169 resultados para Contrast ratio
Resumo:
The physics-based parameter: load/unload response ratio (LURR) was proposed to measure the proximity of a strong earthquake, which achieved good results in earthquake prediction. As LURR can be used to describe the damage degree of the focal media qualitatively, there must be a relationship between LURR and damage variable (D) which describes damaged materials quantitatively in damage mechanics. Hence, based on damage mechanics and LURR theory, taking Weibull distribution as the probability distribution function, the relationship between LURR and D is set up and analyzed. This relationship directs LURR applied in damage analysis of materials quantitatively from being qualitative earlier, which not only provides the LURR method with a more solid basis in physics, but may also give a new approach to the damage evaluation of big scale structures and prediction of engineering catastrophic failure. Copyright (c) 2009 John Wiley & Sons, Ltd.
Resumo:
The physics-based parameter: load/unload response ratio (LURR) was proposed to measure the proximity of a strong earthquake, which achieved good results in earthquake prediction. As LURR can be used to describe the damage degree of the focal media qualitatively, there must be a relationship between LURR and damage variable (D) which describes damaged materials quantitatively in damage mechanics. Hence, based on damage mechanics and LURR theory, taking Weibull distribution as the probability distribution function, the relationship between LURR and D is set up and analyzed. This relationship directs LURR applied in damage analysis of materials quantitatively from being qualitative earlier, which not only provides the LURR method with a more solid basis in physics, but may also give a new approach to the damage evaluation of big scale structures and prediction of engineering catastrophic failure. Copyright (c) 2009 John Wiley & Sons, Ltd.
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In present study, the transition of thermocapillary convection from the axisymmetric stationary flow to oscillatory flow in liquid bridges of 5cst silicon oil (aspect ratio 1.0 and 1.6) is investigated in microgravity conditions by the linear instability analysis. The corresponding marginal instability boundary is closely related to the gas/liquid configuration of the liquid bridge noted as volume ratio. With the increasing volume ratio, the marginal instability boundary consists of the increasing branch and the decreasing branch. A gap region exists between the branches where the critical Marangoni number of the corresponding axisymmetric stationary flow increases drastically. Particularly, a unique axisymmetric oscillatory flow (the critical azimuthal wave number is m=0) in the gap region is reported for the liquid bridge of aspect ratio 1.6. Moreover, the energy transfer between the basic state and the disturbance fields of the thermocapillary convection is analyzed at the corresponding critical Marangoni number, which reveals different major sources of the energy transfer for the development of the disturbances in regimes of the increasing branch, the gap region and the decreasing branch, respectively.
Resumo:
何首乌为常用中药,由何首乌及含何首乌的中成药制剂所引起的不良反应也时见报道,科学阐明不良反应的物质基础并提出解决方案对何首乌的使用十分重要。本论文研究了何首乌炮制前后KM小鼠肝脏毒性基因表达谱、生物活性及化学成分的变化。所获结果支持何首乌炮制的目的是减毒、改性(改变药效),何首乌生、熟异治的观点。制首乌对抑郁症的效果显著优于生首乌,这与本草所记载的何首乌炮制后补肝肾、益精血,归肝、肾经一致。 主要结果如下: 1、 生、制首乌的毒理基因芯片研究结果 何首乌的不良反应主要表现在肝损害方面。本研究建立了生何首乌和制何首乌不同剂量的肝毒性作用模型,体重指标统计发现生何首乌各剂量组平均体重显著下降,中剂量组(10 g/kg.d)体重下降20 %,高剂量组(20 g/kg.d)体重下降42%,50%动物死亡,提示动物机体能量代谢障碍;基因芯片研究结果表明何首乌是CYP450的抑制剂,生何首乌相对于制何首乌CYP3A4、CYP4A5显著下调,导致毒性成分在体内的吸收增加,服用大剂量的生何首乌后产生明显的肝毒性;主要对以下六条Pathway产生影响:①PPAR signaling pathway,主要毒性靶基因有RXRB CYP7a1、Acadl、Apoa2、Cyp4a、 FABP2 、MAPKKK5等基因。②Calcium signaling pathway,主要毒性靶基因有CAMK2B、CACNA1F、S100A1、 F2R、Ryr1、Slc8a2、Camk4 ③Neuroactive ligand-receptor interaction,主要毒性靶基因有Chrm4、 Ntsr2 、 GABRR1、 GRIK3、F2R等基因。④Wnt signaling pathway,主要毒性靶基因有Daam2、Rac1 等基因。⑤Complement and coagulation cascades,主要毒性靶基因有F2R、Serpina1b、Cfi 、FGA等基因。⑥Oxidative hosphorylation,主要毒性靶基因有Atp5e、NDUFA1等基因。生何首乌毒性明显强于制首乌,且生何首乌水煎液的毒性大于生何乌首丙酮提取物的毒性,这一结果表明,何首乌主要的毒性成分很可能并不仅仅是传统所认为的以大黄素为代表的蒽醌类化合物,而是何首乌中大量存在的有效组分二苯乙烯苷与大黄素相互作用的结果,这一研究结果与前述的何首乌对肝药酶的影响是一致的。后续生、制首乌的化学成分差异研究表明,炮制后二苯乙烯苷含量明显降低:生首乌为5.512 %、清蒸制首乌为3.811 %、豆制首乌为3.538 %,大黄素的含量炮制后显著升高,生首乌为0.094 %、清蒸制首乌为0.119 %、豆制首乌为0.126 %。 2 生、制首乌药效差异研究结果 本文采用慢性中等强度不可预知应激刺激模型(chronic unpredictable mild stress, CUMS)和动物行为绝望实验法,研究生、制首乌抗抑郁活性的差异,制首乌(5 g/kg.d)与模型组相比有显著差异(P< 0.01),生首乌制首乌(5g/kg.d)与模型组相比无显著差异,这一结果表明制首乌抗抑郁活性显著优于生首乌。 本文比较了生、制首乌对四氧嘧啶糖尿病模型小鼠血糖的影响的差异,生首乌(5 g/kg.d)与模型组相比有显著差异(P< 0.01),制首乌(5 g/kg.d)与模型组相比无显著差异,这一结果表明生首乌降糖活性优于制首乌。这一结果与历代中医古书中生首乌治疗消渴症(糖尿病)的记载一致。 3生、制首乌化学成分差异的研究结果 本文选用HPLC-DAD指纹图谱技术结合药效成分含量测定来研究生、制首乌化学成分的差异。炮制后,何首乌中的主要化学成分并未消失,只是其含量发生了改变。炮制后二苯乙烯苷含量明显降低:生首乌为5.512 %、清蒸制首乌为3.811 %、豆制首乌为3.538 %,大黄素的含量炮制后显著升高,生首乌为0.094 %、清蒸制首乌为0.119 %、豆制首乌为0.126 %。 综上所述,炮制前后何首乌中二苯乙烯苷和大黄素含量比的变化可能是何首乌炮制减毒、改性的物质基础。 根据上述结果我们建立了生、制首乌的质量控制新模式。 In recent years, some adverse drug reactions (ADR) about some traditional Chinese medicine were reported at times. As a Chinese medicine most in use, the ADRs of Radix Polygoni multiflori (RPM) and the medicines containing the RPM were also mentioned. The resolution of the problems caused by the ADRs is very important for the use of the RPM as a medicine. The process (or preparation) is a significant feature for the clinical use of the Chinese medicine and an important technology for the safe use and good effect of the Chinese medicine. By processing, the toxicity of the Chinese medicine can be reduced, its properties can be changed and curative effect can be enhanced at the same time. The changes of the gene expression profiles for KM mice hepatotoxic effects, and the change of the biological activity and the chemical composition after being processed of the RPm were studied in the present dissertation. The RPm heatotoxicity mechanism and the toxicity target genes were explained on the gene level for the first time. With the antidepressant activity, and the hypoglycemic effect as the target, the differences on the pharmacodynamics between the processed RPm and unprocessed RPm, for the first time, were investigated. The results obtained show that the antidepressant activity of the processed RPM is far higher than the ones of unprocessed RPm. As we know, the results were reported for the first time. The quality control systems (QCS) for the processed and the unprocessed RPm were founded. The HPLC-DAD was used in the systems founded on the basis of the toxicology and the pharmacodynamics experiments. As we know, the OCSs were reported for the first time. The above-mentioned experimental results confirm that the unique process theory of the traditional Chinese medicine (TCM) used for the process of the Radix Polygoni multiflori (RPm) is correct, i.e after being processed the toxicity of the RPm decreases and its Pharmacodynamic effects change. It is known to author that there have been no similar reports in the literatures up to now. The main experimental results are summarized as follows: 1 The results on the mice toxicology gene chip for the unprocessed and processed RPm The KM mice hepatotoxic model