341 resultados para Transpiration
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在陕北黄土丘陵沟壑区用LI-6400光合仪对中国沙棘、俄罗斯沙棘和俄罗斯沙棘×中国沙棘光合特性及影响因子进行了测定。测定结果表明三者的光合速率、气孔导度、胞间CO2浓度、蒸腾速率日变化均为"双峰"曲线。中国沙棘光合"午休"现象较轻,日光合速率和午后光合速率、气孔导度、胞间CO2浓度、蒸腾速率极显著(p<0.01)高于俄罗斯沙棘和俄罗斯沙棘×中国沙棘。中国沙棘光合作用最适气孔导度、大气CO2浓度、空气相对湿度比俄罗斯沙棘小,最适胞间CO2浓度、蒸腾速率、气温、光合有效辐射比俄罗斯沙棘高。中国沙棘在最适气孔导度、胞间CO2浓度、蒸腾速率和大气CO2浓度下的光合速率比俄罗斯沙棘高;在最适气温、空气相对湿度、光合有效辐射下的光合速率比俄罗斯沙棘低。俄罗斯沙棘×中国沙棘光合"午休"现象比中国沙棘强,比俄罗斯沙棘弱,午后光合速率与俄罗斯沙棘相近;俄罗斯沙棘×中国沙棘最适气孔导度、大气CO2浓度高于中国沙棘和俄罗斯沙棘,最适胞间CO2浓度低于中国沙棘和俄罗斯沙棘,最适蒸腾速率、气温、空气相对湿度、光合有效辐射居于中国沙棘和俄罗斯沙棘之间。俄罗斯沙棘×中国沙棘在最适气孔导度下的光合速率高于中国沙棘和俄罗斯沙棘;在最适胞间CO...
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由于人类活动所引起的地球大气层中温室气体的富集已导致全球地表平均温度在20世纪升高了0.6 ℃,并预测在本世纪将上升1.4-5.8 ℃。气候变暖对陆地植物和生态系统影响深远,并已成为全球变化研究的重要议题。高海拔、高纬度地带的生态系统对气候变化最敏感。而在高原和高山极端环境影响下所形成的高寒草甸生态系统极其脆弱,对由于温室效应引起的全球气候变化极其敏感,对这些变化的响应更具有超前性。 本研究以川西北高寒草甸植物群落及几种主要物种为研究对象,采用国际山地综合研究中心(ITEX)普遍所采用的增温方法-----开顶式生长室(OTC)模拟气候变暖来研究增温对高寒草甸植物群落结构、物质分配及其主要物种生长和生理的影响,以探讨高寒草甸植物响应与适应气候变暖的生物学和生态学机制。主要研究结论如下: 1、OTC的增温效果 由于地温、地表温度和气温的平均值在OTC内分别高出对照样地0.28℃、0.46℃和1.4℃,这说明本研究所采用的开顶式生长室(OTC)起到了增温的作用;同时,由于温室内与温室外接受的降水量相同,温室内由于热量条件的改善,土壤蒸发和植被的蒸腾作用增强,直接导致了OTC内土壤表层相对湿度的减少。 2、群落结构对增温的响应 由于增温时间较短,增温内外样地的物种组成并未发生改变;但增温后一定程度上改变了植物群落的小气候环境,从而导致物种间的竞争关系被破坏,种间竞争关系的破坏引起群落优势种组成发生相应的改变,在对照样地,鹅绒委陵菜、甘青老鹳草、遏蓝菜和蚤缀是占绝对优势的物种,而在OTC内,小米草、尼泊尔酸模、垂穗披碱草、发草和羊茅的重要性显著增加。 禾草和杂草由于对增温的生物学特性及其资源利用响应的不同,加之增温造成土壤含水量下降等环境因子的改变。与对照样地相比较,OTC内禾草的盖度及生物量都显著增加,而杂草的盖度和生物量则显著下降。 3、植物生长期对增温的响应 OTC内立枯和调落物的生物量在生长季末(10月份)都要小于对照样地的立枯和调落物生物量,而OTC内的地上鲜体生物量在10月份却略高于对照样地。这说明OTC内植物的衰老或死亡得以延缓,而植物的生长期得以延长。 4、群落生物量及分配对增温的响应 OTC内的地上鲜体生物量(10月份除外)和地下0-30cm的根系生物量与对照样地相比较,都出现了不同程度的减少;土壤根系的分配格局也发生了明显的改变,其中,OTC内0-10cm土层的生物量分配比例增加,而20-30cm土层生物量分配比例的减少。 5、群落碳、氮对增温的响应 增温后,OTC内植物群落地上活体和地下活根的碳浓度不同程度的高于对照样地,植物群落的碳库在OTC内也略高于对照样地;而OTC内植物群落地上活体和地下活根的氮浓度不同程度的低于对照样地,其植物群落的氮库与对照样地相比也略有下降。 6、几种主要植物的生长及物质分配对增温的响应 垂穗披碱草在增温后株高、比叶面积和地上生物量均显著地增加;尼泊尔酸模在增温后比叶面积和单株平均生物量积累显著地增加,而各组分中,增温处理使叶的生物量显著增加,而根的生物量却显著下降;鹅绒委陵菜在增温后株高、比叶面积和单株平均生物量积累显著地减少,而各组分中,增温处理使叶和茎的生物量显著减少,根的生物量却显著地增加。 尼泊尔酸模的LMR、RMR、R/S、根部碳含量、碳和氮在叶片与根部的分配比例在增温后显著地增加,而SMR、根部氮含量、碳和氮在茎部的分配比例在增温后却显著地降低;鹅绒委陵菜的RMR、R/S、碳和氮在根部的分配比例在增温后显著地增加,而SMR、LMR、碳在叶片的分配比例在增温后却显著地降低 7、几种主要植物的光合生理过程对增温的响应 增温使垂穗披碱草和尼泊尔酸模叶片中的叶绿素a、叶绿素b、总叶绿素含量显著增加;而鹅绒委陵菜叶片的叶绿素a、叶绿素b、总叶绿素含量在增温后显著减少,类胡萝卜素含量在增温后却显著增加。 增温对3种植物的气体交换产生了显著影响。其中,垂穗披碱草和尼泊尔酸模叶片的光响应曲线在增温后明显高于对照处理,A、E、gs、Pmax、、Rday、AQY和LSP显著增加,而LCP则显著降低;鹅绒委陵菜的光响应曲线在增温后则明显的低于对照处理,A、E、gs、Pmax、、Rday、AQY和LSP显著减少,而LCP则显著增加。 增温后垂穗披碱草和尼泊尔酸模叶片的Fv/Fm、Yield和qP显著增加;而鹅绒委陵菜叶片的Fv/Fm、Yield和qP则显著减少,qN却显著地增加。 8、几种主要植物的抗氧化酶系统对增温的响应 增温使垂穗披碱草和尼泊尔酸模体内抗氧化酶活性和非酶促作用有所提高,植物膜脂过氧化作用降低;鹅绒委陵菜叶片中酶促反应和非酶促反应在增温后也显著提高,但可能由于增温后的土壤干旱超过了鹅绒委陵菜叶的抗氧化保护能力,抗氧化酶活性及非酶促反应(脯氨酸、类胡萝卜素)的提高不足以完全清除干旱诱导形成的过量活性氧,因此叶片的膜脂过氧化程度仍然显著提高。 Enrichment of atmospheric greenhouse gases resulted from human activities such as fossil fuel burning and deforestation has increased global mean temperature by 0.6 ℃ in the 20th century and is predicted to increase in this century by 1.4-5.8 ℃. The global warming will have profound, long-term impacts on terrestrial plants and ecosystems. The ecoologcial consequences arising from global warming have also become the very important issuses of global change research. The terrestrial habitats of high-elevation and high-latitude ecosystems are regarded as the most sensitive to changing climate. The alpine meadow ecosystme, which resulted from the composite effects of mountain extreme climatic factors in Tibetan Plateau, is thus thought