995 resultados para Gentiana Sect Stenogyne


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In this paper we introduce a new Half-flash analog switch ADC architecture. And we discuss two methods to design the values of the cascaded resistors which generate the reference voltages. Derailed analysis about the effect of analog switches and comparators on reference voltages, and the methods to set the resistor values and correspond;ng voltage errors are given.

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This paper introduces a new highspeed single-way analog switch which has both highspeed high-resolution mono-direction analog transmission gate function and high-speed digital logic gate function with normal bipolar technology. The analysis of static and transient switching performances as an analog transmission gate is emphasized in the paper. In order to reduce the plug-in effect on high-speed high-resolution systems, an optimum design scheme is also given. This scheme is to achieve accelerated dynamic response with very low bias power dissipation. The analysis of PSPICE simulation as well as the circuit test results confirms the feasibility of the scheme. Now, the circuit has been applied effectively to the designs of novel highspeed A/D and D/A converters.

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In this paper, a one-way NMOS analog switch featuring a low plug-in consumption is presented. The performances of analog switch, especially the performances of source follower are simulated under different conditions with PSPICE. Simulation results and factors affecting the deviation between input and output are analyzed, some advice on how to reduce the deviation between input and output is given. Ar the end of the paper, voltage relationship between input and output of the analog switch is obtained. Function of first degree, Vout = kVin + V0, is used to approximate the voltage relationship. The simulation results anti the value achieved from the approximation equation are given as well.

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Self-organized InAs quantum; dots sheets are grown on GaAs(100) substrate and tapped by 80nm GaAs layer with molecular beam epitaxy. Samples were annealed and characterized with Raman spectra, transmission electron microscopy (TEM) and photolumincscence (PL). The Raman spectra indicates arsenic clusters in the GaAs capping layer. The TEM analysis revealed the relaxation of strain in some InAs islands with the introduction of the network of 90 dislocations. In addition, the structural changes also lead to the changes of the PL spectra from me InAs islands. Their correlation was discussed, Our results suggest:est that annealing may be used to intentionally modify me properties of self-organized InAs islands on GaAs.

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This paper presents a detailed PL study of Fe2+ related four zero-phonon(ZP) lines and their related phonon sidebands. Four zero-phonon transitions at approximate to 2800 cm(-1) along with the accompanying phonon sidebands extending down to 2400 cm(-1). There are ta two prominent regions in the phonon sidebands. One is ascribed to coupling to acoustic-type phonons (2700 cm(-1) region), the other is due to coupling to optic-type phonons (2500 cm(-1) region). Beside broad coupling with lattice modes, there are several groups of lines. They are ascribed to resonant modes, impurities induced gap modes and local modes.

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Quantum dot lasers are predicted to have proved lasing characteristics compared to quantum well and quantum wire lasers. We report on quantum dot lasers with active media of vertically stacked InAs quantum dots layers grown by molecular beam epitaxy. The laser diodes were fabricated and the threshold current density of 220 A/cm(2) was achieved at room temperature with lasing wavelength of 951 nm. The characteristic temperature To was measured to be 333K and 157K for the temperature range of 40-180K and 180-300K, respectively.

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This paper gives a condition for the global stability of a continuous-time hopfield neural network when its activation function maybe not monotonically increasing.

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An improved BP algorithm for pattern recognition is proposed in this paper. By a function substitution for error measure, it resolves the inconsistency of BP algorithm for pattern recognition problems, i.e. the quadratic error is not sensitive to whether the training pattern is recognized correctly or not. Trained by this new method, the computer simulation result shows that the convergence speed is increased to treble and performance of the network is better than conventional BP algorithm with momentum and adaptive step size.

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Self-assembled InAs quantum wires (QWRs) embedded in In0.52Al0.48As In0.53Ga0.47As, and (In0.52Al0.48As)(2)/(In(0.53)Ga(0.47)AS)(2)-short-period-lattice matrixes on InP (001) were fabricated with molecular beam epitaxy (MBE). These QWR lines are along [110], x4 direction in the 2x4 reconstructed (001) surface as revealed with high energy electron diffraction (RHEED). Alignment of quantum wires in a multilayer structure depends on the composition of spacer layers.