caused by the RPm at the different dosages was established in the present study. The results obtained show that the mouse average body weight obviously decreased in the groups at the different dosages of the unprocessed RPm: the 10 g/kg.d .group decreased 20%; 20 g/kg.d. group decreased 42%, and 50% mice died at 20 g/kg.d. group. The main experimental results on the mice toxicology gene chip The RPm is the CYP450 inhibitor. As compared with the processd RPm, the CYP3A4, CYP4A5 of the unprocessed RPm demonstrate the marked downregulation, which leads to the increase of the poison absorbtion into the body with the result that the unprocessed RPm yields the marked hepatotoxication. The hepatotoxication was produced because the following 6 pathways were affected: ①PPAR signaling pathway, the chief toxicity target genes are RXRB, CYP7a1, Acadl, Apoa2, Cyp4a, FABP2 and MAPKKK5 etc. ②Calcium signaling pathway, the chief toxicity target genes are CAMK2B, CACNA1F, S100A1, F2R, Ryr1,Slc8a2 and Camk4 etc. ③Neuroactive ligand-receptor interaction, the chief toxicity target genes are Chrm4, Ntsr2, GABRR1, GRIK3 and F2R etc. ④Wnt signaling pathway, the chief toxicity target genes are Daam2, Rac1 etc. ⑤Complement and coagulation cascades, the chief toxicity target genes are F2R, Serpina1b, Cfi and FGA etc. ⑥Oxidative phosphorylation, the chief toxicity target genes are Atp5e, NDUFA1 etc. The above experimental results, for the first time , demonstrate on the gene level that the unprocessed Rpm toxicity is far stronger than the processed RPm one, and the toxicity of the water decoction of the unprocessed RPm is greater than the one of its acetone extracts, which shows that the chief toxicity components of the RPm are probably not only the anthraquinones, for example, the emodin, but the complex compounds produced by the interaction between the emondin and the stilbene glucoside which is the largest component of the unprocessed RPm. The result is accordance with the above effect of the RPm on the hepatic drugenzyme. Aftter being processed, in fact, the content of the stibene glucoside in the RPm markedly decreases. 2. The results on the pharmacodynamic differences between the unprocessed and processed RPm The results obtained show that the effects of processing on RPm pharmacodynamic behaviour received in the Chinese Material Medica are correct. It is known to author that this is the first experimental result in the research materials now available. The chief results are as follows: For the treatment of the antidepressant, the curative effect of the processed RPm is far better than the one of the unprocessed RPm. By contrast with the above results, the hypoblycemic effect of the unprocessed RPm is better than the one of the processed RPm. 3. The results on the Chemical Composition The results obtained by using HPLC-DAD fingerprint and by the determination of effective component content show that the main chemical components in the RPm after being processed do not disappear, but their contents change. The contents of the stilbene glucoside (SG) and emodin in the different samples were determined as follows: SG contents 5.512 % for the unprocessed RPm 3.811 % for the processed RPm (Steamed) 3.588 % for the processed RPm (black soybean) Emodin contents 0.094 % for the unprocessed RPm 0.119 % for the processed RPm (Steamed) 0.126 % for the processed RPm (black soybean) The combination of above experimental results on the toxicity, the pharmacodynamics and the chemical composition indicates that the changes of the content ratio of SG/emodin may be the substance base of the toxicity decrease and pharmacodynamic changes of the RPM by the processing.
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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。本项目以青杨组杨树为模式植物,从形态和生理方面研究了来自不同UV-B背景下的康定杨与青杨在增强UV-B下的反应及其反应差异,并探讨了干旱、施肥对它们抗UV-B能力的影响。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,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。主要研究结果有以下: 1. 在温室中经过增强UV-B处理,杨树的外部形态及生理活动受到了一定程度的影响。增强UV-B导致康定杨、青杨的生物量、叶面积及节间长度降低,叶片增厚,SOD活性升高,膜伤害增加,而对叶片数目、R/S、叶绿素A、叶绿素B及整个叶绿素含量没有影响。两种杨树对UV-B胁迫的响应存在差异:在增强UV-B条件下,青杨的植株高度、生物量、叶面积、脯氨酸含量、长期用水效率受到的影响大于康定杨,相比而言,康定杨在比叶面积、叶片厚度、可溶性糖含量、UV-B吸收物质的含量及SOD和GPX活性方面增加的程度大于青杨。这些区别说明,来自于高海拔的康定杨比来自于低海拔的青杨对增强UV-B 具有更强的耐性。我们认为二者在叶片厚度、比叶面积、UV-B吸收物质含量及SOD、GPX活性差异是导致对增强UV-B耐性不同的原因。 2. 干旱与增强UV-B对杨树的生长和生理特性均产生了影响,而且两种胁迫共同作用时干旱表现减弱或加剧了UV-B对杨树某些形态和生理特性的影响。 据试验结果,干旱显著地降低了杨树的株高、叶片数目、叶面积,增加了叶片厚度,促进ABA的积累,提高了CAT活性。对于干旱,两种杨树之间也表现出了一定的差异性。可溶性蛋白质和脯氨酸在青杨叶片中得到显著积累,而在康定杨中没有变化。此外,CAT、长期用水效率在康定杨中受到的影响更加明显。长期用水效率的不同变化趋势说明两种杨树对水分胁迫采用了不同的用水策略,康定杨采用的是节水用水策略,提高用水效率,而青杨采用的是耗水的用水策略。根据干旱对叶面积、脯氨酸、ABA含量、CAT活性及长期用水效率等方面的影响,我们认为来自高海拔地区的康定杨比来自低海拔的青杨有更大的耐旱性,这是对生长环境长期适应的结果。在高海拔地区,因霜冻常带来土壤水分不可利用,降低了根系对水分的吸收,树木容易受到的生理性干旱。另外,高海拔的地区低的气温使植株对严寒有较强的耐性,减少了水分的需要。 生长于增强UV-B下的康定杨和青杨植株表现为高度降低,叶面积缩小,比叶面积增加;叶片栅栏组织、海绵组织均受到增强UV-B的影响,其厚度的增加导致整个叶片变厚。增强UV-B还显著提高了杨树的APX活性、UV-B吸收物质含量,而对叶片数目、ABA、可溶性蛋白质含量及CAT活性没有产生影响。试验中也观察到了两种杨树对增强UV-B响应的差异:与康定杨相比,在增强UV-B下青杨株高、叶面积降低的程度更大一些,SOD活性显著提高。另外UV-B吸收物质受到的影响不同。根据这些差别,高海拔的康定杨(3500 m)比来自低海拔的青杨(1500 m)增强UV-B有较强的耐性。 与水分充足情况下UV-B对植株的影响相比,干旱对杨树抗增强UV-B产生了一定的影响,表现为加剧或减弱UV-B对植物的影响,但这种影响与形态、生理指标有关。当干旱与增强UV-B共同作用时,杨树植株的株高、叶面积进一步降低、叶片进一步增厚。就脯氨酸的积累的而言,在没有水分胁迫时,增强UV-B促使它显著增加,而在干旱处理下这种效果变得不明显。干旱对增强UV-B的影响还与杨树的种类有一定的关系。在康定杨中,干旱减弱了增强UV-B对栅栏组织与海绵组织的影响,且在植株高度、叶面积上表现出累加效应,而在CAT上交互作用显著;但在青杨中干旱则加剧增强UV-B对栅栏组织与海绵组织的影响,在植株高度、叶面积及比叶面积上表现出显著的交互作用。据碳同素分析,在水分充足的条件下,无论是康定杨,还是青杨,增强UV-B均导致其长期用水效率的提高,然而当两种胁迫共同作用时,长期用水效率则表现出差异,在青杨中,长期用水效率得到进一步增高,而康定杨中干旱的效应被增强UV-B所减轻。 3. 田间试验表明,杨树的生长、生理特征都受到养分和增强UV-B的影响。施肥对杨树的影响表现为:提高了叶面积、生物量及SOD的活性,降低了抗坏血酸含量。对于施肥作用,两种杨树的反应也有区别:在康定杨中施肥显著增加了的叶片长度、宽度及光合色素的含量,降低了净光合速率、气孔导度及胞间CO2浓度;在青杨中,则SOD、GPX、APX活性表现增加。从试验看出,施肥对来自于高海拔地区的康定杨(3500 m)的影响较大,对来自低海拔的青杨(1500 m)影响较小,这与它们对原产地的生境适应有一定关系。在康定杨生长的高海拔地区,低温度和湿度不能为地上凋落物或土壤中的根分解提供理想的条件,造成当地土壤的低养分状况,所以当肥料施用以后,效果显著。 经过增强UV-B处理,杨树叶片中UV-B吸收物质含量、GPX的活性得到提高,而脯氨酸、丙二醛、可溶性蛋白质、叶绿素及类胡萝卜素含量没有受到影响。对于增强UV-B两种杨树受到的影响也有所不同:在青杨中增强UV-B导致叶面积缩小,生物量、净光合速率降低,APX的活性及长期用水效率的提高,而对康定杨的这些指标没有产生显著影响,相反抗氧化酶的活性明显高于青杨。这些差异性是由于两种杨树对原产地不同UV-B背景的长期适应结果。康定杨长期生长在较高UV-B环境中,对UV-B有较强的耐性。而青杨适应于较低的UV-B环境,对增强UV-B较为敏感。 试验中施肥也影响了植株对增强UV-B的反应,不过这种影响与杨树的种类及测定指标有一定的相关性。例如,在缺肥的情况下,青杨的长期用水效率和康定杨的叶绿素含量都受到增强UV-B的显著影响,而施肥以后这种影响变得不显著。在缺肥的条件下,GPX、APX在青杨中的活性、GPX在康定杨中的活性对增加UV-B反应不敏感;而施肥以后则变化显著,同样胞间CO2浓度在康定杨也有类似的变化。 