to be especially vulnerable and sensitive to global warming. In this paper, the response of plant community and several main species in the alpine meadow of Northewst Sichuan to experimemtal warming was studied by using open-top chambers (OTC). The aim of the this study was to research the warming effects on plant community structure, substance allocation, growth and physiological processes of several mian species, and to explore the biological and ecological mechanism of how the alpine meadow plants acclimate and adapt to future global warming. The results were as follows: 1. Warming effects of OTC The mean soil temperature, soil surface temperature and air temperature in OTC manipulation increased by 0.28℃、0.46℃ and 1.4℃ compared to the control during the growing season. This suggested that the OTC used in our study had increased temperature there. Meanwhile, the OTC manipulation slightly altered thermal conditions, but the same amount of precipitation was supplied to both the OTC manipulation and the control, so higher soil evaporation and plant transpiration in OTC manipulation directly lead to the decrease of soil surface water content. 2. The reponse of community structure to experimental warming The species richness was not changed by the short-term effect of OTC manipulation. However, experimental warming changed the microenvironment of plant community, therefore competitive balances among species were shift, leading to changes in species dominance. In the present study, the dominant plant species in the control plots were some forbs including Potentilla anserine, Geranium pylzowianum, Thlaspi arvense and Arenaria serpyllifolia, however, the importance value of some gramineous grasses including Elymus nutans, Deschampsia caespitosa, Festuca ovina, and some forbs including Euphrasia tatarica and Rumex acetosa significantly increased in OTC. The different biology characteristics and resource utilizations between gramineous grasses and forbs, and enhanced temperature caused change in some environment factors such as soil water content. As a result, the coverage and biomass of gramineous grasses significantly increased in OTC compared to the control, however, the coverage and biomass of forbs singnifciantly decreased in OTC compared to the control. 3. The reponse of plant growing season to experimental warming Both the standing dead and fallen litter biomass in OTC were lower than those in the control in October, and the biomass of aboveground live-vegetation in OTC was higher than that of the control. The results indicated that the senescence of plants was postponed, and the growing season was prolonged in our research. 4. The reponse of community biomass accumulation and its allocation to experimental warming Experimental warming caused the decrease of aboveground live biomass and belowground root biomass except for the aboveground live biomass in October. Experimental warming also had pronounced effects on the pattern of root biomass allocation. In the present study, the root biomass in 0-10cm soil layer increased in OTC manipulation compared to the control, however, the root biomass in the 20-30cm soil layer decreased in OTC manipulation compared to the control. 5. The reponse of community C and N content to experimental warming The C concentration and stock in aboveground live and belowground root both increased in OTC manipulation compared to the control. However, the N concentration and stock in aboveground live and belowground root both decreased in OTC manipulation compared to the control. 