892 resultados para The Torrent: Toowoomba and the Lockyer Valley
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We investigated the photoluminescence (PL) of self-assembled In0.55Al0.45As/Al0.5Ga0.5As quantum dots (QDs) grown on (311)A GaAs substrate. The PL peak at 10 K shifts to lower energy by about 30 meV when the excitation power decreases by two orders of magnitude. It has a red-shift under pressure, that is the character of X-like transition. Moreover, its peak energy is smaller than the indirect gap of bulk Al0.5Ga0.5As and In0.55Al0.45As. We then attribute that peak to the type-II transition between electrons in X valley of Al0.5Ga0.5As and heavy holes in In0.55Al0.45As QDs. A new peak appears at the higher energy when temperature is increased above 70 K. It shifts to higher energy with increasing pressure, corresponding to the transition from conduction Gamma band to valence band in QDs. The measurements demonstrate that our In0.55Al0.45As/Al0.5Ga0.5As quantum dots are type-II QDs with X-like conduction-band minimum. To interpret the second X-related peak emerged under pressure, we discuss the X-valley split in QDs briefly. (C) 2000 American Institute of Physics. [S0003-6951(00)04622-2].
Photoluminescence study of multilayer In0.55Al0.45As/Al0.5Ga0.5As quantum dot at various temperature
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The photoluminescence of self-assembled multilayer In0.55Al0.45As/Al0.5Ga0.5As quantum dot (QD) was measured at various temperatures. Strong photoluminescence of wetting layer (WL) and quantum dots were observed at the same time. Furthermore, direct excitons thermal transfer process between the wetting layer and quantum dots was observed. In the study of temperature dependence of PL intensity it was found that the PL peak of wetting layer contains two quenching processes: at low temperature, excitons are thermally activated from localized states to extended two-dimensional states and then trapped by QDs; at high temperature excitons quench through the X valley of barriers. Using rate equation excitons thermal transfer and quenching processes were analyzed quantitatively.
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Room-temperature microwave (MW) oscillations are observed in GaAs/AlAs (10 nm/2 nm) doped weakly coupled superlattices (SLs) in the first plateau of the I-V curve. Oscillations induced by sequential resonant tunneling are detected in a temperature range from 15 to 300 K by applying DC bias on the Si, diodes. The temperature dependence of current at small fixed bias voltage is also measured. Through analysis, it is found that the dominant transport mechanisms are sequential resonant tunneling and phonon-assisted tunneling when the temperature is below 300 K. The low bias voltage at which oscillations are realized is helpful to restrain thermionic emission through the X valley of AlAs barriers in the room-temperature transport. (C) 1999 Published by Elsevier Science B.V. All rights reserved.
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In this paper, we propose an n-type vertical transition bound-to-continuum Ge/SiGe quantum cascade structure utilizing electronic quantum wells in the L and Gamma valleys of the Ge layers. The optical transition levels are located in the quantum wells in the L valley. The Gamma-L intervalley scattering is used to depopulate the lower level and inject the electrons into the upper level. We also show that high quality Si1-yGey pseudosubstrate is obtained by thermal annealing of Si1-xGex/Ge/Si structure. (C) 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
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The empirical pseudopotential method within the virtual crystal approximation is used to calculate the band structure of Mg1-xZnySySe1-y, which has recently been proved to be a potential semiconductor material for optoelectronic device applications in the blue spectral region. It is shown that MgZnSSe can be a direct or an indirect semiconductor depending on the alloy composition. Electron and hole effective masses are calculated for different compositions. Polynomial approximations are obtained for both the energy gap and the effective mass as functions of alloy composition at the GAMMA valley. This information will be useful for the future design of blue wavelength optoelectronic devices as well as for assessment of their properties.
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We have studied the sequential tunneling of doped weakly coupled GaAs/ALAs superlattices (SLs), whose ground state of the X valley in AlAS layers is designed to be located between the ground state (E(GAMMA1)) and the first excited state (E(GAMMA2)) of the GAMMA valley in GaAs wells. The experimental results demonstrate that the high electric field domain in these SLs is attributed to the GAMMA-X sequential tunneling instead of the usual sequential resonant tunneling between subbands in adjacent wells. Within this kind of high field domain, electrons from the ground state in the GaAs well tunnel to the ground state of the X valley in the nearest AlAs layer, then through very rapid real-space transfer relax from the X valley in the AlAs layer to the ground state of the GAMMA valley of the next GaAs well.