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nThermal processing of strained ln(0.2)Ga(0.8)As/GaAs graded-index separate confinement heterostructure single quantum well laser diodes grown by molecular beam epitaxy is investigated. It was found that rapid thermal annealing can improve the 77 K photoluminescence efficiency and electron emission from the active layer, due to removal of nonradiative centers from the InGaAs/GaAs interface. Because of the interdiffusion of Al and Ga atoms, rapid thermal annealing increases simultaneously the density of DX centers in the AlGaAs graded layer. The current stressing experiments of post-growth and annealed laser diodes are indicative of a corresponding increase in the concentration of DX centers, suggesting that DX centers may be responsible for the degradation of laser diode performance.

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本文在前人工作的基础上,对东北产千里光属Senecio植物进行了形态分类、花解剖及花粉形成结构的研究。结果表明:1. 华千里光组Sect. Sinosenecio和狗舌草组sect. Tephroseris在形态和花结构上与千里光属Senecio植物有显著差异,应从千里光属中独立出来,自立为属,即:华千里光属Sinosenecio对狗舌草属Tephroseriso 2. 东北产华千里光属只有一种,即朝鲜千里光Sinosenecio Koreanus (Kom.) B. Nord。3. 东北产狗舌草属分为2组,共5种1变型,其中T. Campestris (Retz.) Reichb.f.spathulatus (Miq.) R. yin et C. y. Li为新组合,并订正了狗舌草T. campestris (Retz.) Reichb的拉丁名。4. 东北产千里光属分为2组,共5种,2变种1变型。5. 狗舌草属和千里光属花粉形态基本相似,都为球形,表面具刺,3孔沟型。花粉形成不能作为划分属的依据。本文还详细描述了每个种的形态学特征,地理分布和文献考证,并附有每个种的叶形。花解剖图及花粉的扫描电镜、光镜照片。还有分属及分种检索表。