6. The reponse of gowth and biomass, C and N alloction of several species to experimental warming Experimental warming significantly increased the height, SLA (specific leaf area) and aboveground biomass of Elymus nutans in OTC manipulation compared to the control. The SLA and total biomass of Rumex acetosa also significantly increased in OTC manipulation compared to control, among the different components of Rumex acetosa, leaf biomass significantly increased, but root biomass significantly decreased in OTC manipulation compared to the control. However, the height, SLA and total biomass of Potentilla anserina significantly decreased in OTC manipulation compared to the control, among the different component of Potentilla anserina, leaf and stem biomass significantly decreased, but root biomass significantly increased in OTC manipulation compared to the control. The LMR (leaf mass ratio), RMR (root mass ratio), R/S (shoot/root biomass ration) and root C concentration of Rumex acetosa significantly increased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively more C and N content to leaf and root in response to experimental warming, however, the SMR (stem mass ration) and root N concentration of Rumex acetosa significantly decreased in OTC manipulation compared to outside control, also, Rumex acetosa allocated relatively less C and N content to stem in response to experimental warming. The RMR and R/S of Potentilla anserina significantly increased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively more C and N content to root in response to experimental warming, however, the SMR and LMR of Potentilla anserina significantly decreased in OTC manipulation compared to outside control, also, Potentilla anserina allocated relatively less C and N content to leaf in response to experimental warming. 7. The reponse of physiological processes of several species to experimental warming Experimental warming significantly increased chlorophyll a, chlorophyll b and total chlorophyll of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control. However, chlorophyll a, chlorophyll b, total chlorophyll and carotenoid of Potentilla anserina in OTC manipulation significantly decreased compared to outside control. Experimental warming had pronounced effects on gas exchange of Elymus nutans, Rumex acetosa and Potentilla anserine. In the present study, warming markedly increased the light response curves of Elymus nutans and Rumex acetosa in OTC manipulation compared to outside control, and also singnificantly increased A (net photosynthesis rate), E (transpiration rate), gs (stomatal conductance), Pmax (maximum net photosynthetic rate), Rday (dark respiration rate), AQY (apparent quantum yield) and LSP (light saturation point), but LCP (photosynthetic light compensation) of Elymus nutans and Rumex acetosa in OTC manipulation singnificantly decreased compared to outside control. However, warming markedly decreased the light response curves of Potentilla anserina in OTC manipulation compared to outside control, and also singnificantly decreased A, E, gs, Pmax, Rday, AQY and LSP, but LCP of Potentilla anserina in OTC manipulation singnificantly increased compared to outside control. Experimental warming singnificantly increased the chlorophyll fluorescence kinetics parameters such as Fv/Fm, Yield and qP of Elymus nutans and Rumex acetosa and qN of Potentilla anserina in OTC manipulation, but Fv/Fm, Yield and qP of Potentilla anserina in OTC manipulation singnificantly decreased. 