plants. 3 Responses to salt stress in P. cathayana Cuttings from a female clone of P. cathayana were treated by Hoagland’s solution: 0, 50, 100 mM NaCl, respectively. Salinity significantly decreased the relative water content of leaves, the contents of chlorophyll a and chlorophyll b, CO2 assimilation rate (A) and stomatal conductance (gs) in both salt stress treatments,which suggested the chloroplast was affected by salt stress. The observed increases of H2O2 and malondialdehyde contents and electrolyte leakage suggested that salinity caused cellular damage, whereas the increases in compatible solutes and in the activities of antioxidant enzymes enhanced the salt tolerance. More than 1,000 protein spots were reproducibly detected on each gel, and 38 salt-responsive proteins were successfully identified by peptide mass fingerprint (PMF). 16 spots (spot 4, 10, 11, 14, 15, 21, 24, 26, 27, 28, 33, 34, 35, 36, 37 and 38) absent in the control sample were induced by the salt treatment, and three spots (spot 10,11 and 35) were present only in the severely salt-stressed treatment. The %vol of the differentially expressed proteins generally increased with progressing salt stress, except for the decreased %vol of two proteins (spot 1 and 2) under salt stress and the presence of spot 1 only in the control sample. Some of the novel salt-responsive proteins identified here may be involved in physiological, biochemical response to salt stress in P. cathayana, the other identified proteins play a role in numerous cellular functions, including signal transduction and protein processing. An integrated physiological, biochemical and proteomic approach was used here to systematically investigate salt acclimation in poplar.
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繁殖更新是植物生活史的重要阶段,在退化生态系统中,植物繁殖更新能力往往较差,是植被恢复的限制环节,因而也成为恢复研究重点和核心。本研究选择岷江干旱河谷广泛分布的三种蔷薇:多苞蔷薇(R. multibracteata)、黄蔷薇(R. hugonis)和川滇蔷薇(R. soulieana)为研究对象,通过野外调查,在查明其生长、繁殖更新状况的基础上,采用控制和模拟实验,对种子和幼苗阶段进行了深入研究,综合分析更新潜力,并提出相对应的促进更新和植被恢复措施。主要结论如下: 1)三种蔷薇在岷江干旱河谷广泛分布,生长和繁殖状况良好,结实量大。各生长指标:株高、基径和冠幅,繁殖指标:结实数量、重量和单果重量都具有显著的空间差异性。基径对多苞蔷薇结实量影响最大;而冠幅对黄蔷薇结实量影响最大。海拔和纬度是对蔷薇生长和繁殖影响最大的环境因素,随着海拔和纬度的升高,植株生长更高大,结实量增加;坡度和坡向对其生长和繁殖也有一定影响,随着坡度 和坡向增加,蔷薇生长和结实受到抑制。 2)三种蔷薇在岷江干旱河谷更新现状不佳, 但更新潜力大。活力种子比率低,动物取食以及两年生幼苗的大量死亡是蔷薇更新的主要限制因素。多苞蔷薇和黄蔷薇的结实率低,川滇蔷薇较高。三种蔷薇种子产量大,但种子质量较差,更新具有充足的种源。三种蔷薇都能形成持久种子库,种子库中种子总量大,但有效种子少,黄蔷薇被动物啃食的比例很高,多苞蔷薇和川滇蔷薇也有一部分种子受到动物破坏。三种蔷薇幼苗库组成特征表现为,当年生幼苗所占比例很高,年龄较大幼苗所占比例小。 3)三种蔷薇都具有不同程度休眠,未经处理种子的发芽率极低。黄蔷薇休眠程度最深,为深度生理休眠;多苞蔷薇为中度生理休眠;川滇蔷薇为非深度生理休眠。三种蔷薇种子在形态上发育成熟,种皮具有透水性。蔷薇果果肉和瘦果中含有抑制物质,其浸泡液抑制了油菜种子萌发,果肉抑制作用更强,果肉和瘦果浸泡液的抑制程度分别为:川滇蔷薇>黄蔷薇>多苞蔷薇。切割和硫酸腐蚀提高了川滇蔷薇种子的发芽能力,而对多苞蔷薇和黄蔷薇没有影响。完全去除瘦果果皮和种皮提高了多苞蔷薇种子发芽率,但对黄蔷薇没有影响。赤霉素和烟水对蔷薇种子萌发没有促进作用。三种蔷薇打破休眠所需低温层积时间分别为:黄蔷薇>多苞蔷薇>川滇蔷薇。对于多苞蔷薇和川滇蔷薇,层积前对种子进行硫酸腐蚀或暖温层积能缩短低温层积时间,提高发芽率。对于多苞蔷薇,变温层积中暖温层积和低温层积具有一定的负补性,即延长暖温层积可以缩短种子萌发对低温层积的需要。 4)多苞蔷薇种子形态特征和种子休眠与萌发在不同海拔梯度间存在较大差异。种子采集时间、采集季节和干藏影响多苞蔷薇和川滇蔷薇的种子休眠。多苞蔷薇果实大小、种子大小和千粒重、种皮厚度随海拔升高而增加,而种子饱满率和活力随海拔升高而降低,种子休眠程度也随海拔升高而增加。种皮厚度与种子大小、千粒重成正相关关系,硫酸腐蚀后的种子经过不同时间的低温层积后,种子发芽率与种皮厚度、种子大小、千粒重、海拔成正相关关系。2006 年采集川滇蔷薇和多苞蔷薇种子休眠程度较2005 年低。种子休眠随种子年龄增加而减弱。高温和干旱能减轻多苞蔷薇和川滇蔷薇种子休眠。 5)三种蔷薇的生长和生物量积累在干旱胁迫条件下受到抑制,而生物量分配、叶片形态特征和水分利用特征等都发生了变化。三种蔷薇的根、茎、叶各器官生物量以及总生物量等在干旱胁迫下明显减小,叶片脱落数量增加。在干旱胁迫条件下,较多的生物量分配到地下部分,从而这使R/S 明显增加。比叶面积(SLA)和冠层面积比(LAR)对干旱胁迫的反应不敏感,仅有部分物种在干旱胁迫条件下发生了变化,并且其变化特点在不同年龄幼苗之间有一定差异。干旱胁迫对WUE 的影响在不同物种间存在差异。多苞蔷薇和黄蔷薇的WUE 随着干旱胁迫的增加而增大, 而川滇蔷薇的WUE 则随干旱胁迫增加而减小。在干旱胁迫条件下,多苞蔷薇和黄蔷薇叶片脱落量和生物量减小幅度较川滇蔷薇大,表明其抗旱能力较强。在干旱胁迫条件,三种蔷薇两年生幼苗的生物量减小幅度较当年生幼苗小,表明两年生幼苗的抗旱能力更强。 6)两种植被恢复措施中,幼苗移栽比播种具有更好的植被恢复效果。播种后,蔷薇种子的发芽率较高,但出苗率都很低,即使出苗,幼苗也几乎在一月内全部死亡。 三种微生境条件下(灌木、半灌木和裸地),种子出苗和幼苗成活没有差异。移栽幼苗总体死亡率都比较低,小于20%。特别是两年生幼苗死亡率更低,小于2%。移栽后的幼苗生长状况良好,在整个生长季中,各生长指标不断增加。生境对幼苗的存活率没有显著影响,但对于幼苗的生长和生物量积累有一定影响,裸地更有利于幼苗生长和生物量积累。与当年生幼苗相比,两年生幼苗具有更高的成活率。总之,三种蔷薇在干旱河谷分布广泛、生长繁殖状况良好,结实量大,具有丰富种源,繁殖更新潜力大,但繁殖更新状况不佳;种子散布后动物对种子的取食、种子的深度休眠过程、种子出苗以及当年生幼苗的存活和定居是更新的主要限制环节。水分是影响结实、种子休眠解除和萌发,幼苗存活和定居的最主要的限制因素。在植被恢复中,应在种子成熟季节大量采集种子,在室内打破休眠后进行人工播种,培育两年生幼苗,通过幼苗移栽方式进行植被恢复。川滇蔷薇应栽种在相对湿润的过渡区,而多苞蔷薇和黄蔷薇可以应用于核心区植被恢复。 Regeneration is an important phase in plant life cycle. It has been a key component of ecological restoration in degradation ecosystem in which plants commonly has poor regeneration. In this paper, we investigated the natural growth, propagation and regeneration status of native three rose species, Rosa multibracteata, R. hugonis and R. soulieana, and analysis the limitation in seed germination and seedling establishment stages. Advice on facilitating the use of these plants in restoration based on the results has been proposed. The results were as follows: 1) Three rose plants widely distributed in the dry valley of the Minjiang River, and made a good performance in growth and propagation. There were significant spatial differences in each growth parameter, such as ramet height, basal diameter, crown diameter and propagation parameters including hip number of a clump, hip mass of a clump and a hip mass. Basa diameter was the most important growth parameter influencing fruit number for R. multibracteata and crown diameter was for R. hugoni. Altitude and latitude had the greatest effect on the growth and propagation of rose plants among environmental conditions. Each parameter of growth and propagation increased with the increase of altitude and latitude. In addition, the increase of slope and aspect limited the growth and propagation. 