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瑞香狼毒(Stellera chamaejasme L.)是瑞香科(Thymelaeaceae)狼毒属的一种多年生野草,有毒。据调查,从20 世纪60 年代开始至今,狼毒在青藏高原东缘的高寒草甸上不断蔓延、密度不断变大,在一些地段甚至成为优势物种。有关狼毒在高寒草甸蔓延的生态系统效应的研究尚未见报道。本文从系统碳、氮循环的角度,分别研究狼毒在生长和非生长季节对高寒草甸生态系统的影响。同时,从花粉化感的角度,深入研究狼毒对当地同花期物种有性繁殖的影响。系统地研究高寒草甸生态系统物质循环过程,特别是非生长季节微生物和土壤碳氮库的动态变化,有助于揭示狼毒在系统物质循环方面的“物种效应”以及这种效应的季节变化,为丰富有关高海拔生态系统,特别是其非生长季的物质循环的科学理论做出贡献。同时,碳氮循环和花粉化感的研究还有助于深刻地理解狼毒作为一种入侵性很强的杂草的特殊的蔓延机制,从而为狼毒的有效防治、高寒草甸的科学管理提供依据。 针对狼毒在青藏高原高寒草甸上蔓延的生态系统碳氮循环方面的影响,开展以下2 方面的研究:(1)在生长季,研究松潘县尕米寺附近(北纬32°53',东经103°40',海拔3190 m)的两种地形(平地和阳坡)条件下狼毒对土壤碳氮循环影响及可能的原因。狼毒和其它几个主要物种(圆穗蓼(Polygonummacrophyllum D. Don var. Macrophyllum),草地早熟禾(Poa pretensis L.),四川嵩草(Kobresia setchwanensis Hand.-Maizz.),鹅绒委陵菜(Potentilla anserina L.var. anserine)和鳞叶龙胆(Gentiana squarrosa Ledeb.)的地上凋落物产量以及地上凋落物和根的化学组成被测量。在有-无狼毒斑块下,各种土壤的库(比如,铵态氮、硝态氮、无机磷和微生物生物量)和周转率(包括净矿化、净硝化、总硝化、反硝化和微生物呼吸速率)被测量和比较。(2)在非生长季节,尤其是春季冻融交替期,选取了两个研究地点——尕米寺和卡卡沟(北纬32°59',东经103°41',海拔3400 m),分别测定有狼毒和无狼毒斑块下土壤微生物生物量碳和氮、可溶性有机碳和氮以及铵态氮和硝态氮的动态变化。同时,分别在上述两个地点有-无狼毒的样地上,研究6 个主要物种(狼毒、圆穗蓼、草地早熟禾、四川嵩草、鹅绒委陵菜和鳞叶龙胆)从秋季开始、为期1 年的凋落物分解过程。 针对狼毒花粉化感对同花期其它物种可能的花粉化感作用开展以下工作:在实验室中,用一系列浓度的狼毒花粉水浸提液对与它同花期的其它物种以及自身花粉进行测试,测定花粉萌发率;在野外自然条件下的其它物种的柱头上施用上述浓度的狼毒花粉水浸提液,观测种子结实率,同时,观察狼毒花粉的种间花粉散布数量。 生长季节的研究结果表明,狼毒地上凋落物氮含量比其它几个主要物种更高,而木质素-总氮比更低。狼毒显著地增加其斑块下表层土壤中有机质的含量,而有-无狼毒的亚表层土壤在有机碳和总磷方面没有显著差异。狼毒表土中硝态氮含量在平地和阳坡比无狼毒土壤分别高113%和90%。狼毒表土中微生物生物量碳和氮量显著高于无狼毒表土。无论是平地还是阳坡,狼毒土壤的总硝化和微生物呼吸速率显著高于无狼毒土壤;而它们的反硝化速率只在平地有显著的差异。狼毒与其它物种间地上凋落物的产量和质量的差异可能是导致有-无狼毒土壤碳氮循环差异的原因。我们假设,狼毒可能通过增加贫氮生态系统土壤中的有效氮含量提高其入侵能力。 非生长季的研究结果表明,在青藏高原东缘的高寒草甸上,土壤微生物生3物量在11 月的秋-冬过渡期达到第一个峰值;在春季的冻融交替期,微生物生物量达到第二个峰值后又迅速降低。无机氮以及可溶性有机碳氮与微生物生物量有相似的变化过程。微生物碳氮比呈现显著的季节性变化。隆冬季节的微生物生物量碳氮比显著高于生长旺季的微生物碳氮比。这种变化可能暗示冬、夏季微生物的群落组成和对资源的利用有所不同。有-无狼毒斑块下土壤微生物和土壤碳、氮库一般只在秋-冬过渡期有显著差异,有狼毒土壤微生物生物量和土壤碳、氮库显著高于无狼毒土壤;而在之后的冬季和春季没有显著差异。所有6 个物种凋落物在非生长季分解率为24%-50%,均高于生长季的10%到30%。其中在秋-冬过渡期,凋落物开始埋藏的两周时间内,分解最快,达10%-20%。不同物种凋落物全年的分解率和分解过程有显著差异。圆穗蓼在全年的分解都较缓慢(非生长季26%,生长季15%),草地早熟禾和四川嵩草等全年的分解速率比较均匀(非生长季和生长季均为30%,非生长季略高),而狼毒在非生长季分解较快(约50%),而在接下来的生长季分解变得缓慢(约12%)。所有物种的凋落物氮含量在非生长季下降,而在随后的生长季上升。 实验室的花粉萌发试验证明,狼毒花粉对自身花粉萌发没有自毒作用,而其它受试的所有物种(圆穗蓼,秦艽(Gentiana macrophylla Pall. var. fetissowii),湿生扁蕾(Gentianopsis paludosa (Hook. f.) Ma var. paludosa),鳞叶龙胆,椭圆叶花锚(Halenia elliptica D. Don var. elliptica),蓝钟花(Cyananthus hookeri C. B.Cl. var. grandiflorus Marq.),小米草(Euphrasia pectinata Ten.),川西翠雀花(Delphinium tongolense Franch.),高原毛茛(Ranunculus tanguticus (Maxim.)Ovcz. var. tanguticus)和鹅绒委陵菜)的花粉萌发率随着狼毒花粉浸提液浓度的增加呈显著的非线性降低。大约3 个狼毒花粉的浸提液就可以抑制受试的多数物种的50%的花粉萌发。在鳞叶龙胆和小米草柱头上狼毒花粉的数量分别为5.76 个和3.35 个。狼毒花粉散布数量的差异最可能的原因在于是否有共同的传粉昆虫。花的形状(辐射对称VS 左右对称)、植株或花的密度以及花期重叠性可以部分解释这种差异。在野外试验中,我们发现6 个物种(秦艽、湿生扁蕾、鳞叶龙胆、椭圆叶花锚、蓝钟花和小米草)的种子结实率随狼毒花粉浸提液浓度的增加呈显著的非线性降低。鳞叶龙胆和小米草柱头上狼毒花粉的数量(分别是5.76 个和3.35 个)分别达到了抑制它们63%和55%种子结实率的水平。因此,狼毒对鳞叶龙胆和小米草可能存在明显的花粉化感抑制作用。狼毒周围的物种可能通过花期在季节或昼夜上的分异避免受到狼毒花粉化感的影响或者通过无性繁殖来维持种群繁衍,因此狼毒通过花粉化感作用对其周围物种繁殖的影响程度还需要进一步地研究。如果狼毒的花粉化感抑制作用确实存在,那么它可能成为一种自然选择压力,进而影响物种的进化。 Stellera chamaejasme L., a perennial toxic weed, has emerged and quicklydominated and spread in the high-frigid meadow on the eastern Tibetan Plateau ofChina since the 1960s. In the present study, effects of S. chamaejasme on carbon andnitrogen cycles on the high-frigid meadow on the eastern Qinghai-Tibetan Plateau ingrowing and non-growing season, and its pollen allelopathic effects on the sympatricspecies were determined. The present study that focused on carbon and nitrogencycles, especially on microbial biomass and pools of carbon and nitrogen innon-growing season, could profoundly illuminate plant-species effects on carbon andnutrient cycles and its seasonal pattern and help to understand spread mechanism ofS. chamaejasme as an aggressive weed. The present study also contributed to furtherunderstand carbon and nutrient cycles on alpine regions in non-growing season andprovide a basis on weed control of S. chamaejasme and scientific management in thehigh-frigid ecosystem. Effects of S. chamaejasme