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Taking the inhomogenous broadening of the electron energy levels into account, a coherent model of the resonant tunneling (RT) of electrons in double quantum wells is presented. The validity of the model is confirmed with the experiments [M. Nido et al., Proc. SPIE 1268, 177 (1990)], and shows that the tunneling process can be explained by the simple coherent theory even in the presence of the carrier scattering. We have discussed the dependence of resonant tunneling on the barrier thickness L(B) by introducing the contrast ratio LAMBDA and the full width at half depth of the RT valley, and found that LAMBDA first increases with increasing barrier thickness, reaches a maximum, and then decreases with a further increase of L(B), in striking contrast to the Fabry-Perot model where a monotonic increase of the peak-to-valley ratio is predicted. We attribute the reduction of LAMBDA with large L(B) to the energy broadening resulting from the carrier scattering. A monotonic decrease of the full width at half depth of the RT valley with an increase of L(R) is also found.
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Photoluminescence of GaInP under hydrostatic pressure is investigated. The Gamma valley of disordered GaInP shifts sublinearly upwards with respect to the top of the valence band with increasing pressure and this sublinearity is caused by the nonlinear relationship between lattice constant and hydrostatic pressure. The Gamma valleys of ordered GaInP rise more slowly than that of the disordered one and the relationship between the band gap and the pressure can not be explained in the same way. Taking into account the interactions between the Gamma valley and the folded L valleys, as well as, the X valleys, the experimental pressure dependences of the band gap of ordered GaInP epilayers are calculated and fitted quite well using first order perturbation theory. The results indicate that simultaneous ordering along [111] and [100] directions can occur in ordered GaInP. (C) 1996 American Institute of Physics.
Resumo:
The photoluminescence (PL) of In0.55Al0.45As/Al0.5Ga0.5As self-assembled quantum dots has been measured at 15 and 80 K under hydrostatic pressure. The lateral size of the dots ranges from 7 to 62 nm. The emissions from the dots with 26, 52 and 62 nm size have a blue shift under pressure, indicating that these quantum dots have the normal type-I structure with lowest conduction band at the Gamma -valley. However, the PL peak of dots with 7 nm diameter moves to lower energy with increasing pressure. It is a typical character for the X-related transition. Then these small dots have a type-II structure with the X-valley as the lowest conduction level. An envelope-function calculation confirms that the Gamma -like exciton transition energy will rise above the X-like transition energy in the In0.55Al0.45As/Al0.5Ga0.5As structure if the dot size is small enough.
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近年来,随着对作物重茬障碍原因的深入研究,植物的化感作用越来越受到国内外众多学者的重视。花椒(Zanthoxy piperitum.)为芸香科植物,是一种收益早、用途广、价值高的经济树种,是川西干旱河谷地区的重要经济作物,其连作障碍也倍受关注,系统研究花椒化感作用将有助于理解和最终解决花椒连作障碍问题。本文首先通过萃取、层析等方法分离花椒主效化感成分;通过外加不同浓度的花椒叶水浸液研究了对土壤氮素养分循环的影响;研究了花椒叶水浸液对苜蓿生理生化、光合作用、氮素养分吸收的影响,并对外施氮肥对这种化感影响的缓解作用做了研究;研究了花椒化感潜力对全球变化——UV-B增强辐射的响应。主要研究结果如下: 1.用不同极性的有机溶剂对花椒叶水浸液浓缩浸膏萃取、柱层析,结合生物活性检测,分离得到主效化感作用组分的一种化感物质——对甲氧基苯酚。采用该物质纯品进行生物活性检测,证明其具有化感作用。 2.花椒叶水浸液处理土壤30天后,土壤硝态氮、铵态氮、无机氮(硝态氮+铵态氮)与对照相比,随着花椒叶水浸液浓度的增加呈现降低的趋势,其中土壤铵态氮含量显著降低,而硝态氮含量的变化则不显著,无机氮含量也显著降低。土壤脲酶和蛋白酶的活性与无机氮含量的变化趋势相同。随着花椒叶水浸液浓度的增加,氨化细菌数量显著降低,固氮菌的数量变化不显著,硝化细菌和反硝化细菌数量有减少的趋势。60天后,硝态氮含量、铵态氮含量、无机氮随水浸液浓度增加的变化趋势与30天时相似;随着花椒叶水浸液浓度的增加,氨化细菌、固氮菌的数量显著减少,硝化细菌数量、反硝化细菌数量仍呈减少趋势;土壤脲酶、蛋白酶活性与第30天的变化趋势相同。第60天与第30天的结果相比,相同水浸液浓度处理的硝态氮、铵态氮、无机氮均有下降的趋势,但除了25g.L-1水浸液处理的外,其它相同浓度的处理间差异均不显著;除了12.5 g.L的处理外土壤脲酶活性均呈增强的趋势;蛋白酶活性都有不同程度的增加;花椒叶水浸液处理的土壤硝化细菌和反硝化细菌数量呈增加趋势。 3.随着花椒叶水浸液浓度的增加,显著抑制了苜蓿根长、地上地下生物量、叶绿素含量、叶片中可溶性蛋白的含量,净光合速率。苜蓿体内四种抗氧化酶(POD、SOD、CAT、APX) 活性随着水浸液浓度的增加而降低,而丙二醛含量则增加。苜蓿氮初级同化相关酶硝酸还原酶(NR)、谷氨酰合成酶(GS)、谷氨酸脱氢酶(GDH)的活性随着水浸液浓度的增加受到不同程度的影响。总的来说,苜蓿硝酸还原酶、谷氨酰合成酶的活性受到抑制,而谷氨酸脱氢酶活性的变化则比较复杂,根呈先降低后增加的趋势,叶片则无显著变化。外施两种不同浓度的硝酸铵氮肥后,对12.5、25 g.L-1花椒叶水浸液处理的苜蓿化感作用有显著的缓解作用,表现在株高、生物量、光合作用等方面,大多达到与对照(0 g.L-1)未施氮肥无显著差异的水平,而对50 g.L-1水浸液处理的苜蓿幼苗,虽有一定的缓解作用,但这种作用均未达到与对照(0 g.L-1)未施氮肥时无显著差异的水平。 4. UV-B增强辐射处理花椒后,花椒的化感潜力显著增强。花椒叶片内UV-B吸收物质的含量和总酚含量均显著增加。 