2) Three rose plants had poor natural regeneration, but great regeneration potential. Low seed viability, predation and higher mortality of current year old seedlings were the limitation in regeneration. R. multibracteata and R. hugonis had higher fruiting rates than R. souliean. All three plants produced a great number of seeds, while their viability was poor. Three rose plants had persistent seed banks, with high total seed number but very low viable seed density. Predation was most severe in R. hugonis, and it also existed to some degree in R. multibracteata and R. soulieana. The seedling age-structure was characteristic of current-year seedlings predominating and few older seedlings were observed. 3) Three rose seeds were dormant and untreated seeds germinated with very low germination percentages. The rose seeds had morphological mature embryos, and achenes were permeable. Some inhabit substances existed in hips and achenes for the extracts of hips and achenes inhibited germination of Brassica campestris. The inhibition effect of the extracts of three rose hip and achenes was R. soulieana>R. hugonis>R. multibracteata. Mechanical and H2SO4 scarification increased R. soulieana germination but had no effect on germination of R. hugonis and R. multibracteata seeds. Full removal of pericarp and testa improved the germination of R. multibracteata but did not affect R. hugonis germination. GA3 and smoke water had no positive effect on rose seed germination. The periods of cold stratification required to released seed dormancy was R. hugonis > R. soulieana >R. multibracteata. H2SO4 scarification and warm stratification shortened cold stratification to release dormancy for R. soulieana and R. multibracteata. Warm stratification had complementary effect for cold stratification, i.e. the longer warm stratification seeds received, the shorter cold stratification were required to obtain the same germination percentage. Three rose seeds had different kinds of dormancy; R. hugonis has deep physiological dormancy, R. multibracteata with intermediate physiological dormancy and R. souliean non-deep physiological dormancy. 4)The seeds traits and dormancy of R. multibracteata showed significant difference across altitudes. Year and season of seed collection had significant effect on seed dormancy for both R. souliean and R. multibracteata. Hip size, seed size, seed weight, seed coat thickness and seed dormancy level increased with the increase of the altitude. There were positive relations between seed coat thickness with seed size and seed weight. Germination percentage of seeds treated with H2SO4 scarification following different periods of cold stratification showed positive relation with seed coat thickness, seed size, seed weight and altitude. Seeds of R. souliean and R. multibracteata collected in 2006 had low dormancy level than those collected in 2005. Seed dormancy decreased with increasing seeds age. High temperature and drought were associated with low dormancy level. 5) Seedling growth, the total dry mass and their components of seedlings were reduced, while leaf senescence accelerated under drought stress. More biomass allocation to root system resulted in higher R/S ratio under drought. Water-use efficiency (WUE) of R. multibracteata and R. hugonis increased, while it declined for R. soulieana under drought stress. R. soulieana seedlings had poor drought-resistance capacity it had more senescent leaves, and its reduction of biomass was stronger than two other rose plants under drought. The reduction degree of one year old seedlings under drought stress was slighter than that of current year seedlings. Therefore, one year old seedling was more drought-resistent compared to current year seedlings. 6)Planting seedlings may have better effect in comparison with direct seeding. Most seeds germinated after seeding, but seedling emergence was very low. More than 80 % seedlings from direct seeding died within a months after emergence. Seedling emergence and survival rate did not show difference among microhabitats. Mortality rates of seedlings artificially planted in microhabitats were general lower than 20 %, and the mortality rate of one year old seedlings was lower than 2 %. Each grow parameter including plant height, leaf number and branch number continually increased after planting. Microhabitat type had effect on the growth parameter and biomass production, but it did not influence the seedling survival. Bare land tended to facilitate seedling growth. One year old seedlings had higher survival rate than current year seedlings. In conculsion, the three rose had wide distribution in the dry valley of the Minjiang River. They produced many seeds and had tolerance to drought stress to some degree. But they had poor regeneration in habitats may be caused by predation, seed dormancy,and high mortality in current year seedlings. We recommend that rose plants should be utilized in restoration by planting two-year old seedlings in spring. A large quantity of seeds should be collected artificially in autumn, release seed dormancy in room, and then cultivate two-year old seedlings by seeding in particular container. R. soulieana seedling probably be planted in transition area, and R. multibracteata and R. hugonis can be used in core area of the dry valley of the Minjiang River.