on carbon and nitrogen cycles on the high-frigidmeadow on the eastern Qinghai-Tibetan Plateau were determined. The study couldbe divided into two parts. (1) In the growing season, we quantified the effects of S.chamaejasme on carbon and nitrogen cycles in two types of topographic habitats, theflat valley and the south-facing slope, where S. chamaejasme was favored to spreadlitter and root were measured to explain the likely effects of S. chamaejasme on soilcarbon and nutrient cycles. The sizes of various soil pools, e.g. nitrite, ammonium,inorganic phosphorus and microbial biomass, and turnover rates including netmineralization, gross nitrification, denitrification and microbial respiration weredetermined. (2) In the non-growing season study, microbial biomass carbon andnitrogen, soluble organic carbon and nitrogen, ammonium and nitrate weredetermined through the non-growing season, especially in the processes offreeze-thaw of spring in two high-frigid sites, i.e. Kaka valley and Gami temple, onthe eastern Qinghai-Tibetan Plateau. Meanwhile, litter decomposition of six commonspecies, including Stellera chamaejasme L., Polygonum macrophyllum D. Don var.Macrophyllum, Poa pretensis L., Kobresia setchwanensis L., Potentilla anserina L.var. anserine and Gentiana squarrosa Ledeb., were also examined under theabove-mentioned experimental design through one whole-year, which began in theautumn in 2006. In the study of pollen allelopathy, several work, including in vitro study oneffects of extract of pollen from S. chamaejasme on sympatric species and pollenfrom itself, field experiments on effects of pollen extract with the same regime ofconcentrations on seed set and field observation on heterospecific pollen transfer ofS. chamaejasme to six of those sympatric species has been done. The results in the growing season showed that aboveground litter of S.chamaejasme had higher tissue nitrogen and lower lignin: nitrogen ratio than thoseco-occurring species. S. chamaejasme significantly increased topsoil organic matter,whereas no significant differences were found for organic C and total P in subsoilbetween under-Stellera and away-Stellera locations. The nitrate in Stellera topsoilwas 113% and 90% higher on the flat valley and on the south-facing slope,respectively. Both microbial biomass C and N were significantly higher in Stelleratopsoil. Gross nitrification and microbial respiration were significantly higher inStellera topsoil both on the flat valley and on the south-facing slope, whereassignificant differences of denitrification were found only on the flat valley. Thedifferences in the quantity and quality of aboveground litter are a likely mechanismresponsible for the changes of soil variables. We assumed that S. chamaejasme couldenhance their spread by increasing nutrient availability in N-deficient ecosystems. The results in the non-growing season showed that microbial biomass achievedthe first summit in late autumn and early winter on the eastern Qinghai-TibetanPlateau. In the stages of freeze-thaw of spring, microbial biomass firstly achieved thesecond summit and subsequently sharply decreased. Inorganic nitrogen, solubleorganic carbon and nitrogen had a similar dynamics with that of microbial biomass.Ratio of microbial biomass carbon and nitrogen had an obviously seasonal pattern.The highest microbial C: N were in the non-growing season, which weresignificantly higher than those in the growing season. The seasonal pattern inmicrobial biomass C: N suggested that large changes in composition of microbialpopulation and in resources those used by microbes between summer and winter.Generally, microbial biomass and pools size of carbon and nitrogen in Stellera soilwere significantly higher than those under adjacent locations in late autumn andearly winter, but there were not significant differences in winter and in spring. Litterof all the focal species (Stellera chamaejasme L., Polygonum macrophyllum D. Donvar. Macrophyllum, Poa pretensis L., Kobresia setchwanensis Hand.