In recent years, with profound research on the reasons of continuous cropping obstacles, allelopathy received increasing attention to many scholars at home and abroad. Zanthoxy bungeanum as a Rutaceae plant is a high economic value species which gains early and uses widely. Zanthoxylum is an important economic crop in the arid valley of western Sichuan region, and its not even has received much concern for the continuous cropping obstacles. The systematic study of allelopathy of Zanthoxylum will contribute to the understanding and final settlement of this issue. The major allelopathic composition was separated through the extraction, chromatography combined with other methods. The impact on soil nutrient cycling was also studied through the addition of different concentrations of water extracts of Zanthoxylum. Furthermore, the effects of water extracts of Zanthoxylum leaves on alfalfa leaf physiological and biochemical indexes, photosynthesis, soil enzymes and nutrient uptake of nitrogen and the mitigation of allelopathy through using external fertilizer were studied to put forward scientific resolvent for Zanthoxylum continuous cropping obstacles .The response of allelopathic potential of Zanthoxylum to global change - UV-B enhanced radiation was studied . The main findings are as follows: 1. Through extraction with different polar organic solvents on concentrated water extract of Zanthoxylum leaf and then using column chromatography combined with detection of biological activity, one of the main allelopathic components- methoxy-phenol was isolated. The biological activity testing of the pure material of methoxy-phenol proved that it does have allelopathic potential. 2. Thirty days after treating soil with water extract of Zanthoxylum leaf, as compared with the control, the contents of soil nitrate, ammonium, nitrate plus ammonium nitrogen showed a trend of decrease with the increase of the concentration of water extract whereas the content of ammonium nitrogen showed a significant reduction, and the content of nitrate did not change significantly, the content of nitrate plus ammonium nitrogen also showed a significant (P <0.05) redction. The activity of soil urease and protease showed the same trend as the content of nitrate nitrogen plus ammonium nitrogen. With the increase in the concentration of water extract, the number of ammonification bacteria significantly reduced but nitrogen-fixing bacteria did not change significantly and there was a decreasing trend in the number of nitrifying bacteria and denitrifying bacteria. Sixty days after the treatment, with the increase in solution concentration of water extract of Zanthoxylum leaf, the content of nitrate、 ammonium nitrogen, nitrate plus ammonium nitrogen showed a similar change trend to 30 days’; the number of ammonification bacteria, nitrogen-fixing bacteria significantly reduced ; the number of nitrifying bacteria, denitrifying bacteria was still an downward trend; the activity of soil urease and protease showed the same trend as the 30th days’. Compared to the results of the 30th days’, the content of nitrate, ammonium, nitrate plus ammonium nitrogen showed a decrease trend between the treatment of same concentration, but there was no significant difference except the treatment of 25g.L-1 between the same concentration; the activity of soil urease showed enhanced trend except the treatment of 12.5 g.L-1; the activity of protease increased to varying degrees; the number of ammonification bacteria、 nitrifying bacteria and denitrifying bacteria were growing while nitrogen-fixing bacteria reduced.. 3. With the increase of the concentration of water extract of Zanthoxylum leaf, the water extract significantly inhibited the root length, aboveground biomass, content of chlorophyll and soluble protein in leaf and net photosynthetic rate. The activity of four antioxidant enzymes (POD, SOD, CAT, APX) reduced with the increase in concentration of the water extract but the content of MDA increased. The activity of enzymes related to primary nitrogen assimilation such nitrate reductase (NR), glutamyl synthetase (GS), glutamate dehydrogenase (GDH) were subject to different degrees with an increase in the concentration of water extracts. In general, the activity of nitrate reductase, glutamyl synthetase were inhibited, while change in the activity of glutamate dehydrogenase was more complex. The activity of glutamate dehydrogenase in leaf was first reduced and then increase,but did not change significantly in root. After using two external different concentrations of nitrogen fertilizer, there was a significant mitigation in inhibiton in plant height, biomass, photosynthesis, etc. in the treatment of 12.5,25 gL-1 of water extract of Zanthoxylum leaf, and most of these indexes showed no significant difference with the control (0 g.L-1, no external fertilizer was added) .Although there showed a certain degree of ease in the treatment of 50 g.L-1 , there was still a significant difference compared with the control (0 gL-1) in which no external fertilizer was used. 4.The allelopathic potential of Zanthoxylum positively responded to enhanced UV-B significantly. The content of UV-B absorbing compounds and the total phenol also significant increased.
