276 resultados para ELECTROLYTE


Relevância:

10.00% 10.00%

Publicador:

Resumo:

本文以复苏植物牛耳草成熟植株的离体叶片为实验材料,以光合作用、蔗糖、抗氧化剂系统和离子渗漏等在脱水复苏过程中的变化为切入点,从生理生化水平上探讨其耐脱水复苏的机制;同时应用mRNA差异显示技术,从分子水平上探讨其耐脱水复苏的机制。 牛耳草叶片光系统II光化学活性参数和叶黄素循环色素在脱水复苏过程中的变化结果表明,极微弱光强(3μmol.m-2.s-1)下,脱水8天的牛耳草叶片诱导了叶黄素循环,叶黄素循环可能介导了牛耳草叶片脱水过程中的光保护作用。 利用不同浓度的磷酸盐溶液处理牛耳草叶片的结果表明,0.1mol/L以上的磷酸盐溶液对牛耳草叶片具有损伤作用,极大的影响了其光系统II的光化学活性,使得牛耳草叶片在脱水后不能很好的复苏。 牛耳草叶片在脱水复苏过程中,抗坏血酸(AsA)、还原型谷胱甘肽(GSH)和蔗糖含量在脱水时很快增加,复苏时又迅速恢复到原来水平,表明它们可能对脱水的牛耳草叶片具有保护作用,但对复苏的牛耳草叶片可能不重要;其离子渗漏情况表明质膜结构的完整性和稳定性在脱水复苏过程中能得到很好的保持,这可能是其耐脱水复苏的重要机制之一。 利用mRNA差异显示技术分离到牛耳草叶片脱水过程中一些脱水和磷酸盐特异诱导表达的cDNA。对其中5个脱水特异诱导表达和3个磷酸盐特异诱导表达的cDNA进行克隆测序、同源性探测和Northern 杂交检测表明,牛耳草脱水过程中诱导表达的基因可能涉及到脱水胁迫的信号转导、调节基因的级联和结构基因产物调节细胞结构在脱水胁迫中的稳定性等。

Relevância:

10.00% 10.00%

Publicador:

Resumo:

Depth profiles of carrier concentrations in GaMnSb/GaSb are investigated by electrochemistry capacitance-voltage profiler and electrolyte of Tiron. The carrier concentration in GaMnSb/GaSb measured by this method is coincident with the results of Hall and X-ray diffraction measurements. It is indicated that most of the Mn atoms in GaMnSb take the site of Ga, play a role of acceptors, and provide shallow acceptor level(s).

Relevância:

10.00% 10.00%

Publicador:

Resumo:

The photovoltaic spectral features and the behaviors of photocurrent versus the electrode potential for near surface In0.15Ga0.85As/GaAs quantum well electrodes have been investigated in nonaqueous solutions of ferrocene and acetylferrocene. The photovoltaic spectrum shows a sharp structure that reflects confined state-to-state exciton transition in the quantum well. Deep dips are observed in the photocurrent versus the electrode potential curves in both electrolytes at the different electrode potentials under the illumination of exciton resonance wavelength. These dips are qualitatively explained by considering the interfacial tunneling transfer of photogenerated electron within the quantum well.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

Poly(3,4-ethylenedioxythiopliene):poly(styrene sulfonate) (PEDOT:PSS) films have been electrochemically polymerized in situ on ITO glass substrate in boron trifluoride diethyl etherate electrolyte (BFEE). Cyclic voltammograms show good redox activity and stability of the PEDOT films. These films had been directly used to fabricate organic-inorganic hybrid solar cells with the structure of ITO/PEDOT/ZnO:MDMC-PPV/Al. The solar cells made of electrochemically polymerized films exhibit higher energy conversion efficiencies compared with that prepared by the spin-coating method, and the highest value is 0.33%. This in-situ electropolymerized method effectively simplifies fabricating procedures and may blaze a facile and economical route for producing high-efficiency solar cells.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