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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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本研究通过我国CDBI、 KUN、PE、SZ等主要标本馆约3, 500份馆藏标本的研究和野外考察相结合,对我国蔷薇属(Rosa L.)芹叶组(Sect. pimpinellifoliae DC. ex Ser.)植物以及相关组的一些种进行了性状特征、形态和微形态的研究,对该组的一些种的形态特征描述进行了补充,同时给出详细的地理和海拔范围分布图。综合花粉以及种子(瘦果)形态的研究结果重新制订了分种检索表,同时,对该组一些形态相近容易混淆的种进行了对比研究,特别对一直存在争议的绢毛复合体(绢毛蔷薇R. sericea Lindl.和峨眉蔷薇R. omeiensis Rolfe)进行了大量宏观形态特征的研究,并用光学显微镜(LM)和扫描电镜(SEM)对二者的花粉及种子形态、微形态进行对比研究和分析,主要研究内容包括: 1. 芹叶组孢粉研究 对芹叶组的10个种及相关的4个组共17个种(18个样品)的植物花粉进行了光镜和扫描电镜观察和比较研究。研究结果表明:蔷薇属植物花粉粒大小为中等偏小,极轴长23.98[21.82(R. graciliflora Rehd. et Wils.)~29.18(R. tsinglingensis Pax. et Hoffm.)] μm,赤道轴长28.65[24.15(R. graciliflora)~34.70(R. davidii Crép.)] μm;花粉属辐射对称等极单花粉,花粉形态赤道面观呈球形到超长球形;极面观为三裂圆形或近圆形,三孔沟,孔缘加厚,具中部突起的桥状盖。花粉外壁纹饰为条纹状,光镜下形态特征相差不大;在电镜下外壁条纹和脊沟内穿孔的形状、大小和频度等特征,常具组至种水平上的可见变异,可作为组至种水平划分的依据。 根据花粉外壁条纹特征及穿孔形状和数目等特征,本研究将这些植物的花粉归为5个类型,并编制了分组检索表。同时,根据条纹状的清晰度,排列方式、条纹形状、穿孔大小及其频度等方面的差异,各有特点,对该组的10个种编制了分种检索表。 2. 芹叶组种子形态研究 应用光学显微镜和扫描电镜对我国蔷薇属芹叶组14个种及相关组5个组共36种植物的种子宏观形态及种皮微形态特征进行了观察研究。结果显示,蔷薇属种子形态多样,形状分别为肾形、卵形或锥形等;种子颜色以淡棕色、褐色以及土黄色为主;种子大小种间相差悬殊,相对体积为(长×宽×厚)36.66(4.79~114.47) mm3。光镜下,种子宏观形态特征具组内一致性,在扫描电镜下种子表面结构特征因种而异,其纹饰以网纹为主,可分为3种类型,即近平滑型、负网纹型和网纹型。研究结果表明,蔷薇属种子表面纹饰与地理分布关系不大,具有组及种内稳定性。其种子形态、大小、表而纹饰类型等特征可作为蔷薇属组及种水平上的分类依据。 结合蔷薇属花粉形态研究结果,得出蔷薇属种皮微形态特征与花粉外壁纹饰特征相吻合,在代表组及种的特征上具相关性的结论。同时根据种子形态、微形态结构特征的组间区别和种间差异编制了分组及芹叶组14个种的分种检索表。 3. 绢毛蔷薇复合体的研究 通过对大量标本的研究、野外观察以及扫描电镜对绢毛蔷薇复合体的花粉形态和种皮表面结构进行研究,通过对小叶、花粉及种子的形态定量分析结果支持Rowley (1959)的观点,将峨眉蔷薇处理为绢毛蔷薇的一个变种。 综上研究结果得出,蔷薇属植物的小叶片数目、花被基数以及花粉及种子形态等性状是较为稳定的,这些特征可很好的作为分类学依据。 The morphology, pollen exine sculpture and seed coat structure of the species of Rosa sect. Pimpinellifoliae and related sections were studied.About 3,500 herbarium specimens at CDBI, KUN, PE, and SZ were examined. Field work in Sichuan and Yunnan were conducted. Revisions of some species were carried out and a new key to species of sect. Pimpinellifoliae was proposed based on morphology, pollen exine sculpture and seed coat structure, Detailed morphological descriptions, geographical distributions and the altitudinal ranges of some taxa are given. The systematics of the species complex, the Rosa sericea complex (R. sericea Lindl. & R. omeiensis Rolfe), was emphasized. This thesis focused on the following three aspects: 1. Pollen morphology of Rosa sect. Pimpinellifoliae The pollen morphology of 18 samples representing 10 species of the Eurasian Rosa sect. Pimpinellifoliae and 7 additional species of related sections was investigated under LM and SEM. The pollen grains are monadic, actinomorphic, equipolar, medium-sized, spheroidal to perprolate in equatorial view, 3-lobed circular or semi-circular in polar view, crassimarginate, pontoperculate, and with striate exine sculpture. The striate sculpture varies among sections and species. The equatorial axis ranges from 17.97 μm (R. sikangensis) to 29.18 μm (R. tsinglingensis) with an average of 23.98 μm in length, while polar axis varies from 24.15 μm (R. gracilifolra) to 34.70 μm (R. davidii) with an average of 28.65 μm in length. The pollens can be divided into five types based on striate sculpture and a key to the sections sampled was proposed accordingly. The pollen morphology of species of sect. Pimpinellifoliae is more homogeneous and different from other sections sampled and did not support the two-series subdivisions in sect. Pimpinellifoliae. A key is also provided based on characers of pollen morphology among species in sect. Pimpinellifoliae. 2. Seed coat structure of Rosa sect. Pimpinellifoliae The seed coat structure of 39 samples representing 14 species of Rosa sect. Pimpinellifoliae and 12 additional species of related sections was investigated under LE and SEM. The seed relative volume (Length × width × thickness) ranges from 4.79 to 114.47 mm3 with an average of 36.66. mm3. The seeds are reniform, ovate or oblong in shape, with orange-brown, light brown or deep brown color. Seed coat sculpture was reticulate or striate-like reticulate. There was no difference in sculpture character of various speices under LM, while three types of seed coat sculpture were identified under SEM and a key to species based on the seed coat sculpture was provided. The three types of seed coat sculpture were nearly smooth, areolate and reticulate. The study of the seed coat sculpture of same species sampled from different populations showed that characters on the seed coat are stable, and thus the size, shape and seed coat sculpture can be used in species level identification. Interestingly, characters in the seed coat sculpture and the pollen morphology in sect. Pimpinellifoliae are consistent at in specific or sectional levels. A key to the 14 species sampled was given based on seed coat sculpture. 3. The study on Rosa sericea complex The Rosa sericea complex contains R. omeiensis and R. sericea. They are morphologically similar to one another and the systematic status of R. omeiensis has been controversial. In this study we examined large numbers of herbarium specimens of R. omeiensis and R. sericea and conducted field observations in the Hengduan Mts.. We also performed SEM study of pollen morphology and seed coat structure of R. omeiensis and R. sericea. We further carried out intensive morphometric study on the leaflet, pollen, and seed morphology. Our results showed that R. omeiensis should be sunk to be a variety of R. sericea, just as Rowley’s treatment in 1959. In conclusion, the features in the number of leaflet and petal, and the morphological character on pollen and seed are relatively stable. Therefore these characters are very useful in taxon delimition.