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青藏高原东部分布着世界最高的林线,该区域也是由欧亚北温带物种形成的林线的南界。在大面积野外踏勘的基础上,选择青藏高原东部具有典型高山林线分布的三个地点(滇西北白马雪山、川西北鹧鸪山及岷江源地区)作为研究区,从种群的结构、生存特征、分布格局及分形特征等方面对青藏高原东缘高山林线乔木种群生态学特征进行了研究,并在此基础上探讨了人类活动对林线种群生态特征及林线格局的影响。结果表明,林线区乔木树种多以单种群形式存在,林线区群落结构简单,乔木层多为单一树种组成,其生长型较之郁闭林发生了急剧的变化:树木高度急剧下降,而发展多茎多分枝的生长型。生长型的转变是高山林线乔木对恶劣自然条件的形态适应。 研究发现,在青藏高原东缘,阴坡林线乔木主要是冷杉(Abies spp.),阳坡主要由圆柏(Sabina spp.)组成,少数地方还有云杉(Picea spp.)。阴坡乔木种群结构多表现为增长型,幼苗和幼树在种群中占较大比重,种群潜在自然更新能力较强,但幼龄个体死亡率非常高,存活曲线多接近Deevey-Ⅲ型;阳坡乔木种群幼苗个体数极少,幼树相对增加。野外调查表明,人为活动较频繁的阳坡林线区幼苗数量极少甚至缺失,而受人为活动干扰较小的样地中幼苗和幼树数量明显增多,从一个侧面说明放牧等人类活动可能对林线种群的更新带来较大影响,而对卡卡沟围栏内外的样地分析也进一步证明了这一结果。 所研究林线乔木种群各龄级的空间格局在不同尺度上表现为聚集、随机和均匀分布,以聚集分布为主;各龄级在不同尺度上表现出显著的相关性,幼苗通常与另外两个龄级的关联性较密切。各龄级间显著的相关性表明不同龄级个体在空间交错分布,有利于对各种资源的充分利用,对种群的生存和发展非常有利,反映了高山生态系统恶劣生境中种群的一种适应对策。 林线乔木种群各龄级分布格局的计盒维数有差别,林线种群的计盒维数总是小于郁闭林种群的计盒维数。另外,郁闭林各龄级计盒维数通常也高于林线各龄级,表明不同海拔或者不同群落类型中的乔木树种具有不同的水平空间占据能力。林线区种群分布格局的计盒维数都很低,占据现实水平空间的程度较低,具有相对较高的生态间隙维,其潜在占据空间的能力较高,群落还可提供给种群的最大空间限度较大,但实际上由于受群落中种内、种间的竞争及林线区恶劣的生态环境条件的限制,其潜在空间占据能力可能难以表现出来。 青藏高原东缘高海拔地带以季节性游牧为主要的资源利用和生产方式,阳坡森林郁闭度低于阴坡,灌丛数量和种类较阴坡少,融雪早且积雪时间短,所以阳坡包括高山林线区成为当地牧民游牧路线的必经之地。牲畜的践踏、啃食使得幼龄乔木树种个体数量大大减少,严重阻碍了林线乔木种群的自然更新,同时种群占据空间的能力也明显降低。因此可以认为,在青藏高原东部地区,山地游牧等人为干扰叠加于恶劣的自然条件,阳坡林线的自然更新潜力受到抑制,其生存状态较之阴坡林线显著恶化,并可使阳坡林线高度逐渐降低。高山林线区森林一旦破坏在短时间内很难有效更新和恢复,因此,对于处于恶劣高山生境中的乔木种群应加强保护,同时适度控制人为干扰强度和幅度以减少其直接和间接破坏,防止阳坡林线退化并促进高山生态系统的自然恢复。 Eastern Qinghai-Tibetan Plateau has the highest timberline of the world. On the basis of field surveys and literature reviews, three typical alpine timberlines were chosen for in-depth studies, i.e., Baima Snow Mountain in northwest Yunnan, Zhegu Mountain and the waterhead area of Minjiang River in west Sichuan. Using the methodologis of population ecology, we analyzed the population structure, survival characteristics, spatial point patterns and fractal dimensions of the timberline tree populations and discussed the impacts of grazing on the structure and spatial pattern of alpine timberline. Compared with closed forests, the community structure of timberline is simpler, usually with one or two species constituting the tree layer. Differences also exist in the growth forms: the trees were significantly shorter with more stems and branches, reflecting morphological adaptation of trees to the severe conditions at timberline. In the eastern Qinghai-Tibetan Plateau, Abies spp. often formed alpine timberline in the north-facing slope while Sabina spp. and sometimes Picea spp. in the south- facing slope. The population structures of north-facing slope showed an increasing trend, with numerous seedlings and saplings. However, the survival curves tend to follow Deevy-III because of high dead ratio of young individuals. There are only few seedlings in the south-facing slope with heavy grazing, demonstrating that human disturbance may prevent regeneration at alpine timberline, which was confirmed by comparisons between fenced enclosures and control plots in the Kaka Valley. Depending on the spatial scales on consideration, the individuals of different age-classes showed clumping, random or even distribution, but mostly with clumping distribution. At all scales, individuals in different age-classes were all significantly correlated with each other while the seedlings were usually more correlated to two other age classes. This high degree of correlation among different age classes indicates that individuals of different age classes are spatially interlocked with each other, which helps sufficient utilization of various resources and is