For past decades, 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. In this experiment, different species of Populus section Tacamahaca Spach from different UV-B background were selected as a model plant to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B were observed and the different responses between P. kangdingensis and P. cathayana were discussed, furthermore the influences of drought and fertilizer on responses induced by enhanced UV-B were studied. Since poplars play an important role in lumber supply, and are important component of ecosystems due to their fast growth and wide adaptation, the study could provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem. The results are as follows: 1. The experiment conducted in a greenhouse indicated that morphological and physiological traits of two poplars were affected by enhanced UV-B radiation. Enhanced UV-B radiation not only reduced biomass, leave area and internode length, but also increased leaf thickness and SOD activity as well as MDA concentration and electrolyte rate. However, no significant changes in leaf numbers, root shoot ratio, and total chlorophyll and chlorophyll component were observed. There were different responses to enhanced UV-B radiation between two species. Compared with P. kangdingensis, cuttings of P. cathayana, exhibited lower height increment and smaller leaf area. In addition, there were significant differences in free proline, soluble protein, and UV-B absorbing compounds, and the activity of SOD and GPX, long-term WUE between them. Differences in plant height, biomass, leaf area, free proline concentration, and long-termed WUE showed that P. cathayana were more affected by enhanced UV-B radiation than P. kangdingensis. In contrast, more increase of specific leaf mass, leaf thickness, and soluble sugar, and UV-B absorbing compounds, and activity of SOD and GPX were observed in P. kangdingensis. According to these results, we suggested that P. kangdingensis from high elevation, which adapted to higher UV-B environments, had more tolerance to enhanced UV-B than P. cathayana from low elevation, which adapted to lower UV-B environment. We believe it was the difference of leaf thickness, specific leaf mass, and UV-B absorbing compounds as well as the activity of SOD and GPX resulted in lower adaptation of P. cathayana to enhanced UV-B radiation. 2. Growth and physiological traits of two poplars were affected by both drought and enhanced UV-B radiation. Moreover, it was observed that when two stresses applied together drought could exacerbate UV-B effects or decrease sensitivity to UV-B. In the experiment, drought significantly decreased plant height, leaf numbers, leaf area, and increased leaf thickness, and ABA, and CAT activity of two poplars. There were significant interspecific differences to drought stress. Exposed to drought, soluble protein and proline concentration were increased in P. cathayana but not in P. kangdingensis. However, more changes in CAT and long-term WUE were observed in kangdingensis. Different change in long-term WUE suggests that two poplars adapted different water-use strategies. P. kangdingensis employ a conservative water-use strategy, whereas P. cathayana employ a prodigal water-use strategy. Based on the differences in leaf area, accumulation of free proline and ABA, CAT activity as well as long-term WUE, we believed that P. kangdingensis from high elevation had a greater tolerance to drought than P. cathayana from low elevation,which is the result of adaptation to local environment. In high elevation area, trees are prone to suffer from physiological drought because of un-movable water caused by frost. Besides lower temperature enable the plants had greater adaptability to frost as a results the requirement of water is reduced Enhanced UV-B radiation decreased shoots height, leaf area, and increased specific leaf mass and thickness of palisade and sponge layer as well as APX activity and UV-B absorbing compounds in both species. Whereas, leaf numbers, ABA content, soluble protein and CAT activity showed no differences to enhanced UV-B radiation. Interspecific differences were also observed. Compared with P. kangdingensis, P. cathayana showed lower shoot height and smaller leaf area, higher SOD activity. Besides, variation in UV-B absorbing compounds was found. These differences suggested that P. kangdingensis from high elevation (3500 m) was more tolerant to enhanced UV-B radiation than P. cathayana from low elevation (1500 m). Compared with morphological and physiological changes induced by enhanced UV-B radiation under well-watered conditions, drought exacerbated or decreased these changes. However, these effects vary with parameters measured. When two stresses applied together, shoot height and leaf area further decreased while leaf thickness further increased. Under well-watered conditions, enhanced UV-B radiation significantly increased proline content, but such effect was not observed under drought conditions. The effect of drought on enhanced UV-B radiation was related to species. For example, drought reduced the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll in P. kangdingensis, and additive effects in shoot height and leaf area and interactive effect CAT activity were observed. In contrast, for P. cathayana drought significantly exacerbated the effects of enhanced UV-B radiation on palisade parenchyma and sponge mesophyll; there were noticeable interaction in shoot height, leaf area and specific leaf mass. As far as long-term WUE is concerned, it was increased by enhanced UV-B radiation under well-watered conditions in both species. While different effect was observed between two species in combination of two stresses. Long-term water use efficiency was further increased in P. cathayana whereas the effect was less significant in P. kangdingensis. 3. The field experiment showed that growth and physiological traits of poplars were affected by nutrition and enhanced UV-B radiation. Fertilization significantly increased leaf area, biomass and SOD activity, reduced Ascorbic acid concentration. There was interspecific difference in response to fertilization. For P. kangdingensis, fertilization significantly increased leaf width, leaf length and photosynthetic pigments content while net photosynthetic rate and stomatal conductance, intercellular CO2 concentration were significantly decreased. However, for P. cathayana, these parameters were unaffected except the increase of SOD, GPX and APX activity. From above, it could concluded that P. kangdingensis from high elevation was more affected by fertilization than P. cathayana, This difference was due to adaptation to local environment., The low temperature and moisture where P. kangdingensis was collected can not provided optimum to decompose roots and litter fall as a result the nutrition in soil was poor. Exposed to enhanced UV-B radiation, for both species UV-B absorbing compounds and GPX activity were significantly increased while proline, MDA, soluble protein, chlorophyll, carotenoids were not affected. Different responses were also observed between the two species. Enhanced UV-B radiation caused significant decreases in leaf area, biomass, net photosynthetic rate and increase in APX activity and long-term WUE in P. cathayana but not in P. kangdingensis. In addition, activity in antioxidant enzymes was much higher in P. kangdingensis than in P. cathayana. In the experiment fertilization affected responses of cuttings to enhanced UV-B radiation, but it concern species and parameters measured. Long-term WUE in P. cathayana and chlorophyll in P. kangdingensis were significantly increased by enhanced UV-B radiation under non-fertilization treatments while the increase was not found under fertilization treatment. In contrast, under no fertilization treatment enhanced UV-B radiation did not affected GPX and APX activity in P. cathayana and GPX in P. kangdingensis while significant increase appeared after application of fertilization. Similar effect of enhanced UV-B radiation on intercellular CO2 concentration in P. kangdingensis was observed.