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九寨沟自然保护区地处四川盆地北部岷山山脉南段,属川西高原气候中的暖温带。保护区内分布着118 个串珠状排列的高山湖泊,核心景区共有湖泊20 个(组),集中在海拔2000 米到3000 米之间。水资源是九寨沟的灵魂,保护水生生态系统是九寨沟生态旅游可持续发展的立足之本。本研究以九寨沟湿地生物多样性保护项目为依托,于2007 年5 月~2008 年4 月对九寨沟核心景区湖泊水体硅藻植物进行调查,分析了其密度与环境因子的相互关系,旨在为九寨沟湖泊生态系统保护提供科学依据和基础资料。主要有以下几点结论: (1)九寨沟硅藻植物丰富,调查统计为2 纲6 目10 科35 属159 种(包括变种和变型),水体硅藻中真性浮游种类较少,着生种类多,羽纹纲的脆杆藻属、舟形藻属、桥弯藻属、异极藻属、曲壳藻属的种类最多,常为优势属组成湖泊硅藻群落,只在长海是小环藻属占绝对优势。 (2)九寨沟水体中硅藻细胞密度不大,多在4×104~30×104 个/L,长海最高,五花海最低。在年变化上,不同的湖泊水体中硅藻的生长高峰季节不尽相同,在夏季、秋末和冬末春初均有可能出现高密度。在空间上,整体看来密度与海拔有一定的正相关,但由于水温不是单一主导因素,密度受多种环境因子影响,在海拔梯度上呈现一定的波浪状变化。 (3)影响九寨沟湖泊水体硅藻细胞密度较重要的环境因子是海拔、水温和pH。硅藻细胞密度年变化幅度大的湖泊水体,年平均细胞密度大。因此,环境因素不太稳定的湖泊水体,更可能出现高的硅藻细胞密度。11 月份响应于水量剧烈的减少,细齿菱形藻大量出现在五彩池成为优势种群,造成水体硅藻细胞密度的大幅增加。 (4)九寨沟湖泊水体水质几乎达到国家地表水I 类标准,但是,硅藻商指数以及代表多样性的Cairns 指数计算的结果不能反映水质的情况,这是由于水质状况不是决定九寨沟湖泊水体中硅藻生存的种类和数量的主要因素。因此在用各种指数分析水质的时候,都应该先判断计算该指数的参数是否主要由水质状况决定,然后再综合理化性质等其他指标分析和比较。 (5)人类的活动对九寨沟湖泊水生态系统有一定的影响。一方面,在低海拔景点人类活动的干扰比海拔高处相对要大,干扰引起低海拔景点湖泊硅藻Cairns指数较高;另一方面,旺季的影响比淡季大,水体中出现了典型富营养种类。因此,应加强低海拔景点和旺季的管理,并限制淡季的人类活动以保证九寨沟生态系统的自我恢复。 Jiuzhaigou nature reserve is located in north Sichuan, at the south section of Minshan Mountain, having a warm temperate climate which belongs to West Sichuan plateau climate. It includes 118 mountain lakes, and 20 of them between 2000 and 3000 altitude degree constitute the core area of Jiuzhaigou. The water is Jiuzhaigou's soul, so it is the basis of sustainable tourism development of Jiuzhaigou to protect the water ecological environment. Research depends on Jiuzhaigou wetland biodiversity protection program in this paper, invested the diatom plants in 17 lakes at the core area from May 2007 to April 2008. The composition and the correlation of diatom density and the environment were studied, which could provide scientific and basic information for the protection of the ecological environment of Jiuzhaigou. The main conclusions were: 1 The diatom species were in abundance, investigation showed that there were 159 species belong to 10 families and 35 genera. Most diatoms were benthos, euplankton were less. Genera Fragilaria, Navicula, Cymbella, Gomphonema and Achnanthes had the most species, which always were the dominant genera. Only in The Long Lake genus Cyclotella possessed a predominant status. 2 The diatom cell density was not very high, always between 4×104 cell/L and 30×104 cell/L, highest in The Long Lake and lowest in The Colorful Lake. In one year's time, diatom in different lakes had different growth type. The maximum could be in summer, late autumn and late winter early spring. The cell density showed a positive correlation to altitude in the whole, but fluctuated in details along altitude because of comprehensive influence by many environmental factors. 3 Altitude, water temperature and pH were the more important factors to determine the cell density. There was a positive correlation between the range of cell density in one year and the average cell density. So high annual cell average density probably will appear in lakes which has an unstable environmental condition. The boom of Denticula elegans in The Jade-Colored Lake in November was corresponding to the large decrease in water quantity. 4 The water quality of Jiuzhaigou was near the National Surface Water Standard rank I, but the Diatom Quotient and the Cairns Index didn't reflect the water quality accordingly. It was because the water quality wasn't the most important factor here which determined the species number and species quantity, so the index based on these items would not reflect it. Therefore, it is recommended that, when we use indices to evaluate the water quality, we should know first whether the water quality is the main determinative factor inflects the items used to compute the indices, and then combined with other chemical indices to analyze and do comparison. 5 The tourists' activities had influenced the water ecosystem of Jiuzhaigou. On the one hand, disturbance was relatively high at the lower altitude, it resulted in higher Cairns index; on the other hand, influence in high season was more severe than in low season, some typical species indicating eutrophication were observed in the water. It is suggested that the management at lower altitude and in high season must be strengthen, and less people activity in low season is necessary to fulfill the recovery of the water ecosystem.