conducive to the survival and development of population. It is another adaptation strategy for trees at the severe environment. The spatial patterns of different age classes had different box dimension. In general, the box dimensions of total individuals and each age class at timberline are always smaller than that of closed forests, suggesting that space occupation capacity is not the same for populations at different altitude or in different communities. Populations on both the south- and the north-facing slopes had a very low box dimensions (far away from the max., 2), however, the lower the box dimension, the bigger the potential space provided by community. In fact, because of inner- and inter- competition as well as the severe conditions at timberline, this kind of potential ability can hardly be realized. Mountain pastoralism is the major type of as well as the only most effective way of resource uses in the high elevation regions of the eastern Qinghai-Tibetan Plateau. Due to lower canopy cover, less bushes and short snow-cover time, south-facing slopes became the favorite pastures. Damages from livestock through tramping, browsing and others have greatly reduced the number of young individuals. As a result, the potential of timberline trees to regenerate and their ability to occupy more space are greatly inhitibted. We conclude that human disturbances (mountain pastoralism) as well as harsh environmental conditions co-worked to inhibit the regeneration of tree populations in the south-facing slope and made south slopes more difficult than the north-facing slopes for trees to survive and develop, resulting a gradual retreat of timberline in the north-facing slopes. Forests at alpine timberline are susceptible to disturbance and difficult to regenerate and restore once damaged and controlling human disturbances is important for protecting the forest ecosystems at the timberline area.
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干旱胁迫是全球范围内影响植物生存、生长和分布的重要环境因子。岷江上游干旱河谷区,由于生态环境的脆弱性和长期人类活动的干扰和过度利用,导致植被严重退化,水土流失加剧,山地灾害频繁,干旱化和荒漠化趋势明显。这种趋势若不能遏制,将严重阻碍区域社会经济的快速协调发展,并且威胁成都平原地区的发展和长江中下游地区的生态安全。因而开展干旱河谷生态恢复研究成为解决这些问题的关键。水分匮乏是限制干旱河谷生态恢复的关键因子,在全球气候变化的背景下,干旱胁迫在区域尺度上可能会更加严重,并使干旱河谷的生态环境更加恶化。因此,深入研究干旱河谷乡土植物对干旱胁迫的响应和适应机理,具有非常重要的理论和实践意义。 本论文以岷江上游干旱河谷的三种乡土豆科灌木,白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha)理论和实践意义。为研究对象,在人工控制条件下设计了4-5个连续性干旱胁迫处理,系统地研究了灌木幼苗的生长、生物量积累和水分利用效率(WUE)、形态结构和生理过程等对干旱胁迫的反应,揭示了幼苗的干旱适应能力及种间差异。主要研究结论如下: 1) 灌木生长和繁殖对干旱胁迫的反应 在干旱胁迫下,幼苗生长速率显著减小,叶片衰老和脱落比率增大,这些变化随着胁迫强度的增加具有累积效应。叶片比茎对干旱胁迫的反应更敏感。在严重干旱胁迫下,幼苗的有性繁殖被限制,但在中等程度干旱胁迫下,幼苗的有性繁殖能力被提高。 2) 灌木生物量积累及其分配和WUE对干旱胁迫的反应 在干旱胁迫下,灌木各器官的生物量都显著减小,但是生物量的分配侧重于地下部分,使得根茎比在干旱条件下增大。幼苗的耗水量(WU)随着干旱胁迫的增加而显著减少。白刺花和小马鞍羊蹄甲WUE在干旱胁迫下降低;小雀花的WUE在中等干旱胁迫下升高。 3) 灌木叶片结构特征对干旱胁迫反应 白刺花叶片具有较为典型的旱生型结构,而小马鞍羊蹄甲和小雀花则为中生型结构。在1至2年的干旱胁迫下,灌木叶片结构组成未发生本质性的改变,主要是细胞大小的变化。在中等和严重干旱胁迫下,叶肉组织厚度明显减小;并且气孔和表皮细胞面积也显著减小,气孔和表皮细胞密度显著增大;叶肉细胞层数、P/S值、表皮厚度等无显著变化。 4) 灌木对干旱胁迫的生理响应 气体交换参数和叶片相对含水量(RWC)在中等干旱胁迫下发生了明显的改变,而叶绿素荧光参数和光合色素含量在严重干旱胁迫下才发生显著变化。在干旱胁迫下,净光合作用速率(Pn)、气孔导度(gs)和RWC呈下降趋势,而叶片温度(Tl)呈增加趋势,蒸腾速率(Tr)的变化不明显。除了日最大Pn减小以外,干旱胁迫对气体交换参数的日变化无显著影响,但是对光合-光响应曲线有显著的影响,使有效光合时间缩短。在严重干旱胁迫下光系统受到损害而代谢减弱,PSⅡ中心的内禀光能转换效率(Fv/Fm)、量子产量(Yield)、光化学淬灭参数(qP)显著降低,而非光化学淬灭参数(NPQ)明显增加。气孔限制和非气孔限制对Pn的影响与干旱胁迫强度有关。在中度胁迫下,气孔限制起主导作用,在严重胁迫下非气孔限制起主导作用,40% FC水分条件可能是灌木由气孔限制向非气孔限制的转折点。 5) 灌木对干旱胁迫的适应能力及其种间差异 三种灌木对干旱胁迫具有较好的适应能力,即使在20% FC,幼苗未因干旱胁迫III而死亡;80% FC适宜于幼苗生长。白刺花生长速率慢,耗水量较少,具有较强的耐旱和耐贫瘠能力,并具有干旱忍受机制,能够在较干旱的环境中定居和生长。小马鞍羊蹄甲和小雀花,生长快,水分消耗量较大,尤其是小雀花,对干旱胁迫的忍受能力较弱,具有干旱回避机制,因而适宜于在较为湿润的生境中生长。综合分析表明,生长速率较慢的物种抗旱能力较强,其更适宜于作为干旱地区植被恢复物种。 