-Maizz.,Potentilla anserina L. var. anserine and G. squarrosa Ledeb.) decomposed about24%-50% in the non-growing season, which were higher than those in the growingseason (ranged from 10% to 30%). Litter decomposed 10%-20% within the first twoweeks in late autumn and early winter. Significant differences in the whole-yeardecomposition rate and in the processes of decomposition were found among species.Polygonum macrophyllum decomposed slowly through the whole year (26% and15% in the non-growing season and in the growing season, respectively). Certainspecies, such as P. pretensis L. and K. setchwanensis, decomposed at a similar rate(30% both in the non-growing and in the growing season, slightly higher in the8growing season than those in the growing season), whereas S. chamaejasmedecomposed more rapidly (about 50%) in the non-growing season and subsequentlydecomposition became slow (about 12%) in the growing season. Litter nitrogencontents of all the focal species firstly decreased in the non-growing season and thenincreased in the growing season. In vitro experiments of pollen allelopathy, the results showed that pollen from S.chamaejasme was not autotoxic, whereas pollen germination in all the sympatricspecies (Polygonum macrophyllum D. Don var. Macrophyllum, Gentianamacrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Ma var. paludosa,Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica, Cyananthushookeri C. B. Cl. var. grandiflorus Marq., Euphrasia pectinata Ten., Delphiniumtongolense Franch., Ranunculus tanguticus (Maxim.) Ovcz. var. tanguticus andPotentilla anserina L. var. anserina) decreased nonlinearly as the increasingconcentrations of extract of pollen from S. chamaejasme. Pollen Extract of threepollens from S. chamaejasme generally inhibited 50% pollen germination of most ofthe focal species. 5.76 and 3.35 pollens from S. chamaejasme were observed in fieldon stigmas of G. squarrosa and E. pectinata, respectively. Differences inheterospecific pollen transfer of S. chamaejasme could be attributed to the primaryreason whether they shared common pollinators. Flower morphology (e.g.zygomorphic or actinomorphic), plant or floral density and concurrence in floweringphonologies could explain, in part, the differences in heterospecific pollen transfer.In field experiments, the results showed that seed set in six sympatric species(Gentiana macrophylla Pall. var. fetissowii, Gentianopsis paludosa (Hook. f.) Mavar. paludosa, Gentiana squarrosa Ledeb., Halenia elliptica D. Don var. elliptica,Cyananthus hookeri C. B. Cl. var. grandiflorus Marq. and Euphrasia pectinata Ten.)decreased nonlinearly as the increasing concentrations of extract of pollen from S.chamaejasme. According to the nonlinear curves, the amounts of pollens from S.chamaejasme on stigmas of G. squarrosa and of E. pectinata (i.e. 5.76 grains and3.35 grains, respectively) could reduce 63% and 55% seed set of G. squarrosa and ofE. pectinata, respectively. Thus, allelopathic effects of S. chamaejasme on G.squarrosa and E. pectinata could be realistic. The sympatric species of S.chamaejasme could avoid pollen allelopathy of S. chamaejasme to sustainthemselves. This highlights the need to study how much pollen allelopathy in S.chamaejasme influences the sympatric species through divergence in seasonal ordiurnal flowering phonologies or through asexual reproduction. If pollen allelopathyin S. chamaejasme was confirmed, it could be as a pressure of natural selection andthus play an important role in species evolution.

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

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Natl Tech Univ Ukraine, Huazhong Univ Sci & Technol, Huazhong Normal Univ, Wuhan Univ, Ternopil Natl Econ Univ, IEEE Ukraine Sect, I&M CI Joint Chapter