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全球气候变化已经成为不争的事实,其中全球变暖是近年来国内外的研究热点之一。土壤碳库作为陆地生态系统最大的碳库,气温升高必然会导致一系列的土壤碳储量和碳通量的变化,这些微小的变化又可能导致大气CO2浓度的变化并强化这种变暖的趋势。目前,土壤碳循环对温度升高的响应仍然是陆地碳循环研究最缺乏的部分,对土壤有机碳动态变化的研究仍存在着很大的不确定性与争议。四川西部的亚高山人工针叶林是青藏高原东部高寒林区的重要组成部分,是研究全球变化对森林生态系统影响的关键地区和重要森林类型。本研究通过采用原位人工模拟增温装置(Open-top chambers,OTCs)对川西米亚罗60年人工云杉林土壤实施增温,研究高海拔地区森林,尤其是人工森林系统下的土壤有机碳 含量、土壤呼吸及土壤酶活性对温度升高的响应。结果表明: 1. 增温处理的660天(2005年11月至2007年9月)期间,增温条件下的平均气温和土壤平均温度分别比对照提高0.43 ℃和0.27 ℃;0~10 cm土壤含水量在增温的不同时期均有不同程度的降低。 2. 土壤蔗糖酶、蛋白酶和脲酶活性在温度升高的不同阶段均有不同程度的提高。在增温处理300天(2006.09)、540天(2007.05)、600天(2007.07)和660天(2007.05)后,0~10 cm层的蔗糖酶活性分别比对照提高了36.36%(P<0.05)、24.31%、14.54%(P<0.05)和7.22%,脲酶活性分别提高了12.90%、24.19%(P<0.01)、34.48%(P<0.05)和14.64%(P<0.05),蛋白酶活性分别提高了31.37%、1.99%、3.70%和17.80%。10~20 cm层的土壤酶活性也均有不同程度的提高,但均没有显著差异。蔗糖酶、脲酶和蛋白酶活性均呈现出随土层加深而减弱的趋势。 3. 土壤过氧化氢酶和多酚氧化酶活性在增温的第1年内均有不同程度的提高,但在增温的第2年内比对照有所降低。增温300天后(2006.09),过氧化氢酶和多酚氧化酶在0~10 cm层分别比对照增加3.76%和49.25%(P<0.05),10~20 cm层分别增加了5.54%和29.67%。在增温的第2年内,增温540天(2007.05)、600天(2007.07)和660天(2007.09)后,0~10 cm层的过氧化氢酶活性分别比对照降低了27.70%(P<0.05)、4.34%和1.47%,多酚氧化酶活性分别降低了5.86%、11.76%(P<0.05)和7.47%。增温的第2年内,10~20 cm层的过氧化氢酶和多酚氧化酶活性也均有不同程度的降低,但差异均未达到显著水平。不同土层之间相比较,过氧化氢酶活性随土层加深而降低,多酚氧化酶活性随土层加深而增加。 4. 土壤有机碳和有机质在增温的不同阶段,含量比对照均有所降低;且随增温时间的延长,降低的幅度下降。0~10 cm层的土壤有机碳和土壤有机质在增温300天(2006.09)、540天(2007.05)、600天(2007.07)和660天(2007.09)后分别降低了8.69%、4.35%、3.80%和2.44%,差异均未达到显著水平。土壤全氮含量在增温后与对照相比无明显的增加或者降低趋势。增温条件下的土壤C/N比与对照相比有所降低,但在增温各阶段的差异均不显著。10~20 cm层的有机碳、有机质和C/N比也有不同程度的降低趋势,但差异均不显著。不同土层之间相比,0~10 cm层的有机碳、有机质、全氮含量和C/N比均高于10~20 cm层,呈现出随土层加深而降低的趋势。 5. 土壤呼吸速率在增温第1年内,与对照相比明显提高,但在增温处理2年后,与对照相比无显著变化。增温300天(2006.09)和360天(2006.11)后分别提高了13.32%和21.17%,差异显著。增温处理540天(2007.05)到660天(2007.09)期间,与对照相比,不仅没有明显的提升,反而有些月份比对照有所降低,对温度升高的敏感性降低,呈现出对温度升高的适应性。土壤呼吸的日呼吸速率呈现单峰曲线形式,在14:00~20:00期间达到最大值,在4:00~10:00期间具有最低值。土壤呼吸的季节变化,呈现出与外界环境温度相一致的趋势,在7月份(夏季) 最高,11月份(冬季)最低。土壤呼吸与2 cm土壤温度、5 cm土壤温度和空气温度均呈极显著指数相关,与0~10 cm土壤含水量呈线性相关,相关性达到显著水平,但低于土壤呼吸与温度的相关性。 The past century has seen a marked increase in atmospheric carbon dioxide concentrations and a concomitant warming that has drawn scientific attention to the link between global carbon stocks and climate change. In particular, the decomposition and turnover of soil organic matter is recognised as an important determinant of carbon driven climate change. The slightly variation in soil organic carbon will result in the increase of atmospheric carbon dioxide concentrations and reinforce the tendency of warming. The experiment was conducted in Subalpine coniferous forest in western Sichuan province. Subalpine coniferous forest in western Sichuan was a important part of eastern Qinghai-Tibetan Plateau, which play a important role in reseaching the sensitivity of forest ecosystem to climate change. To investigate the effects of elevated temperature on soil organic carbon content, soil respiration rates, and soil enzyme activities in subalpine Picea asperata plantations, a esimulated warming measure was applied with Open-top chambers. The results were as followed: 1) During the period from Nov. 2005 to Sep. 2007, mean air temperature and soil temperature were respectively 0.43℃ and 0.27℃ the ambient higher. Soil water content decreased to different exent in different months in warmed plots than in unwarned plots at depth of 0-10 cm. 2) In general, elevated temperature enhanced the soil enzyme activities of invertase, protease, and urease. In the first year of warming—after 300 days’ treatment (in Sep,2006), the activities of invertase, protease, and urease increased by 36.36%, 12.90% and 31.37% respectively at the depths of 0-10 cm,among which the activity of invertase reached statistic significance. In the second year of warming, invertase activity increased by 24.31% after 540 days’ treament (in May, 2007), 14.54% after 600 days’ treament (in Jul, 2007) and 7.22% after 660 days’ treatment (in Sep, 2007) at the depths of 0-10 cm, and the differences in July and Septemmber were statistically significant. Elveated temperature also increased the activity of urease in the second year of warming and had significant effects in May and July. The activity of protease in warmed plots was also higher than in unwarmed plots at depths of 0-10 cm, but there was no significant difference. Elevated temperature had no significant effects on all soil enzyme acitivities at the depths of 10-20 cm in the first and sencond year. The values of above-mentioned soil enzyme all decreased with soil layers. 