Drought is often a key factor limiting plant establishment, growth and distribution inmany regions of the world. The harsh environmental conditions and long-termanthropogenic disturbance had resulted in habitat destruction in the dry valley ofMinjiang river, southwest China. Recently, it tended to be more severe on the vegetationdegradation, soil erosion and water loss, natural disaster, as well as desertification, whichimpact on regional booming economy and harmonious development, and would be verydangerous to the environmental security in the middle and lower reaches of Yangzi River.Therefore, ecological restoration in the dry valley is one of the vital tasks in China. Waterdeficit is known to affect adversely vegetation restoration in this place. Moreover, in thecontext of climate change, an increased frequency of drought stress might occur at aregional scale in the dry valleys of Minjiang River. The selection of appropriate plantingspecies for vegetation restoration in regard to regional conditions is an important issue atpresent and in further. The research on responses of indigenous species to drought stresscould provide insights into the improvement of the vegetation restoration in the dry valleys of Minjiang River. In this paper, the responses of three indigenous leguminous shrubs, Sophora davidii,Bauhinia faberi var. microphylla and Campylotropics polyantha, to various soil watersupplies were studied in order to assess drought tolerance of seedlings, and to compare interspecific differences in seedlings’ responses to drought stress. The results were as follows: 1 Growth and reproduction of shrubs in response to drought stress Seedling growth reduced significantly while leaf senescence accelerated underdrought stress, the cumulative responses to prolonged drought were found. The capacityfor reproduction was limited by severe drought stress, and improved by moderate droughtstress. Leaf responses were more sensitive than shoot to various water supplies. 2 WUE, biomass production and its partitioning of shrubs in response to drought stress Drought stress reduced significantly the total dry mass and their components ofseedlings, and altered more biomass allocation to root system, showing higher R/S ratiounder drought. Water use (WU) and water-use efficiency (WUE) of both S. davidii and B.faberi var. microphylla declined strongly with drought stress. The WU C. polyantha ofalso declined with drought stress, but WUE improved under moderate drought stress. 3 Anatomical characteristics and ultrastructures of leaves in response to drought stress There were xeromorphic for S. davidii leaves and mesomorphic for B. faberi var.microphylla and C. polyantha at the all water supplies. The foundational changes in leafstructures were not found with drought stress. However, mesophyll thickness, the areas ofstomatal and epidermis reduced slightly while the densities of stomatal and epidermisincreased under severe drought stress. Variations in these parameters could mainly be duoto cell size. Other structures did not displayed significant changes with drought stress. 