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由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20 世纪升高了0.6 ¡æ,并预测在本世纪将上升1.4-5.8 ¡æ。气候变暖对陆地植物和生态系统产生深远影响,并已成为全球变化研究的重要议题。位于青藏高原东部的川西亚高山针叶林是研究气候变暖对陆地生态系统影响的重要森林类型。森林采伐迹地和人工云杉林下作为目前该区人工造林和森林更新的两种重要生境,二者截然不同的光环境对亚高山针叶林不同物种更新及森林动态有非常重要的影响。 本文以青藏高原东部亚高山针叶林几种主要森林树种为研究对象,采用开顶式增温法(OTCs)模拟气候变暖来研究增温对生长在两种不同光环境下(全光条件和林下低光环境)的几种幼苗早期生长和生理的影响,旨在从更新角度探讨亚高山针叶林生态系统不同树种对气候变暖在形态或生理上的响应差异,其研究结果可在一定程度上为预测气候变暖对亚高山针叶林物种组成和演替动态提供科学依据,同时也可为未来林业生产管理者提供科学指导。 1、与框外对照相比,OTCs 框内微环境发生了一些变化。OTCs 框内与框外对照气温年平均值分别为5.72 ¡æ和5.21 ¡æ,而地表温度年平均值分别为5.34 ¡æ和5.04 ¡æ,OTCs 使气温和地表年平均温度分别提高了0.51 ¡æ和0.34 ¡æ;OTCs框内空气湿度年平均值约高于框外对照,二者分别为90.4 %和85.3 %。 2、增温促进了三种幼苗生长和生物量的积累,但增温效果与幼苗种类及所处的光环境有关。无论在全光或林下低光条件下,增温条件下云杉幼苗株高、地径、分支数、总生物量及组分生物量(根、茎、叶重)都显著地增加;增温仅在全光条件下使红桦幼苗株高、地径、总生物量及组分生物量(根、茎、叶重)等参数显著地增加,而在林下低光条件下增温对幼苗生长和生物量积累的影响效果不明显;冷杉幼苗生长对增温的响应则与红桦幼苗相反,增温仅在林下低光条件下对冷杉幼苗生长和形态的影响才有明显的促进作用。 增温对三种幼苗的生物量分配模式产生了不同的影响,并且这种影响也与幼苗所处的光环境有关。无论在全光或林下低光环境下,增温都促使云杉幼苗将更多的生物量分配到植物地下部分,从而导致幼苗在增温条件下有更高的R/S 比;增温仅在林下低光条件下促使冷杉幼苗将更多的生物量投入到植物叶部,从而使幼苗R/S 比显著地降低;增温在全光条件下对红桦幼苗生物量分配的影响趋势与冷杉幼苗在低光条件下相似,即增温在全光条件下促使红桦幼苗分配更多的生物量到植物同化部分—叶部。 3、增温对亚高山针叶林生态系统中三种幼苗气体交换和生理表现的影响总体表现为正效应(Positive),即增温促进了几种幼苗的生理活动及其表现:(i)无论在全光或林下低光环境下,增温使三种幼苗的光合色素含量都有所增加;(ii)增温在一定程度上提高了三种使幼苗的PSII 光系统效率(Fv/Fm),从而使幼苗具有更强的光合电子传递活性;增温在一定程度使三种幼苗潜在的热耗散能力(NPQ)都有所增强,从而提高幼苗防御光氧化的能力;(iii)从研究结果来看,增温通过增加光合色素含量和表观量子效率等参数而促进幼苗的光合作用过程。总体来说增温对幼苗生理过程的影响效果与幼苗种类及所处的光环境有关,增温仅在全光条件下对红桦幼苗光合过程的影响才有明显的效果,而冷杉幼苗则相反,增温仅在低光条件下才对幼苗的生理过程有显著的影响。 4、增温对三种幼苗的抗氧化酶系统产生了一定的影响。从总体来说,增温使几种幼苗活性氧含量及膜脂过氧化作用降低,从而在一定程度上减轻了该区低温对植物生长的消极影响;增温倾向表明使三种幼苗体内抗氧化酶活性和非酶促作用有所提高,从而有利于维持活性氧代谢平衡。但增温影响效果与幼苗种类所处的光环境及抗氧化酶种类有关,增温对冷杉幼苗抗氧化酶活性的影响仅在林下低光环境下效果明显,而对红桦幼苗抗氧化酶活性的影响仅在全光条件下才有明显的效果。 总之,增温促进了亚高山针叶林生态系统中三种幼苗的生长和生理表现,但幼苗生长和生理对增温的响应随植物种类及所处的光环境不同而变化,这种响应差可能异赋予了不同植物种类在未来气候变暖背景下面对不同环境条件时具有不同的适应力和竞争优势,从而对亚高山针叶林生态系统物种组成和森林动态产生潜在的影响。 Enrichment of atmospheric greenhouse gases resulted from human activities suchas fossil fuel burning and deforestation has increased global mean temperature by 0.6¡æ in the 20th century and is predicted to increase it by 1.4-5.8 ¡æ. The globalwarming will have profound, long-term impacts on terrestrial plants and ecosystems.The ecoologcial consequences arising from global warming have also become thevery important issuses of global change research. The subalpine coniferous forests inthe eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying theeffects of climate warming on terrestrial ecosystems. The light environment differssignificantly between clear-outs and spruce plantations, which is particularlyimportant for plant regeneration and forest dynamics in the subalpine coniferous forests. In this paper, the short-term effects of two levels of air temperature (ambient andwarmed) and light (full light and ca. 10% of full light regimes) on the early growthand physiology of Picea asperata, Abies faxoniana and Betula albo-sinensis seedlingswas determined using open-top chambers (OTCs). The aim of the present study wasto understand the differences between tree species in their responses to experimentalwarming from the perspective of regeneration. Our results could provide insights intothe effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientificdirection for the production and management under future climate change. 1. The OTCs manipulation slightly altered thermal conditions during the growingseason compared with the outside chambers. The annual mean air temperature andsoil surface temperature was 5.72 and 5.34 ¡æ (within the chambers), and 5.21 and5.04 ¡æ (outside the chambers), respectively. The OTCs manipulation increased airtemperature and soil surface temperature by 0.51 and 0.34 ¡æ on average, respectively.Air relative humidity was slightly higher inside the OTCs compared with the controlplots, with 90.4 and 85.3 %, respectively. 2. Warming generally stimulated the growth and biomass accumulation of thethree tree species, but the effects of warming on growth and development variedbetween light conditions and species. Irrespective of light regimes, warmingsignificantly increased plant height, root collar diameter, total biomass, componentbiomass (stem, foliar and root biomass) and the number of branches in P. asperataseedlings; For A. faxoniana seedlings, significant effects of warming on all the tested parameters (plant height, root collar diameter, total biomass, and component biomass) were found only under low light conditions; In contrast, the growth responses of B.albo-sinensis seedlings to warming were found only under full light conditions. Warming had pronounced effects on the pattern of carbon allocation. Irrespectiveof light regimes, the P. asperata seedlings allocated relatively more biomass to rootsin responses to warming, which led to a higher R/S. Significant effects of warming onbiomass allocation were only found for the A. faxoniana seedlings grown under lowlight conditions, with significantly increased in leaf mass ratio (LMR) and decreasedin R/S in responses to warming manipulation. The carbon allocation responses of B.albo-sinensis seedling to warming under full light conditions were similar with theresponse of A. faxoniana seedlings grown under low light conditions. Warmingsignificantly decreased root mass ratio (RMR), and increased leaf mass ratio (LMR)and shoot/root biomass ratio (S/R) for the B. albo-sinensis seedlings grown under full light conditions. 3. Warming generally had a beneficial effect on physiological processes of dominant tree species in subalpine coniferous forest ecosystems: (i) Warming markedincreased the concentrations of photosynthetic pigments in both tree species, but theeffects of warming on photosynthetic pigments were greater under low lightconditions than under full light conditions for the two conifers; (ii) Warming tended toenhance the efficiency of PSII in terms of increase in Fv/Fm, which was related tohigher chloroplast electron transport activity; and enhance non-radiative energydissipation in terms of in increase in NPQ, which may reflect an increased capacity inpreventing photooxidation; (iii) Warming may enhance photosynthesis and advancephysiological activity in plants by increasing photosynthetic pigment concentration,the efficiency of PSII and apparent quantum yield (Φ) etc. From the results, theeffects of warming on seedlings’ physiological performance varied between lightenvironment and species. The effects of warming on photosynthesis performance of B.albo-sinesis seedlings were pronounced only under full light conditions, while thephysiological responses of A. faxoniana seedlings to warming were found only underthe 60-year plantation. These results provided further support for the observationsabove on growth responses of seedlings to warming. 