3) Eleavted temperature enhanced the activities of catalase and polyphenol oxidase in the first year of warming while they turned out downtrend in the second year. The activity of catalase increased by 3.76% and 5.54% at depths of 0-10 cm and 10-20 cm respectively in the first year—after 300 days’ warming (in Sep, 2006), the differences of which had no statistical significance. The activity of polyphenol oxidase was significantly increased by 49.25% at depths of 0-10 cm and not significantly increased by 29.67% at depths of 10-20 cm after 300 days’ warming. In the second year of warming, the catalase activity was significantly decreased by 27.70% after 540 days’ treament (in May, 2007) and not significantly decreased by 4.34% and 1.47% after 600 days’ (in Jul, 2007) and 660 days’ treament (in Sep, 2007) respectively. The activities of catalase and polyphenol oxidase at depths of 10-20 cm were decreased to different extent, but there was no significant difference. Catalase activity stepped down with soil layers while polyphenol oxidase activity stepped up. 4) Increased temperature in both the first year and the second year resulted tendency of decrease in the contents of soil organic carbon and soil organic matter, and C/N ratios at soil depths of 0-10 cm and 10-20 cm. However, with the prolonged warming, the tendency of decrease gradually tapered off and the extent of decrease in the second year of experiment were lower than that in the first year. The contents of soil organic carbon and soil organic matter were all decreased 8.69% by warming in the first year and dcreased 4.35%, 3.80% and 2.44% in May, July and September of the second year, but no significant difference were found. The C/N ratios increased 8.52% in the first year of warming and had less increment in the second year, all of which were not statistical significant. Eleveated temperature had no obvious effect on the content of tatol N in two year consecutive warming experiment. The contents of soil organic carbon and soil organic matter, total N and C/N ratios all had the tendency of dcreasing with soil layers. 5) Soil respiration rates were significantly enhanced by 13.32% and 21.17% after 300 days’ (in Sep, 2006) and 360 days’ (in Nov, 2006) treament in the first year of warming, but the same showed no obvious difference in the second year of treatment, which was assumed the adaptability of soil respiration with a certain heightened temperature. Diurnal soil resspiration showed a daily variation with a minimum value between 4:00 and 10:00 h and a maximum value between 14:00 and 20:00 h, coinciding with the minimum and maximum values of soil temperature at 2 cm. Soil respiration rates exhibited a pronounced seasonal variation with minimum values in Novmber and a maximum value in July, approximately coinciding with the seasonal variation of air and soil temperature. An exponential function provided the best fit for soil respiration with temperature while a quadric equation was used to estimate the effect of soil moisture on soil respiration, which were all significantly correlated. Soil respiraion rate was more highly correlated with the soil temperature than soil moisture.
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横断山区干旱河谷地区的土壤是制约植被恢复的一个关键要素,但干旱条件下土壤质量与动态演变状况并不清楚,本研究以岷江干旱河谷核心地段的土壤为对象,研究了阳坡褐土土壤的物理、化学以及生物学特征在海拔梯度(1800m、2000m、2200m)格局及其时间动态(2005-2007)变化,应用主成分分析综合评价了土壤质量及其时空差异性,明确了土壤质量的变化趋势。主要研究结论如下: 1) 海拔梯度上土壤物理性状的变化,2005 年和2007 年土壤物理性状综合质量随着海拔的升高均得到了优化,即海拔2200 m>海拔2000 m>海拔1800 m。 2) 海拔梯度上土壤化学性质的变化,土壤化学综合性质2005 年随海拔升高而趋于变得更优,而2007 年海拔2000 m 最优,海拔1800 m 则最差。 3) 海拔梯度上土壤生物学性质的变化,2005 年土壤生物学性质随海拔升高表现出趋于更好,2007 年海拔2000 m 最优而海拔1800 m 地段最差。 4) 从土壤物理、化学、生物学三方面出发,应用主成分分析,分别分析得出2005 年和2007 年不同海拔高度的土壤质量综合得分。根据综合得分得出土壤质量综合评价的排列顺序为:2005 海拔2200 m>2007 年海拔2000 m>2005 年海拔2000 m>2007 年海拔1800 m>2007 海拔2200 m>2005 年海拔1800 m。2005年土壤综合质量随海拔升高而趋好,2007 年则以海拔2000 m 最优,海拔1800 m和2200 m 差异不大。 5)排除人为干扰后,干旱河谷土壤物理性状在海拔1800 m 略有恢复,海拔2000 m 变化不明显,而海拔2200 m 仍有退化趋势;土壤化学性质在海拔1800 m和2000 m 地段得到恢复,而海拔2200 m 处仍有退化;土壤生物学性质在海拔2000 m 地段呈恢复趋势,而1800 m 和2200 m 仍处于退化状态。综合质量分析表明,与2005 年相比,2007 年海拔1800 m 和2000 m 地段土壤质量趋于变优而海拔2200 m 地段仍有退化迹象。 Soil is a key factor that affect the restoration of vegetation in the Hengduan Mountains dry valley area. But the dynamics and quality of soil is not knowed in dry area. In this study, soil physiochemical and biological characteristics ranging from 1800~2200m above sea level from a typical south-facing slope at the Minjiang River dry valley area had been studied, and characteristics of changes in soil quality along altitudinal gradients and time scales were also discussed. The principal component analysis was used to assess the soil quality. The main results were as follows: 1) Changes in soil physical properties along altitudes. Soil physical properties obtained the optimization along with the elevation in 2005 and 2007. 2) Changes in soil chemical properties. It was summarized that soil chemical properties increased with elevation in 2005, but the soil of 2000 m was the best in 2007. 3) Changes in soil biological characteristics. Soil biological properties increased with elevation in 2005, but the soil of 2000 m was the best and 1800 m was the worst in 2007. 4) Change tendency of soil quality. With the soil physics, chemistry and biology characteristics, we analysised the change tendency of soil quality in altitudes. The result indicated that soil quality increased with altituded in 2005, and soil quality of 2000 m was the best in 2007. 5) In brief, the soil quality is by physics, chemistry as well as the biology synthesizes the influence the final outcome. And the soil quality's change was manifested by soil physics, chemistry and the biology characteristics. All the results indicated soil quality still degenerated at 2200 m in in the dry valley of Minjiang River. And soil quality of 1800 m and 2000 m resumed slight.
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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.