A thin oriented bacteriorhodopsin (bR) him is deposited on a stainless steel slide by use of the electrophoretic sedimentation method. A junction is made with electrolyte gels having a counterelectrode to construct a bR-based photoelectric detector;. The photoelectric response signal to a 10 ns laser pulse is measured. A theory on the photoelectric kinetics of bR is developed based on the concept of the charge displacement current and the bR photocycle rate equations. Comparison between the theoretical and experimental results proves that the bR photoelectric response to a short laser pulse is a multi-exponential process. The decay time constants and amplitudes of each, exponential component are obtained by data fitting.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

聚合物电解质在制备高能密度全固态铿电池、光电化学器件、气体传感器和电化学半导体等器件显示了重要的应用前景,成为近期的研究热点之一。然而,人们目前对聚合物电解质中离子传输、导电机理及电极界面动力学等问题的了解有限,这严重阻碍高性能聚合物电解质的研究和开发。本论文研究工作主要集中在纳米复合聚合物电解质中离子传输和导电机理的研究,内容和创新点表现在以下几个方面:1、设计制备了形状可控的纳米级金微带电极和铂、碳纤维微盘阵列电极。操作相对简单,在普通实验室中就可以制备,电极通过研磨就可以重复多次使用。2、选择四种不同分子量的PEG作为聚合物本体,杂多酸纳米粒子作为添加剂,和铿盐组成了有机一无机纳米复合聚合物电解质。用微盘电极的计时电流方法求出了杂多酸纳米粒子在聚合物电解质中的扩散系数,利用DSC技术和FT-IR光谱解释了离子之间相互作用的机理和电导率提高的原因。3、合成出不同侧链长的梳状丙烯酸铿/甲基丙烯酸聚乙二醇单甲醚醋聚合物电解质。用DSC,TGA和交流阻抗研究了纳米SiO2掺杂的梳状聚合物电解质的电化学和热力学性质。给出了铿离子在纳米复合聚合物电解质中随纳米SiO2含量变化的传输模型。黑抉巍一淤呱4、将聚环氧乙烷,盐和多壁碳纳米管掺杂在一起,制备出一种离子一电子混合导电的聚合物电解质纳米杂化薄膜。用贫流阻抗和DSC技术表征了杂化薄膜的导电性质和热力学行为,给出了体系的等效电路,解释了电导率增加的原因。5、将多壁碳纳米管和碳微珠分别和室温离子液体混合制备了两种无机/有机的纳米复合材料:多壁碳纳米管/离子液体和碳微珠/离子液体。利用交流阻抗技术研究了无机/有机的纳米复合材料的导电机理。用这两种复合材料制备的酶电极可以获得酶的直接电化学,且能够催化O2和H2O2的还原。

Relevância:

10.00% 10.00%

Publicador:

Resumo:

由极性聚合物-碱金属盐构成的聚合物固体电解质(SPE)是一类很有前途的离子导电材料。但此类SPE,比通常非水介质电解质溶液约低3个数量级,故极需发展电导率更高的新型SPE材料。目前,由极性(或交联)聚合物、碱金属盐和高介电常数溶剂三组份构成的新一代SPE正在崛起,其室温电导率的达10~(-3)S·cm~(-1)。本工作以甲基丙烯酸多缩乙二醇二酯为大分子单体,研究了三组份凝胶电解质的制备、各组分的变化和温度等对膜强度和电导率的影响;并研究了该凝胶体系中离子传导机理。

Relevância:

10.00% 10.00%

Publicador:

Resumo:

Three causes involved in the instability of the ISFET are proposed in this study. First, it is ascertained that hydroxyl group resident at the surface of the Si3N4 film or in the electrolyte solution is most active and subject to gain or loss of electrons. This is one of the main causes for ISFET structural instability. Secondly, the stability of the pH-sensitive FET varies with deposition conditions in the fabrication process of the ISFET. This proves to be another cause of ISFET instability. Thirdly, the pH of the measured solution varies with the measuring process and time, contributing to the instability, but is not a cause of the instability of the pH-ISFET itself. We utilized the technique of readjusting and controlling the ratio of hydroxyl groups to amine groups to enhance the stability of the ISFET. Our techniques to improve stability characteristics proved to be effective in practice.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