4. Warming had marked effects on antioxidative systems of the three seedlings.Warming generally decreased H2O2 accumulation and the rate of O2- production, andalleviated degree of lipid peroxidation in terms of decreased MDA content, whichalleviated to some extent the negative effects of low temperature on the plant growthand development in this region; Warming tended to increase the activities ofantioxidative enzymes and stimulate the role of non-enzymatic AOS scavenging,which helped to create an balance in maintaining AOS metabolites for the threeseedlings. Nevertheless, the effects of warming on antioxidative defense systems werepronounced only under the 60-year plantation for the A. faxoniana seedlings. Incontrast, the marked effects of warming on antioxidative defense systems for the B.albo-sinesis seedlings were found only under the full light conditions. In sum, warming is considered to be generally positive in terms of growth andphysiological process. However, the responses of growth and physiology performanceto warming manipulation varied between species and light regimes. Competitive and adaptive relationships between tree species may be altered as a result of responsedifferences to warming manipulation, which is one mechanism by which globalwarming will alter species composition and forest dynamics of subalpine coniferousforest ecosystems under future climate change.
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杨树具有分布广、适应性强的特征,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,也是生长较迅速、易繁殖的重要杨树资源。本研究选取了来自不同气候地区的青杨两种群为材料,采用植物生态学、生理学和生物化学的研究方法,系统地研究了青杨对干旱与遮荫、干旱与外源脱落酸(ABA)喷施的生长、形态、生理和生化响应及种群间差异,研究成果可为我国干旱半干旱地区的造林以及生态恢复提供理论依据和科学指导。主要研究结论如下:1.青杨在干旱胁迫下的适应机制为:生长性状及生物量的分配变化:干旱胁迫下虽然植株生长受抑,株高、基茎及各部分生物量都显著减小,但有相对较多的生物量向根部分配,根/冠比以及细/粗根比增加。青杨对干旱胁迫的光合作用表现为:干旱胁迫降低了青杨的净光合速率、蒸腾速率、气孔导度以及光合氮利用效率,提高了瞬时用水效率。干旱还引起了活性氧的产生,使得膜脂过氧化产物丙二醛(MDA)增加,同时也增强了植物抗氧化酶系统(如超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性的增加)及非酶系统的能力(如抗坏血酸(AsA)含量的增加)。干旱降低了植物叶片的相对含水量,而促进了渗透调节物质(游离脯氨酸及可溶性糖)的积累,增加了植物的渗调能力。干旱下青杨两种群的内源ABA含量显著增加,碳同位素组分(δ13C)也显著提高。这些结果证明植物遭受干旱胁迫后发生一系列的形态、生理和生化响应,这些变化能提高植物在干旱下的存活和生长能力。2.青杨两种群对干旱胁迫反应的种群差异:与来自湿润地区的汉源种群相比,来自干旱地区的乐都种群在干旱条件下生物量向根系分配的可塑性更强,同时具有更强的抗氧化系统能力,所受到活性氧的伤害也更少,并且累积更多的脯胺酸和ABA,具有更高的δ13C。这些都说明了乐都种群对干旱的适应性比汉源种群更强。两种群对干旱的响应差异应归于它们的用水策略的不同:汉源种群来自湿润地区,采用了耗水型的用水策略,抗旱能力较弱;而乐都种群,来自干旱地区,通常采用节水型的用水策略,有更强的抗旱能力。3.遮荫对青杨两种群抗旱性的影响:遮荫对青杨抗旱性的影响决定于遮荫程度的不同,我们的结果表明中度的遮荫可以有效的提高干旱下植物的生长,对干旱胁迫有明显的缓解作用,具体体现在中度遮荫下受旱植物的叶片相对含水量得到提高,使得植物体内水分状况得到了改善;光合速率并未降低,植物光合氮利用效率增加,说明中度的遮荫并未明显限制植物的碳获得;抗氧化酶活性与膜脂过氧化产物MDA含量的同时降低,说明中度遮荫下所受到的活性氧伤害减少;中度遮荫下的ABA及δ13C的变化也不如在全光下变化明显,这也说明中度遮荫缓解了干旱胁迫。但是重度的遮荫却对干旱胁迫有明显的加剧作用,主要表现在重度遮荫降低了植物的光合速率,严重抑制了植物的生长;同时重度遮荫下脯胺酸含量和抗氧化酶活性的急剧下降,导致了植物渗调能力的下降及膜脂过氧化产物MDA的显著升高;重度遮荫还显著降低了内源ABA的累积和δ13C,降低了植物的抗旱能力。此外,青杨两种群在对干旱和遮荫的响应中,也表现出种群差异。汉源种群,来自湿润且年日照辐射较少的地区,表现出相对更强的耐荫性和需水性。而乐都种群,来自干旱且年日照辐射丰富的地区,表现出相对更强的耐旱性和需光性。这说明了植物对环境胁迫的耐受性是其长期适应原生境的结果,并且来自不同气候地区的两种群在面临环境胁迫时会采取不同的生存策略。4. 外源ABA喷施对青杨两种群抗旱性的影响:外源ABA的喷施可以提高两种群的抗旱性,具体表现为外源ABA喷施促进了青杨根系的生长,显著提高了干旱下植物的根/冠比和细/粗根比,减少了比叶面积;在生理生化方面,外源ABA降低了干旱下植物叶片的气孔导度,降低了蒸腾速率和净光合速率,但提高了瞬时用水效率,提高了叶片的相对含水量,增加了干旱下植物的保水能力。外源ABA进一步增加了干旱下植物内源ABA的积累,促进了植物渗调物质如脯胺酸和可溶性糖的积累,增加了抗氧化酶系统(如SOD、APX、CAT)的活性和非酶系统AsA的含量,降低了活性氧(如超氧阴离子(O2和过氧化氢(H2O2))对植株的伤害。此外,外源ABA还进一步提高了干旱下植物的δ13C,提高了植物的长期用水效率,由此提高了植物的抗旱能力。另一方面,两种群对外源ABA和干旱的响应也有所差别。来自湿润地区的汉源种群,对干旱较为敏感,所受干旱的影响也较大,而外源ABA的喷施对汉源种群抗旱性的提高作用也更为突出。乐都种群,由于其长期适应干旱地区的生长,本身已具有较强的抗旱能力,因此外源ABA喷施对其抗旱性的提高不如对汉源种群的效果明显。由此我们可以得出对于一些抗性弱或干旱敏感的物种或者种群,可以采用外施ABA的方法来提高其抗性。Poplars play an important role in lumber supply, and are important component ofecosystems 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 ofpoplars for its fast growth and reproductive. In this study, different populations of P.cathayana were used as experiment material to investigate the adaptability to drought stressand population differences in adaptability, and the effects of shade and exogenous abscisicacid (ABA) application on the drought tolerance. Our results could provide a strongtheoretical evidence and scientific direction for the afforestation, and rehabilitation ofecosystem in the arid and semi-arid area, and provide a strong evidence for adaptivedifferentiation of different populations, and so may be used as criteria for species selectionand tree improvement. The results are as follows:1. A large set of parallel response to drought stress: Drought stress caused pronouncedinhibition of the growth and increased relatively dry matter allocation into the root. For thetwo populations, the shoot height, basal diameter and total biomass were decreased but theroot/shoot ratio and fine root/coarse root ratio were increased under drought conditions;Drought stress caused pronounced inhibition of photosynthesis, decreased the stomatalconductance, transpiration rate, and photosynthetic nitrogen-use efficiency (PNUE) butincreased the instantaneous water use efficiency. Drought significantly improved the levels ofreactive oxygen species and malondialdehyde (MDA) and to induce the entire set ofantioxidative systems including the increase of activities of superoxide dismutase (SOD),ascorbate peroxidase (APX), catalase (CAT) and ascorbate (AsA) content. Drought decreased the leaf relative water content (RWC) but improved the capability of osmotic adjustmentindicated by the higher proline accumulation. Drought also increased the ABA content andcarbon isotope composition (δ13C), which indicating the long period water use efficiency wasimproved under drought. These results demonstrate that there are a large set of parallelchanges in the morphological, physiological and biochemical responses when plants areexposed to drought stress; these changes may enhance the capability of plants to survive andgrow during drought periods.2. Difference in adaptation to drought stress between contrasting populations of P.cathayana: Compared with the Hanyuan population (wet climate), the Ledu population (dryclimate) showed higher root/shoot ratio and water use efficiency, exhibited higherantioxidative systems capability thus resulting in less oxidative damage, accumulated moreABA and free proline content under drought conditions. The results suggested that there weredifferent water-use strategies between the two populations. The Ledu population, whichcomes from dry climate region, with higher drought tolerance, may employ a conservativewater-use strategy, whereas the Hanyuan