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植物功能生态学研究不仅提供了植物生理生态学与生态系统生态学的连接,还为植物种群生活史对策研究提供了材料。Westoby 等 (2002) 提出了利用植物功能性状变量的主导维度来确定和量化植物生活史的生态适应策略。在他们所提出四个主导维度中,叶大小-小枝大小是研究相对较少的一维;其内部各组分的关系、对环境的响应,以及与其它重要维度的关系,目前的理解非常有限。 本研究以贡嘎山不同海拔不同功能群物种为研究对象,采用种间比较和系统发生独立性比较等研究方法,系统研究了植物的功能特征及其相关性在不同生境及不同功能群间的差异,旨在分析不同功能群物种的叶大小-小枝大小的成本和收益。其研究结果将有助于我们理解植物生活史对策的进化,进而理解物种共存和维持物种多样性的机制。主要研究结果如下: 1. 叶大小-小枝大小关系 小枝茎横截面积与单叶面积和总叶面积均呈异速生长关系,即总叶面积和单叶面积的增加比茎横截面积的增加速度快。但是,总叶面积和叶片干重的增加却基本上与小枝茎干重的增加等速。系统发生独立性比较研究的结果与此相一致。表明,在某一给定的茎投入时,至少大叶大枝物种不比小叶小枝物种在支撑叶面积和叶片干重方面具有优势。同时,在某一给定的小枝茎投入时,常绿阔叶物种比落叶阔叶物种支撑更少的叶面积。在茎干重与总叶面积的关系中,落叶复叶物种比落叶单叶物种具有更高的y轴截距,表明复叶物种比单叶物种在展叶面积方面更有效。复叶物种与单叶物种相比,通常具有较大的叶大小和小枝大小。 2. 叶大小-叶数量关系 叶大小与数量间在不同的叶片习性、不同的叶片形态以及不同的生境类型的物种间均存在稳定的负的等速生长关系,且这种关系在系统发生独立性比较时依然成立。然而,在某一给定的出叶强度 (单位小枝的叶数量) 时,常绿阔叶物种比落叶物种具有更小的叶面积。而在给定体积基础上的出叶强度时,落叶复叶物种的叶面积显著大于落叶单叶物种,且复叶物种比单叶物种具有更大的叶大小和更小的出叶强度。但是,叶大小与数量间的关系在不同的海拔间并没有显著的差异。 3. 小枝大小-总叶面积关系 在不同的生活型或不同的生境下,小枝上总叶面积与茎干重和小枝干重均呈正的异速生长关系,且斜率显著小于1.0,表明小枝上总叶面积的增加都不能赶上小枝及茎大小的增加。这种“收益递减”表明随着小枝干重的增加,光截取的收益递减。此外,叶面积比 (总叶面积与小枝干重的比值) 与单叶干重呈显著负相关关系,系统发生独立性比较的结果与此相一致。根据以上结果,可以推测,大叶的物种在质量较好的生境中出现,而群落内部小枝茎的寿命较长的物种可以拥有较大的叶片。 4. 叶片色素浓度-LMA关系 随着海拔的升高,阔叶木本植物和草本植物的叶片色素浓度减少,叶绿素a/b和类胡萝卜素/叶绿素比值以及比叶重 (LMA) 增加。然而,在草本植物中的色素浓度、色素比值和LMA的变化比阔叶木本植物的更明显。同时,LMA与叶片色素浓度呈负相关关系,但是在落叶物种中的LMA对色素浓度的影响比常绿阔叶物种更强烈。总之,草本植物的叶片特征对海拔梯度的变化似乎比木本植物更敏感,LMA对叶片色素的保护作用在落叶物种中比在常绿阔叶物种显得更重要。这些结果表明不同生活型物种可能采取不同的保护机制来降低叶绿体器官的损伤和增加他们的碳获取能力。 Studies on plant functional ecology not only bridge plant eco-physiology and ecosystem functioning, but also enrich plant population biology. As pointed out by Westoby et al (2002), plant life history strategies can be identified and quantified by four leading dimensions of variations in plant functional traits, i.e., seed size/output, leaf mass per area and leaf life span, plant height, and leaf size-twig size. Compared to the other dimensions, the cost/benefit of the leaf size-twig size spectrum has scarcely been analyzed in relation to environmental gradients and life form types, and the adaptive significance of this spectrum is not fully understood. In the present study, the relationships between functional traits of plant twigs are determined for the species with different life forms along an altitudinal gradient of Gongga Mountain with both cross-species analysis and evolutionary divergence analysis. The primary objective of this study is to examine the cost/benefit of leaf size-twig size in plants. The study results are supposed to provide insights into the understanding of the mechanism of species coexistences. The results are shown in the following. 1. The relationship between leaf size and twig size Twig cross-sectional area allometrically scaled with both individual leaf area and total leaf area supported by the twigs. However, the increase in total lamina mass/area was generally proportional to the increase in stem mass. These correlations between trait variations were significant in both interspecies analysis and phylogenetically independent comparison (PIC) analysis, which indicated that thick-twigged/large-leaved species, at least, do not have an advantage in supporting leaf/lamina area and lamina mass for the same twig stem investment than thin-twigged/ small-leaved species. Meanwhile, the evergreen broad-leaved species supported a smaller leaf area for the same twig stem investment in terms of both cross-sectional area and stem mass than the deciduous species. The deciduous compound-leaved species have a higher y-intercept in the scaling relationship of twig stem mass versus total leaf area than the deciduous simple-leaved species, indicating that compound-leaved species were more efficient in displaying leaf area. The compound-leaved species were larger in both leaf size and twig size than their counterpart in the present study. 2. The relationship between leaf size and leaf number Significantly negative and isometric scaling relationships between leaf size and leafing intensity (leaf number per twig mass or volume) were found to be consistently conserved across species independent of leaf habit, leaf form and habitat type. The negative correlations between leaf size and leafing intensity were also observed across correlated evolutionary divergences. However, leaf area was smaller in the evergreen broad-leaved species at a given leafing intensity than in the deciduous species. The deciduous compound-leaved deciduous species were higher in leaf area than deciduous simple-laved species at a given volume-based leafing intensity. Moreover, the compound-leaved deciduous species were larger in leaf size but smaller in leafing intensity than their simple counterparts. No significant difference was found in the scaling relationships between altitudes. 3. The relationship between twig size and total leaf area Leaf area was found to scale positively and allometrically with both stem and twig mass (stem mass plus leaf mass) with slopes significantly smaller than 1.0, independent of life form and habitat type, indicating that the increase in total leaf area fails to keep pace with increasing twig size and stem size. This ‘diminishing returns’ suggests that the benefit of light intercept decreased with increasing twig mass. Moreover, the leaf area ratio (the ratio of total leaf area to stem or twig mass) correlated negatively with individual leaf mass. The results of PIC were consistent with the correlations. According to the results, it is speculated that large-leaved species may be favored when habitat is good and when stem longevity are long within community. 4. The relationship between leaf pigment concentrations and leaf mass per area With increasing altitude, the concentrations of pigments decreased, but the ratios of chlorophyll a/b and carotenoid/chlorophyll, and LMA increased, in both the broad-leaved woody species and herbaceous species groups. However, the changes in the pigment concentrations, ratios and LMA were more profound in the herbaceous species than in the woody species. In addition, pigment concentrations were negatively correlated with LMA in each life form type and in the pooled dataset. However, the LMA effect on leaf pigment concentrations was more profound in the deciduous species than in the evergreen braode-leaved species. In general, herbaceous species seemed more sensitive to the increasing altitude compared to woody species, and LMA seemed to be a more important mechanism for protecting leaf pigments in deciduous species than in evergreen broad-leaved species. These results suggested that the species with different life forms may employ different protective mechanisms to decrease the chloroplast apparatus damage and increase their carbon gain.