The photoelectric properties of the lattice-matched GaAs/AlxGa1-xAs quantum well electrodes and the influence of the electrode structure such as well width, the thickness of outer barrier and the number of period were studied in a nonaqueous electrolyte. A new kind of structure of multiple quantum well electrode with varied well width, possessing the quantum yield three times that of GaAs bulk materials, was designed and fabricated.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

The interfacial behavior of the single quantum well (SQW) GaAs/AlxGa1-xAs electrode in HQ/BQ and Fc/Fc(+) electrolytes was characterized respectively by studying the quantum confined Stark effect and Franz-Keldysh oscillation with electrolyte electroreflectance spectroscopy. The interaction of the surface state of the SQW electrode with redox species and its effects on the distribution of external bias at the interface of the SQW electrode are discussed.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

Transient photocurrents induced by short light pulses at lattice-matched GaAs/AlxGa1-xAs multiple quantum well (MQW) electrodes were studied as a function of electrode potential. Dual exponential photocurrent decay transients were observed at various potentials. By analysis of the dual exponential decay transients, information on steady state photocurrents (I-s), surface collection of photoexcited minority carriers (G(0)) and lifetimes of surface states (T-s) was obtained. The kinetic behaviors of photoprocesses at illuminated MQW/electrolyte interface were discussed.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

A monolithic integrated CMOS preamplifier is presented for neural recording applications. Two AC-coupied capacitors are used to eliminate the large and random DC offsets existing in the electrode-electrolyte interface. Diode-connected nMOS transistors with a negative voltage between the gate and source are candidates for the large resistors necessary for the preamplifier. A novel analysis is given to determine the noise power spectral density. Simulation results show that the two-stage CMOS preamplifier in a closed-loop capacitive feedback configuration provides an AC in-band gain of 38.8dB,a DC gain of 0,and an input-referred noise of 277nVmax, integrated from 0. 1Hz to 1kHz. The preamplifier can eliminate the DC offset voltage and has low input-referred noise by novel circuit configuration and theoretical analysis.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

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.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