population, which comes from wet climate, withlower drought tolerance, may employ a prodigal water-use strategy. These variations indrought responses may be used as criteria for species selection and tree improvement.3. The effects of shade on the drought tolerance: The reduction in the availability of lightand water affected the morphological and physiological responses of the two P. cathayanapopulations. In addition, the light environment modified the growth responses of P.cathayana seedlings to varying water environments in different ways depending upon theintensity of the light levels considered. There is an apparent alleviation to drought effects bymoderate shade in P. cathayana seedlings, as indicated by the higher leaf RWC, and unchanged net photosynthesis and PNUE, as well as by the lower antioxditative enzymeactivity, MDA, ABA and δ13C levels, which implied moderate shade did not significantlylimited the carbon acquisition or inhibited the plant growth, but ameliorated the detrimentaleffects of drought. On the other hand, an apparent aggravation to drought effects by severeshade was also observed, as indicated by the pronounced decrease of plant growth and net photosynthesis, the lower total biomass, ABA level, δ13C, free proline content andantioxditative enzyme activity and higher MDA accumulation. By contrast, the twopopulations showed different responses to shade and drought. The Hanyuan population,which comes from a riparian basin having a relatively wet climate and less annual solarradiation, is more sensitive to drought but more tolerant to shade. The Ledu population, whichcomes from a mountainous plateau with less rainfall and with more annual solar radiation, ismore tolerant to drought but more sensitive to shade. The results demonstrated that theendurance of plants to stress is a result of long-term evolution and adaptation to theenvironment, as suggested by the different strategies employed by the P. cathayanapopulations originating from contrasting habitats when they were exposed to drought andshade.4. The effects of exogenous ABA application on the drought tolerance: For bothpopulations under drought conditions tested, exogenous ABA application significantlyimproved the root/shoot ratio, fine root/coarse root ratio, and decreased the specifical leaf area.On the physiological and biochemical traits, exogenous ABA application significantlydecreased stomatal conductance, transpiration rate and net photosythesis but increased theinstance water use efficiency and leaf RWC. On the other hand, exogenous ABA applicationsignificantly increased endogenous ABA, proline, solube sugar and AsA content, as well asSOD, APX and CAT activities, thus reduced the damage of reactive oxygen species. Moreover,the long period water use efficiency as indicated by δ13C was also improved by exogenousABA application. In additionally, there was different responsive between the two populationsto drought and exogenous ABA application. The Hanyuan population, which comes from wetclimate region, is more sensitive to drought, and the effect of exogenous ABA is moreobviously than that in the Ledu population, which comes from dry climate region and is moredrought-responsive. Therefore, we can use exogenous ABA application to improve theresistance of plants, especially for the drought- sensitive species or populations.
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克隆植物被认为比非克隆植物更宜于利用异质性环境。在复杂的空间异质环境中,克隆植物可能形成了各种有效利用环境异质性的适应对策。对于克隆植物适应机制的研究,前人已做了大量的工作,特别是从形态和生物量分配等方面对简单异质生境下克隆植物的克隆整合和克隆分工进行了详细的研究。本研究以分布广泛的克隆植物东方草莓(Fragaria orientalis)作为研究对象,应用野外调查和实验生态学方法,采用多对比度单资源模型和不同向双资源模型,从形态和生理生态的角度,研究复杂异质生境下克隆植物的整合和分工及其耗益问题,分析不同类型的生境对克隆植物整合和分工的修饰作用,进而探讨克隆植物对异质生境的适应策略。克隆构型和分株种群特征是植物克隆生长及其生态适应对策研究的基本内容。本文通过野外调查,研究在不同光照条件下东方草莓克隆构型、分株种群特征以及点分布格局。结果表明:东方草莓的克隆构型随光照发生相应的变化,低光照下其匍匐茎节间长和分枝角度均增大而分枝强度减小;随光照减弱,东方草莓分株种群的生物量、根冠比和分株种群密度显著降低;不同光照下东方草莓分株均以随机分布为主但不同尺度下有所差异,其分布格局强度依次为旷地<林缘<林下。结合克隆植物对资源的利用对策,探讨了克隆构型和分株种群特征以及分布格局随环境条件变化的生态适应意义。不同生境斑块条件下克隆植物可能采取不同的适应对策。采用盆栽实验,研究不同水分对比度下克隆整合及其生理生态特征,并对单向和交互资源中东方草莓的克隆整合做了对比研究。结果显示:高的水分对比度能够促进东方草莓的克隆整合,并能刺激相连分株增加光合作用,东方草莓体内的氧化—抗氧化系统也II随对比度做出相应的反应。耗-益分析表明胁迫分株的受益是以供给分株的损耗为代价的,但从克隆片段总体来说是受益的。单向资源中东方草莓生长的绝对值高于交互资源,但耗-益分析表明生长于交互资源下东方草莓的克隆整合获益大于生长于单向资源下东方草莓的克隆整合获益。长期生长于特定生境的克隆植物,在进化过程中其克隆整合和克隆分工在对资源异质性的适应策略方面可能有所侧重。采用盆栽实验对来自不同海拔梯度的东方草莓的克隆整合和克隆分工对异质资源的适应对策进行了研究。实验结果表明,来自高海拔的东方草莓可塑性较差。来自两个海拔的东方草莓对切断匍匐茎的表现有所差异,总体上切断匍匐茎对来自高海拔的东方草莓影响更大些。另外,来自高海拔的东方草莓表现出更高的克隆分工。IIIClonal plants are known to be more suitable for the habitats of heterogeneousresources than nonclonal plants, perhaps due to their well developed adaptivestrategies to environmental heterogeneity. Many studies have been done on theadaptive mechanisms of clonal plants, especially on the clonal integration anddivision of labor with morphology and biomass allocation under simpleheterogeneous habitats. Based on field surveys, laboratory experiments, multi-contrastunidirectional resource model and reciprocal resource model, Fragaria orientalis, aRosaceae stoloniferous herb that widely distributes in China, was used to study thisplant’s morphological and physiological responses to complicated heterogeneoushabitats in terms of its clonal integration, division of labor and cost-benefit, as well astheir modifications by different habitats, so as to better understand the adaptivestrategies of clonal plants under heterogeneous environments.Clonal architecture and ramet population characteristics are of the major concernin the studies on growth and adaptive strategies of clonal plants. Clonal architecture,ramet population characteristics and spatial point pattern of F. orientalis underdifferent light intensity were studied with field observations. The results showed that,clonal architecture changed with light availability: Internode-lengths and branchangels of stolons were larger while branch intensities were smaller under lower lightintensity than those under higher light intensity; Biomass of ramet population,root-shoot ratio and density of ramet population decreased significantly with reduce oflight intensity; Under all light intensities, spatial pattern of ramets was mainlyrandomly distributed but it changed with different scales, with pattern intensity as:open space < forest edge < understory. Adaptation significance of the clonal architecture, the ramet population characteristics and the spatial pattern changing withdifferent environments was discussed according to these results.Clonal plants may take different adaptive strategies under different patches. Withpot culture, clonal integration