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雌雄异株植物对环境的不同响应一直是一个有趣而新颖的研究领域,由于雌雄个体不同的繁殖成本及不同的生存策略,使得雌雄植株在生长、存活、生殖格局、空间分布、资源配置等方面已经表现出明显的不同,在生理和分子水平上也表现出明显的性别间差异。干旱是制约农林业发展的环境因子之一,叶锈病是对杨树危害最严重的病害之一,由于长期进化的结果,不同性别的植物必然对生物和非生物胁迫有着不同的响应。本文以雌雄异株的青杨为模式植物,研究雌雄间在生理、生化、亚细胞结构和蛋白质水平上对生物和非生物胁迫的差异响应。主要研究结果如下: (1) 青杨雌雄植株对锈病胁迫的生理生化差异响应 在正常的对照组中,雄株叶片比雌株叶片有着较高的活性氧自由基产生速率、较高的SOD、POD、PPO 和较低的CAT 活性;在锈病感染的早期阶段, SOD、POD、CAT 活性、活性氧自由基产生速率、H2O2 含量、膜脂过氧化程度和细胞膜的电渗率在雌雄株中都增加,而PPO 仅在雄株中增加明显,APX 仅在雌株中增加明显,并且雌株比雄株有着更严重的锈病感染程度、细胞膜的伤害程度和光合系统II 的破坏程度,雌株有更多的净光合速率、气孔导度和叶绿素a 含量的降低,在同工酶变化上,雌雄间对锈病也显示出不同的表达模式。结果显示,雄株比雌株对锈病有着更好的抗性和更有效的ROS 清除系统。 (2) 青杨雌雄植株对干旱胁迫的生理生化及亚细胞结构的差异响应 与较好水分条件相比,干旱下雄株比雌株有着更高的A-Ci 响应参数,如Rubisco 最大羧化速率、光呼吸速率、暗呼吸速率和最大电子传递速率等。干旱显著地增加了膜脂过氧化程度和游离脯氨酸含量,并且雄株比雌株表现出较低的膜脂过氧化程度,较高的总蛋白和游离脯氨酸含量。无论是中度干旱还是极度干旱,除了CAT 外,雄株比雌株表现为较强的抗氧化酶活性,在同工酶谱带上,雌雄间表现出不同的变化模式,并且有些条带是干旱影响应的,而有些条带是性别特异性的,这些性别特异性条带能够作为鉴定性别快速而准确的标记。干旱显著地影响了线粒体、叶绿体和细胞壁的结构,尤其在中度干旱胁迫下,雄株线粒体和叶绿体比雌株呈现出较好的完整性,并且雄株细胞壁要比雌株更厚。因此, 雄株比雌株表现出更强的干旱忍耐性和更高效的抗氧化酶系统。 (3) 青杨雌雄植株对干旱胁迫的蛋白质组差异响应 用双相电泳检测到雌雄间近1000 个蛋白点,通过对比发现对照组雌雄间有54 个差异蛋白点,干旱下雌雄间有108 个差异点,其中102 个被质谱成功鉴定。对照组雌雄间的差异蛋白主要集中在与光合作用相关蛋白、抗氧化酶、胁迫防御蛋白和一些调节基因表达的蛋白;干旱胁迫下雌雄间差异蛋白明显增多,主要有参与信号转导、调节基因表达、蛋白质加工、转录产物的转录翻译后修饰的调节性蛋白蛋白和参与氧化还原平衡、抗胁迫、细胞壁合成、光合作用、能量代谢、氨基酸代谢和脂肪酸代谢等的功能性蛋白。干旱下这些蛋白的表达量在雌雄中有的表现出相同的表达模式,如干旱下雌雄株中Rubisco 激活酶、小热激蛋白等表达都增加,而有的表现出相反的表达模式,如Rubisco 大亚基的降解片段、羰酸酯酶等在雄株中表达量上调而在雌株中却是下调。因此,雌雄间在蛋白质水平上对干旱胁迫响应的差异是显著的,也是复杂的。 It is an interesting and novel topic that dioecious plants possess different responses to environmental stress. As for the different productive cost and different survive strategy, different sexual plants have shown obviously morphological, physiological and molecular differences. Drought is one of the most worldwidely important environmental stress factors that limit plant growth and ecosystem productivity. Rust disease is one of the economically important diseases in many trees. As a result of the long evolutionary process, male and female plants should show different responses to abiotic and biotic stress. In this paper, using a dioeious tree of Populus cathayana Rehd as a model, we study the sexual differences to drought and rust disease stress in physiological, biochemical, sub-cellular and proteomics levels. The main results are follows: (1) The sexual differences in physiology and biochemistry of poplar to rust disease In controls, males showed higher production of superoxide radicals, higher activities of SOD, POD, PPO and lower CAT activity. Under rust disease, the activities of antioxidant, the content of ROS and the degree of cellular member destroyed were increased in both sexes, except for PPO in diseased males and APX in diseased females. However, females showed more seriously disease severity and cellular member and PS II destroyed degrees. Net photosynthesis rate, transpiration rate and chlorophyll a content were decreased more in diseased females than in males. There were also some different changes inantioxidant isozymes under rust disease. The results suggested that male poplar possessed a more effectively antioxidant system and were more resistant to rut disease than females. (2) The sexual differences in physiology and biochemistry of poplar to drought stress Under drought stress, there were higher rates of RuBP-saturated CO2 assimilation, dark respiration, photorespiratory release of oxygen, the max electron transportrate in CO2-saturated and carboxylation efficiency in males than in females. And males showed lower TBARS and higher proline content. Except for CAT, the activities of other antioxidants were higher in males than in females. Meanwhile, there were obviously differences in isozyme changes between teo sexes. Drought stress obviously destroyed the integralities of chloroplasts and mitochondria and the sexual differences in sub-cellular level were obviously under the moderate water stress. Male cell walls were more sensitive to drought stress than did female. The results suggested males were more resistant to drought stress. (3) The sexual differences in proteomics of poplar to drought stress By 2-D and MS analysis, we identified 102 different protein spots between males and females. Under control conditions, the different proteins were mainly in photosynthesis related proteins, antioxidants, stress response proteins and some gene expression related proteins. Under drought stress, the different proteins were focused on (i) regulated proteins such as signaling conduction, kinase, HSP, gene expressional regulation and protein modification, (ii) functional proteins such as photosynthesis, energy metabolism, antioxidant, redox, stress response, lipid metabolism and amino acid metabolism. Some protein showed the same expressional pattern, while some showed contrary expressional pattern. Thus, the results suggested that sexual differences in proteomics were significant and complex.