本研究通过粗枝云杉不同种群进行的温室半控制试验,采用植物生态学、生理学和生物化学的研究方法,系统地研究了粗枝云杉不同种群抗旱性的生长、形态、生理和生化机理,并结合有关研究进行综合分析,得出主要研究结论如下: 1.粗枝云杉对干旱胁迫的综合反应 粗枝云杉在干旱胁迫下的适应机制为:(1)相对生长速率及植株结构的调整:干旱胁迫下虽然植株相对生长速率显著降低,且有相对较多的生物量向根部分配,但并未发现细根/总根比增加。(2)粗枝云杉对干旱胁迫的光合作用表现为:干旱胁迫显著地降低了控制的理想条件下的气体交换,但干旱胁迫对PSII最大光化学效率(Fv/Fm)没有影响,表明干旱并未影响到光合机构。(3)干旱还影响了很多生理生化过程,包括渗透调解物质(游离脯氨酸)、膜脂过氧化产物、脱落酸(ABA)含量的增加,以及保护酶活性的升高。这些结果证明植物遭受干旱胁迫后发生了一系列的形态、生理和生化响应,这些变化能提高干旱时期植物的存活和生长能力。 2.粗枝云杉不同种群对干旱胁迫反应的种群差异 粗枝云杉三个种群-干旱种群(四川丹巴和甘肃迭部)和湿润种群(四川黑水)对干旱适应不同,这种不同应归因于它们采用的用水策略不同:在水分良好和干旱胁迫条件下,受试种群在相对生长速率和水分利用效率(WUE)方面都表现出显著的种群间差异。与湿润种群相比,干旱种群在两种水分条件下有更高的WUE。粗枝云杉不同种群的碳同位素组分(δ13C)只在干旱胁迫下有显著差异,并且这种差异在水分良好时比干旱胁迫条件下小,说明生理响应和干旱适应性之间的关系受植物内部抗旱机制和外部环境条件(如水分可利用性)或两者互作效应的影响。这些结果说明干旱种群和湿润种群所采用的用水策略不同。干旱种群有更强的抗旱能力,采用的是节水型的用水策略,而湿润种群抗旱能力较弱,采用的是耗水型的用水策略。 3. 遮荫对粗枝云杉不同种群抗旱性影响 干旱胁迫显著降低了全光条件下叶相对含水量(RWC)、相对生长速率、气体交换参数、PSII的有效量子产量(Y),提高了非光化学猝灭效率(qN)、水分利用效率、脯氨酸(PRO)积累、脱落酸(ABA)含量及保护酶活性。然而这种变化在遮荫条件下不明显。我们得出结论适度遮荫降低了干旱对植物的胁迫作用。另一方面,在干旱条件下,与湿润种群相比,干旱种群抗旱性更强,表现在干旱种群净光合速率与单位重量上叶氮含量(Nmass)降低较少。另外,干旱种群表现出更为敏感的气孔导度,更高的热耗散能力(qN)能力、用水效率、ABA积累、保护酶活性,以及更低的总用水量、相对生长速率。这一结果表明这两种群采用不同的生理策略对干旱和遮荫做出反应。许多生长和生理反应差异与这两个种群原产地气候条件相适应。 4. 外源脱落酸(ABA)喷施对粗枝云杉不同种群抗旱性影响 外源ABA喷施在干旱和水分良好条件下均不同程度地提高了根/茎比,表明根和茎对ABA敏感程度不同。实验结果还表明,外源ABA喷施对这两个种群在干旱胁迫期间影响不同。干旱胁迫期间,伴随着ABA喷施,湿润种群净光合速率(A)显著降低,而干旱种群净光合速率变化不明显。另一方面,外源ABA喷施显著提高了干旱条件下干旱种群的单位叶面积重(LMA)、根/茎比、细根/总根(Ft)比、水分利用效率(WUE)、ABA含量, 以及保护酶活性。然而,外源ABA喷施对湿润种群的上述测定指标没有显著影响。这一结果表明干旱种群对外源ABA喷施更为敏感, 反应在更大的气孔导度降低,更高的生物量可塑性,及更高的水分利用效率、ABA含量和保护酶活性。综上所述,我们得出结论,粗枝云杉对外源ABA敏感性因种群的不同而不同。该研究结果可为两个明显不同种群在适应分化方面提供强有力的证据。 Arid or semi-arid land covers more than half of China's land territory. In arid systems, severe shortages of soil water often coincide with periods of high temperatures and high solar radiation, producing multiple stresses on plant performance. Protection from high radiation loads in shaded microenvironments during drought may compensate for a loss of productivity due to reduced irradiance when water is available. Additionally, ABA, a well-known stress-inducible plant hormone, has long been studied as a potential mediator for induction of drought tolerance in plants. Picea asperata Mast., which is one of the most important tree species used for the production of pulp wood and timber, is a prime reforestation species in western China. In this experiment, different population of P. asperata were used as experiment material to study the adaptability to drought stress and population differences in adaptabiliy, and the effects of shade and exogenous abscisic acid (ABA) application on the drought tolerance. Our results cold provide a strong theoretical evidence and scientific direction for the afforestation, and rehabilitation of ecosystem in the arid and semi-arid area, and provide a strong evidence for adaptive differentiation of different populations, and so may be used as criteria for species selection and tree improvement. The results are as follows: 1. A large set of parallel response to drought stress Drought stress caused pronounced inhibition of the growth and increased relatively dry matter allocation into the root; drought stress also caused pronounced inhibition of photosynthesis, while drought showed no effects on the maximal quantum yield of PSII photochemistry (Fv/Fm) in dark-adapted leaves, indicating that drought had no effects on the primary photochemistry of PSII. However, in light-adapted leaves, drought reduced the quantum yield of PSII electron transport (Y) and increased the non-photochemical quenching (qN). Drought also affected many physiological and biochemical processes, including increases in superoxide dismutase (SOD), ascorbate peroxidase (APX) activities, malondialdehyde and ABA content. These results demonstrate that there are a large set of parallel changes in the morphological, physiological and biochemical responses when plants are exposed to drought stress; these changes may enhance the capability of plants to survive and grow during drought periods. 