and physiological parameters of F. orientalis underdifferent water contrasts were studied, and clonal integration under unilateralresources and reciprocal resources were also compared. The results suggested that,high water contrast improve the clonal integration of F. orientalis and increase thephotosynthesis of connected ramets. Oxidative and antioxidative system of F.orientalis also responded with changing water contrasts. According to cost-benefitanalysis, the drought-stressed ramets obtained benefits from the connectedwell-watered ramets, and as a whole, the clonal fragment could also get benefits.Growth of F. orientalis in homogeneous resources was better than that inheterogeneous resources, but the whole plant got more benefit through clonalintegration in heterogeneous resources than in homogeneous resources.Pot culture experiments were also used to study the adaptive strategies inutilizing heterogeneous resources by the plant populations from different altitudes.The results showed that, F. orientalis from alpine zones were shorter and lessexpanded with poorer clonal plasticity than those from middle mountains. F.orientalis from two different altitudes showed different responses to stolon severing,and as a whole, stolon severing had more influence on F. orientalis from alpine zones.In addition, F. orientalis from alpine zones exhibited higher division of labor, whichsuggested that clonal plants from different habitats develop their own adaptivemechanisms in their clonal integration and division of labor in response toenvironmental heterogeneity.
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干旱环境常常由于多变的降水事件和贫瘠土壤的综合作用,表现出较低的生产力和较低的植被覆盖度。全球性的气候变暖和人类干扰必将使得干旱地区缺水现状越来越严竣。贫瘠土壤环境中已经很低的有效养分含量也将会随着干旱的扩大而越来越低。干旱与半干旱系统中不断加剧的水分与养分的缺失将严重限制植物的生长和植被的更新,必然会使得已经恶化的环境恶化速率的加快、恶化范围的加大。如何抑制这种趋势,逐步改善已经恶化的环境是现在和将来干旱系统管理者面临的主要关键问题。了解干旱系统本土植物对未来气候变化的适应机制,不仅是植物生态学研究的重要内容,也对人为调节干旱环境,改善干旱系统植被条件,提高植被覆盖度具有重要的实践意义。 本研究以干旱河谷优势灌木白刺花(Sophora davidii)为研究对象,通过两年大棚水分和施N控制实验和一个生长季野外施N半控制实验,从植物生长-生理-资源利用以及植物生长土壤环境特征入手,系统的研究了白刺花幼苗生长特性对干旱胁迫和施N的响应与适应机制,并试图探讨施N是否可调节干旱系统土壤环境,人工促进干旱条件下幼苗定居,最终贡献于促进植被更新实践。初步研究结论如下: 1)白刺花幼苗生长、生物量积累与分配以及水分利用效率对干旱胁迫和施N处理的适应白刺花幼苗株高、基径、叶片数目、叶面积、根长、生物量生产、相对含水量和水分利用效率随着干旱胁迫程度的增加而明显降低,但地下部分生物量比例和R/S随着干旱胁迫程度的增加而增加。轻度施N处理下幼苗株高、基径、叶片数目、叶片面积和生物量生产有所增加。但重度施N处理下这些生长指标表现出微弱甚至降低的趋势。严重干旱胁迫条件下,幼苗叶面积率、R/S、相对含水量和水分利用效率也以轻度施N处理为最高。 2)白刺花幼苗叶片光合生理特征对干旱胁迫和施N处理的适应叶片光合色素含量和叶片光合效率随着干旱胁迫程度的增加而显著降低,并且PS2系统在干旱胁迫条件下表现出一定程度的光损害。但是比叶面积随着干旱胁迫程度的增加而增加。在相对较好水分条件下幼苗净光合速率的降低可能是因为气孔限制作用,而严重干旱胁迫条件下非气孔限制可能是导致幼苗叶片光合速率下降的主要原因。叶片叶绿素含量、潜在光合能力、羧化效率、光合效率以及RUBP再生能力等在施N处理下得到提高,并因而改善干旱胁迫条件下光合能力和效率。虽然各荧光参数对施N处理并无显著的反应,但是干旱胁迫条件下qN和Fv/Fm在轻度施N处理下维持相对较高的水平,而两年连续处理后在严重干旱胁迫条件下幼苗叶片光合效率受到重度施N处理的抑制,并且Fv/Fm和qN也在重度施N处理下降低。 3)白刺花幼苗C、N和P积累以及N、P利用效率对干旱胁迫和施N处理的适应白刺花幼苗C、N和P的积累,P利用效率以及N和P吸收效率随干旱胁迫程度的增加而显著降低,C、N和P的分配格局也随之改变。在相同水分处理下,C、N和P的积累量、P利用效率以及N和P吸收效率在轻度施N处理下表现为较高的水平。然而,C、N和P的积累量和P利用效率在重度施N处理下不仅没有表现出显著的正效应,而且有降低的趋势。另外,在相同水分条件下白刺花幼苗N利用效率随着施N强度的增加而降低。 4)白刺花幼苗生长土壤化学与微生物特性对干旱胁迫和施N的适应白刺花幼苗生长土壤有机C、有效N和P含量也随干旱胁迫程度的增加而明显降低。干旱胁迫条件下土壤C/N、C/P、转化酶、脲酶和碱性磷酸酶活性的降低可能表明较低的N和P矿化速率。尽管微生物生物量C、N和P对一个生长季干旱胁迫处理无显著反应,但微生物生物量C和N在两年连续干旱胁迫后显著降低。土壤有机C和有效P含量在轻度施N处理下大于重度施N处理,但是有效N含量随着施N强度的增加而增加。微生物生物量C和N、碱性磷酸酶和转化酶活性也在轻度施N处理下有所增加。但是碱性磷酸酶活性在重度施N处理下降低。 5)野外条件下白刺花幼苗生长特征及生长土壤生化特性对施N的适应植物生长、生物生产量、C的固定、N、P等资源的吸收和积累、其它受限资源的利用效率(如P)在轻度施N处理下均有所增加,但N利用效率有所降低。幼苗生物生产量及C、N和P等资源的分配格局在轻度施N处理下也没有明显的改变。白刺花幼苗叶片数目、生物生产量和C、N、P的积累量在重度施N处理下虽然也相对于对照有所增加,但幼苗根系长度显著降低。生物量及资源(生物量、C、N、P)在重度施N处理下较多地分配给地上部分(主要是叶片)。另外,土壤有机C、全N和有效N含量随外源施N的增加而显著增加,土壤pH随之降低,但土壤全P含量并无显著反应。其中有机C含量和有效P含量以轻度施N处理最高。微生物生物量C、N和P在轻度施N处理下也显著增加,而微生物生物量C在重度施N处理下显著降低。同时,转化酶、脲酶、碱性磷酸酶和中性磷酸酶活性在施N处理下也明显的提高,但酸性磷酸酶和过氧化氢酶活性显著降低,其中碱性磷酸酶和中性磷酸酶活性以轻度施N处理最高。 综合分析表明,干旱河谷水分和N严重限制了白刺花幼苗的生长。施N不能完全改变干旱胁迫对白刺花幼苗的抑制的作用,但是由于施N增加土壤N有效性,改善土壤一系列生物与化学过程,幼苗的生长特性也对施N表现出强烈的反应,表现为植物结构与资源分配格局的改善,植物叶片光合能力与效率的提高,植物生长以及利用其他受限资源(如水分和P)的效率的增加,致使植物自身生长及其生长环境在干旱环境下得到改善。但是过度施N不仅不能起到改善干旱胁迫下植物生长环境、促进植物生长的作用,反而在土壤过程以及植物生长过程中加重干旱胁迫对植物的伤害。因此,建议在采用白刺花作为先锋种改善干旱河谷系统环境的实践中,可适当施加N以改善土壤环境,调节植物利用与分配资源的效率,促进植物定居,得到人工促进种群更新的目的。但在实践过程中也要避免过度施N。 Arid regions of the world are generally noted for their low primary productivity which is due to a combination of low, unpredictable water supply and low soil nutrient concentrations. The most serious effects of global climate change and human disturbances may well be those which related to increasing drought since drought stress has already been the principal constraint in plant growth. The decline in total rainfall and/or soil water availability expected for the next decades may turn out to be even more drastic under future warmer conditions. Nevertheless, water deficit is not the only limiting factor in arid and semiarid environments. Soils often suffer from nutrient (especially N and P) deficiencies in these ecosystems, which can also be worsened by climate change. How to improve the poor soil quality and enhance the vegetation coverage is always the problem facing ecosystem managers. The adaptive mechanisms of native plant to future climate change is always the focus in plant ecology, it also plays important roles in improving vegetation coverage by manual controlled programmes. Sophora davidii is a native perennial shrub of arid valleys, which is often predominant on eroded slopes and plays a vital role in retaining ecological stability in this region. It has been found that S. davidii was better adapted to dry environment than other shrubs, prompting its use for re-vegetation of arid lands. A two-years greenhouse experiment and a field experiment were conducted in order to understand the adaptation responses of Sophora davidii seedlings to different water and N conditions, and further explore if additional N supply as a modified role could enhance the adaptation ability of S. davidii seedlings to dry and infertile environment. Two-month old seedlings were subjected to a completely randome design with three water (80%, 40% and 20% water field capacity (FC)) and three N supply (N0: 0, Nl: 92 and Nh: 184 mg N kg-1 soil) regimes. Field experiment was arranged only by three N supplies in the dry valley. 