4 Physiological responses of shrubs to drought stress The gas exchange parameters and leaf relative water content (RWC) were affectedby moderate stress, while chlorophyll fluorescence and chlorophyll content were onlyaffected by severe stress. Drought stress decreased net photosynthesis rate (Pn), stomatalconductance, light-use efficiency and RWC, and increased leaf temperature. Therespiration rates (Tr) were kept within a narrower range than Pn, resulting in aprogressively increased instantaneous water use effiecency (WUEi) under drought stress.Moreover, drought stress also affected the response curve of Pn to RAR, there was adepression light saturation point (Lsat) and maximum Pn (Pnmax) for moderate andsevere stressed seedling. However, diurnal changes of gas exchange parameters did notdiffer among water supplies although maximum daily Pn declined under severe stress.VISevere stress reduced Fv/Fm, Yield and qP while increased NPQ and chlorophyll content.Photosynthetic activity decreased during drought stress period due to stomatal andnon-stomatal limitations. The relative contribution of these limitations was associatedwith the severity of stress. The limitation to Pn was caused mainly by stomatal limitationunder moderate drought stress, and by the predominance of non-stomatal limitation undersevere stress. In this case, 40% FC water supply may be a non-stomatal limitation 5 Interspecific differences in drought tolerance of shrubs Three shrubs exhibited good performance throughout the experiment process, evenif at 20% FC treatment there were no any seedlings died, 80% FC water supply wassuitable for their establishment and growth. S. davidii minimized their water loss byreducing total leaf area and growth rate, as well as maintained higher RWC and Pncompared to the other two species under drought stress, thus they might be more tolerantto the drought stress than the other two species. On the contrary, it was found that C.polyantha and B. faberi var. microphylla had higher water loss because of their stomatalconductance and higher leaf area ratios. They reduced water loss with shedding theirleaves and changing leaf orientation under drought stress. Based on their responses, thestudied species could be categorized into two: (1) S. davidii with a tolerance mechanismin response to drought stress; (2) C. polyantha and B. faberi var. microphylla withdrought avoidance mechanism. These results indicated that slow-growing shrub speciesare better adapted to drought stress than intermediate or fast-growing species in present orpredicted drought conditions. Therefore, selecting rapid-growing species might leavethese seedlings relatively at a risk of extreme drought.
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岷江上游干旱河谷区水土流失强烈,地质灾害频繁,生态环境十分脆弱,而土壤条件恶劣(水分不足和养分缺乏)是阻碍该区植被恢复的关键因子,因此研究水分和乡土灌木生长对土壤的影响对该区的生态恢复具有指导意义。本文通过定点模拟实验,选取三种优势豆科灌木为研究对象,分别是白刺花(Sophora davidii)、小马鞍羊蹄甲(Bauhinia faberi var. microphylla)和小雀花(Campylotropics polyantha),设置5 个水分梯度,分别为100%、80%、60%、40%和20%田间持水量(FC),对栽种植物与不种植物下土壤理化性质和酶活性进行测定分析,系统比较和研究了不同水分条件和不同乡土灌木生长对干旱河谷区土壤结构、养分循环、酶活性以及微生物量的影响。主要结果如下:1. 无论生长植物与否,土壤的毛管持水量和毛管孔隙度都随着水分含量的减少而降低,最大持水量、总孔隙度和容重变化不大,相应地,土壤中的非毛管孔隙随含水量的减少而升高。各水分条件下,种植植物的毛管持水量和毛管孔隙度低于无植物生长的土壤,非毛管孔隙度相应地高于无植物土壤。土壤含水量在100%-40% FC 时,三种豆科灌木的毛管持水量和毛管孔隙度存在差异,而20% FC 条件下,三种豆科灌木土壤的物理性质基本相同。2. 水分胁迫影响土壤中养分的矿化和积累,主要表现在降低了水溶性碳和铵态氮的含量,中等程度胁迫时(60% FC)促进了有机碳和硝态氮的富集,对速效钾和有效磷没有明显作用。种植豆科灌木后各水分梯度上都增加了有机碳、铵态氮、速效钾和有效磷的积累。增加程度上三种豆科灌木间有一定差异,对于土壤有机碳总量,种植白刺花和小马鞍羊蹄甲明显高于小雀花,同样的情况还出现在铵态氮和速效钾上,但是对于有效磷,种植小雀花后的增加程度则明显高于白刺花和小马鞍羊蹄甲。种植豆科灌木不仅增加了土壤养分的相对含量,也改变了其在水分梯度上的变化趋势及其变化幅度,这种作用主要体现在碳元素和氮元素上。3. 无植物生长时脲酶活性随水分含量的减少而升高,水分胁迫对磷酸酶和过氧化氢酶的作用不显著,蔗糖酶也保持在相对较高的水平。种植植物后,蔗糖酶、磷酸酶活性与无植物时相比有较大幅度的提高,种植白刺花的脲酶活性也升高,其升高的程度在不同水分含量时不同。种植植物还降低了酶活性在水分梯度上的变幅,使之在水分梯度间的差异显著性降低。脲酶活性在指示土壤性质改变方面是较敏感的指标,其它三种酶在不同植物间的差异不明显。4. 在无植物生长时,中等程度的水分胁迫(60% FC)提高了土壤微生物量碳含量,过高或过低的土壤水分均不利于微生物碳的积累。种植小马鞍羊蹄甲后微生物量碳在水分梯度上的变化趋势与无植物生长时一致,而种植白刺花和小雀花后微生物量碳随着水分含量的减少而降低。