2. Difference in adaptation to drought stress between contrasting populations of Picea asperata There were significant population differences in growth, dry matter allocation and water use efficiency. Compared with the wet climate population (Heishui), the dry climate population (Dan ba and Jiebu) showed higher LMA, fine root/total root ratio and water use efficiency under drought-stressed treatments. The results suggested that there were different water-use strategies between the dry population and the wet population. The dry climate population with higher drought tolerance may employ a conservative water-use strategy, whereas the wet climate population with lower drought tolerance may employ a prodigal water-use strategy. These variations in drought responses may be used as criteria for species selection and tree improvement. 3. The effects of shade on the drought tolerance For both populations tested, drought resulted in lower needle relative water content (RWC), relative growth rate (RGR), gas exchange parameters and effective PSII quantum yield (Y), and higher non-photochemical quenching (qN), water use efficiency (WUE), proline (PRO) and abscisic acid (ABA) accumulation, superoxide dismutase (SOD), ascorbate peroxidase (APX) activities as well as malondialdehyde (MDA) levels and electrolyte leakage in sun plants, whereas these changes were not significant in shade plants. Our study results implied that shade, applied together with drought, ameliorated the detrimental effects of drought. On the other hand, compared with the wet climate population, the dry climate population was more tolerant to drought in the sun treatment, as indicated by less decreases in A and mass-based leaf nitrogen content (Nmass), more responsive stomata, greater capacity for non-radiative dissipation of excitation energy as heat (analysed by qN), and higher WUE,higher level of antioxidant enzyme activities,higher ABA accumulation as well as lower MDA content and electrolyte leakage. Many of the differences in growth and physiological responses reported here are consistent with the climatic differences between the locations of the populations of P. asperata. 4. The effects of exogenous abscisic acid (ABA) application on the drought tolerance For both populations tested, exogenous ABA application increased root/shoot ratio (Rs) under well-watered and drought-stressed conditions, indicating that there was differential sensitivity to ABA in the roots and shoots. However, it appeared that ABA application affected the two P. asperata populations very differently during drought. CO2 assimilation rate (A) was significantly decreased in the wet climate population, but only to a minor extent in the dry climate population following ABA application during soil drying. On the other hand, ABA application significantly decreased stomatal conductance (gs), transpiration rate (E) and malondialdehyde (MDA) content, and significantly increased leaf mass per area (LMA), Rs, fine root/total root ratio (Ft), water use efficiency (WUE), ABA contents, superoxide dismutase (SOD), ascorbate peroxidase (APX) and catalase (CAT) activities under drought condition in the dry climate population, whereas ABA application did not significantly affect these parameters in the wet population plants. The results clearly demonstrated that the dry climate population was more responsive to ABA application than the wet climate population, as indicated by the strong stomata closure and by greater plasticity of LMA and biomass allocation, as well as by higher WUE, ABA content and anti-oxidative capacity to defense against oxidative stress, possibly predominantly by APX. We concluded that sensitivity to exogenous ABA application is population dependent in P. asperata. Our results provide strong evidence for adaptive differentiation between populations of P. asperata.

Relevância:

10.00% 10.00%

Publicador:

Resumo:

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.