1) The growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings in respond to drought stress and N supply Seedlings height, basal diameter, leaf number, leaf area, root length, biomass production, relative water content (RWC) and WUE were decreased with increase of drought stress. An increase in below-ground biomass was observed indicating a higher root/shoot ratio (R/S) under drought stress conditions. Low N supply increased seedlings height, basal diameter, leaf number, leaf area, and biomass production, but decreased root length. In contrast, these growth characteristics showed little or negative effect to high N supply treatment. Leaf percentages increased with increase of N supply, but fine root percentages decreased. In addition, Low N supply rather than the other two N treatments increased leaf area ratio (LAR), leaf/fine root mass ratio (L/FR), R/S and RWC under severe drought stress (20%FC), even though these parameters could increase with the high N supply treatment under well-watered condition (80%FC). Moreover, Low N supply also increased WUE under three water conditions, but high N supply had little effect on WUE under drought stress conditions (40%FC and 20%FC). 2) Leaf gas exchange and fluorescence parameters of Sophora davidii seedlings in respond to drought stress and N supply Leaf area (LA), photosynthetic pigment contents, and photosynthetic efficiency were decreased with increase of drought stress, but specific leaf area (SLA) increased. Photodamage in photosystem 2 (PS2) was also observed under drought stress condition. The decreased net photosynthetic rate (PN) under relative well-watered water conditions might result from stomatal limitations, but the decreased PN under other hand, photosynthetic capacity by increasing LA, photosynthetic chlorophyll contents, Pnmax, CE, Jmax were increased with increase N supply, and photosynthetic efficiency was improved with N supply treatment under water deficit. Although N supply did a little in alleviating photodamages to PS2 caused by drought stress, low N supply enhanced qN and kept relative high Fv/Fm under drought stress condition. However, high N supply inhibited leaf photosynthetic efficiency, and declined Fv/Fm and qN under severe drought stress condition after two year continues drought stress and N supply. 3) Carbon accumulation, nitrogen and phosphorus use efficiency of Sophora davidii seedlings in respond to drought stress and N supply C, N and P accumulation, NUE , N and P uptake efficiency (NUtE and NUtE ) P N P were decreased with increase of drought stress regardless of N supply. On the other hand, the S. davidii seedlings exhibited strong responses to N supply, but the responses were inconsistent with the various N supply levels. Low N supply rather than the other two N treatments increased C, N and P accumulation, improved NUEP, NUtE and NUtE under corresponding water condition. In contrast, high N supply N P did few even depressed effects on C, N and P accumulation, and NUEP, although NUtEN and NUtEP could increase with high N supply under corresponding water conditions. Even so, a decrease of NUEN was observed with increase of N supply under corresponding water conditions. 4) Soil microbial and chemical characters in respond to drought stress and N supply The content of soil organic C, available N and P were decreased with increase of drought stress. Decreases in C/N and C/P, and invertase, urea and alkaline phosphatase activity were also observed under drought stress conditions, indicating a lower N and P mineralization rate. Although microbial biomass C, N and P showed slight responses to drought stress after one growth period treatment, microbial biomass C and N were also decreased with increase of drought stress after two year continuous treatment. The content of soil organic C and available P showed the stronger positive responses to low N supply than which to high N supply, although than the other two N treatments increased microbial biomass N and invertase activity under severe drought stress condition, even though invertase activity could increase with high N supply treatment under relative well-water conditions. Moreover, low N supply treatment also increased C/P and alkaline phosphatase activity which might result from higher P mineralization, but high N supply did negative effects on alkaline phosphatase activity. 5) The growth characteristics of Sophora davidii seedlings and soil microbial and chemical characters in respond to N supply under field condition Low N supply facilitated seedlings growth by increasing leaf number, basal diameter, root length, biomass production, C, N and P accumulation and absorption, and enhancing the use efficiency of other limited resources as P. Compared to control, however, low N supply did little effect on altering biomass, C, N and P portioning in seedlings components. On the contrary, high N supply treatment also increased leaf number, biomass and C, N and P accumulation relative to control, but significantly decreased root length, and altered more biomass and resources to above-ground, which strongly reduced the ability of absorbing water under drought condition, and thus which might deep the drought stress. In addition, N supply increased soil C, N and available N content, but declined pH and showed little effects on P content. Low N supply showed higher values of soil C and available P content. Low N supply also increased microbial biomass C, N and P, although high N supply decreased microbial biomass C. N supply significantly enhanced soil invertase, urea, alkaline and neutral phosphratase activity, while declined acid phosphratase and catalase activity. Low N supply exhibited higher alkaline and neutral phosphratase activity compared to the others. The results from this study indicated that both drought and N limited the growth of S. davidii seedlings and their biomass production. Regardless of N supply levels, drought stress dramatically reduced the seedlings growth and biomass production. Although plant growth parameters, including basal diameter, height, leaf number, and biomass and their components were observed to be positive responses to low N supply, N supply alone can not alter the diminishing tendency which is caused by drought. available N content increased with increase N supply. In addition, low N supply rather These findings imply that drought played a primary limitation role and N was only the secondary. Even so, appropriate N supply was seemed to enhance the ability that S. davidii seedlings adapted to the xeric and infertile environment by improving soil processes, stimulating plant growth, increasing recourses accumulation, enhancing use efficiency of other limited resources, and balancing biomass and resources partitioning. Appropriate N supply, therefore, would be recommended to improve S. davidii seedling establishment in this region, but excess N supply should be avoided.
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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.