不同种类植物的微生物量碳在水分梯度上的变化特征也不同,100% FC 条件下三种植物间没有差异,80%和60% FC 条件下小马鞍羊蹄甲显著高于白刺花和小雀花,40%和20% FC 条件下白刺花和小马鞍羊蹄甲也显著高于小雀花,说明不同种类植物随着干旱胁迫程度的加深微生物量碳的降低幅度不同,在极度干旱时,白刺花和小马鞍羊蹄甲土壤依然保持了较高的微生物活性,而小雀花土壤微生物量则明显下降。The dry valley of the upper reaches of the Minjiang River is seriously degradedmountain ecosystem. It was endangered by extremely soil lost and frequentlygeological disaster. Previous studies showed that short of water and nutrients in soilwas the principal limiting factors of vegetation restoration in this area. The typical soiland three dominant leguminous shrubs Sophora davidii, Bauhinia faberi var.microphylla and Campylotropics polyantha in upper reaches of arid Minjiang Rivervalley were considered as experimental material. Two-month old seedlings of eachspecies were exposed to five water supplies (100%, 80%, 60%, 40% and 20% waterfield capacity (FC)) in a temperature and light-controlled greenhouse. Afterthree-month water treatment, soil physiochemical variables and soil microbialactivities were determined by conventional methods. The main results showed that:1. Soil capillary capacity and capillary porosity decreased along water supplyregimes in all treatments, while saturated water capacity, total porosity and bulkdensity kept in a relatively stable level, as a result, the non-capillary porosity andcapacity increased with decrease of water supply. Compared to non-planted soil, theplant-soil systems had a higher non-capillary porosity and capacity, suggestingappropriate oxygen was present in soil to maintain the living of microorganism. Soilof three type shrub species shared the same capillary capacity and capillary porosityunder 20% FC.2. Water soluble carbon and NH4+-N decreased in response to water stress, whiletotal organic carbon and NO3--N promoted by moderate water stress and inhibited by 100% and 20% FC. Total organic carbon, NH4+-N, rapidly available K and availableP increased after the planting of leguminous shrubs in five water supply regimescompared to non-planted soil. For TOC, NH4+-N and rapidly available K, thepromotion effect was higher in S. davidii and B. faberi var. microphylla than C.polyantha planted soil, while available P displayed the opposite side. The planting ofshrubs also reduced the variance of observed traits along water supply gradients.3. Drought stress increased urease activity in non-planted soil, while insignificantdifferences were observed in phosphatase and catalase activity among five watersupply regimes. The planting of leguminous shrubs facilitated the β-glucosidase andphosphatase activity compared to the non-planted soil. It also reduced the variance ofenzyme activity along water supply gradients. Urease was more sensitive to waterstress than other three enzymes.4. Soil water content significantly affected microbial biomass carbon andCmic:Corg. S. davidii and B. faberi var. microphylla showed more drought toleranceability than C. polyantha, attributing not only to their relatively smaller variance ofmicrobial biomass carbon along soil water supply gradients, but also to the highlevel of microbial activity under severe water stress. S. davidii and B. faberi var.microphylla benefited reproduction of soil microorganism at 60%-80% FC, whilesevere drought limited it due to the competition of water and nutrients between plantand soil microorganism.
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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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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.