8. The reponse of antioxidative systems of several species to experimental warming Experimental warming tended to increase the activities of antioxidative enzymes and stimulate the role of non-enzymes of Elymus nutans and Rumex acetosa. As a result, MDA content of Elymus nutans and Rumex acetosa decreased. The activities of antioxidative enzymes and non-enzymes of Potentilla anserina also significantly increased in OTC manipulation, but more O2- was produced because of lower soil water content, and the O2- accumulation exceeded the defense ability of antioxidative systems and non-enzymes fuctions. As a result, MDA content of Potentilla anserine still increased in OTC manipulation compared to outside control.
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大气CO2浓度的增加已经成为不可争议的事实。预计本世纪末大气CO2浓度将增加到约700µmol mol-1。森林年光合产量约占陆地生态系统年光合产量的70%。森林树木是一个巨大的生物碳库,约占全球陆地生物碳库的85%。森林树木对CO2的固定潜力是缓解由大气CO2浓度升高引起的未来全球气候变化问题的决定性因子之一。红桦(Betula albosinensis Burk.)是川西亚高山采伐迹地自然或人工恢复的重要树种。本研究以1a红桦幼苗为模式植物,采用人工模拟的方法,研究CO2浓度升高对不同种内竞争强度(种群水平)下红桦幼苗的生理特征、生长、干物质积累及其分配的影响,探讨在种内竞争生长条件下红桦幼苗的“光合适应机理”与生长特征,为西南亚高山森林生产力对未来全球变化的预测提供重要参考。 本研究的主要结果如下: 1)在种内竞争生长条件下红桦幼苗经过CO2浓度升高熏蒸4个月后,叶片出现“光合适应”现象。与对照相比,低种植密度(28株m-2)和高种植密度(84株m-2)条件下的红桦幼苗净光合速率(A)、气孔导度(gs)、蒸腾速率(E)、表观量子产量(AQY)和羧化速率(CE)显著降低,而水分利用效率(WUE)则显著提高。CO2浓度升高处理的红桦幼苗叶片Rubisco活性、单位叶面积N浓度、叶绿素a、叶绿素b和类胡萝卜素浓度都显著降低。但CO2浓度对红桦幼苗的叶绿素a与叶绿素b的比值没有显著影响。CO2浓度升高显著增加红桦幼苗单位叶面积的非结构性碳水化合物(TNC)浓度,结果是红桦幼苗的比叶面积(SLA,cm2 g-1)显著降低。 2)与对照相比,CO2浓度升高处理的红桦幼苗高、基径、单叶面积和侧枝的相对生长速率(R GR)显著提高,尤其在试验处理的早期。CO2浓度升高既增加单株红桦幼苗总叶片数量又增加单叶面积,结果是单株红桦幼苗的总叶面积比对照显著增加。 3)CO2浓度升高处理显著增加红桦幼苗干物质积累(尤其是细根生物量),改变了红桦幼苗生物量的分配格局。与对照相比,CO2浓度升高处理的红桦幼苗叶重比(LWR)、叶面积比(LAR)、叶根重比(Wl/Wr)和源汇重比(leaf weight to non-leaf weight ratio, Wsource/Wsink)显著下降(高种植密度的LWR除外),而根冠比(R/S)则显著增加。在两种种植密度条件下,CO2浓度升高显著增加红桦幼苗根生物量的分配比率,显著降低叶片的生物量分配比率,对主茎、侧枝以及地上生物量的分配比率不变或约有下降。 总之,长期生长在CO2浓度升高条件下的红桦幼苗光合能力下降,并伴随Rubisco活性、叶N浓度、光合色素浓度的显著降低以及TNC浓度的显著增加。支持树木光合速率下降与Rubisco活性、叶N浓度下降以及TNC浓度增加紧密相关的假设。CO2浓度升高处理红桦幼苗的早期相对生长速率大大高于对照,而后期迅速下降,说明红桦幼苗生物量的显著增加主要归功于CO2浓度升高的早期促进作用和叶面积的显著增加。CO2浓度升高显著增加红桦幼苗根系生物量和根冠比,表明红桦幼苗“额外”固定的C向根系转移。 The steady increae of atmospheric CO2 concentration([CO2])has been inevitable fact. Models predict that the atmospheric [CO2] will increase to about 700µmol mol-1 at the end of the twenty-first century. As trees constitute a majoor carbon reservoir–85% of total plant carbon is found in forest, and their ability to sequester carbon is a key determinant of future global change problems caused by increases in atmospheric CO2. In addition to the role of forests in the global carbon cycle, inceased growth could be of economic benefit, for example, offsetting deleterious effects of climatic changes. Betula albosinensis (Burk.) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of mountain forest area, and also is one of important tree species for afforestation in logged area, in southwesten China. In this experinment, Betula albosinensis seedling (one-year-old) was used as the model plant. B. albosinensis seedlings were grown under two all-day [CO2], ambient (about 350 µmol·mol-1) and elevated [CO2] (about 700 µmol·mol-1), and two planting densities of 28 plants per m2 and 84 plants per m2. The objectives were to characterize birch mature leaf photosynthesis, growth, mass accumulation and allocation responses to long-tern elevated growth [CO2] under the influences of neighbouring plants, and to assess whether elevated [CO2] regulated birch mature leaf photosynthetic capacity, in terms of leaf nitrogen concentration (leaf [N]), activity of ribulose bisphosphate carboxygenase (Rubisco), Rubisco photosynthetic efficiency, and total nonstructural carbohydrates (TNC) concentration, and also to provide a strong reference to predict the productivity of subalpine forests under the future global changes. The results are as follows: 1) B.albosinensis seedlings exposed to elevated [CO2] for 120 days, photosynthetic acclimation phenomena occurred. At two planting densities, leaves of birch seedlings grown under elevated [CO2] had lower net photosynthetic rate (A), stomatal conductance (gs), transpiration (E), apparent quantum yield (AQY) and carboxylated efficiency (CE) and higher water use efficiency (WUE), compared to those of B.albosinensis seedlings grown under ambient [CO2]. Based on the leaf area, leaf [N], Rubisco activity and photosynthetic pigments concentrations of B. albosinensis seedlings grown under elevated [CO2] were significantly lower than those grown under ambient [CO2]. The ratio of chlorophyll a to chlorophyll b concentration was not affected by elevated [CO2]. Under elevated [CO2], the TNC concentration per unit leaf area significantly increased, resulting in significant decrease in specific leaf area. Thus leaf photosynthetic capacity of B. albosinensis seedlings would perform worse under rising atmospheric [CO2] and the influences of neighbouring plants. 2) Under elevated [CO2], the relative growth rate (RGR) of B. albosinensis seedlings height, basal diameter, a leaf area and branch length significantly increased, especially at the initial stage of exposure to elevated [CO2], and a leaf area and leaf numbers per B. albosinensis seedling also significantly increased. Thus the total leaf area per B. albosinensis seedling was significantly increased under elevated [CO2]. 3) As the increase of RGR and total leaf area, biomass of B. albosinensis seedling grown elevated [CO2] was higher, compared to that of B.albosinensis seedlings grown at ambient [CO2]. Elevated [CO2] changed the biomass allocation pattern of B. albosinensis seedling. At two planting densities, B. albosinensis seedlings grown elevated [CO2] had lower leaf weight to total weight ratio (LWR), leaf area to total weight ratio (LAR) and leaf weight to non-leaf weight ratio (Wsource/Wsink), but higher root weight to shoot weight ratio (R/S), compared to those of B.albosinensis seedlings grown at ambient [CO2]. Under elevated [CO2], roots biomass to total biomass ratio was signigicantly increased, leaves biomass to total biomass ratio was significantly decreased. The main stem and branch biomass to total biomass ratio were not affected by elevated [CO2]. In conclusion, our results supported the hypothesis that the decline in photosynthetic capacity of C3 plants will appear after long-term exposure to elevated [CO2], accompanying with the significant decrease in Rubisco activity, leaf N concentration, photosynthetic pigments concentration, and significant increase in total non-structural carbohydrates concentration. Our results also have shown that the increase of biomass of B. albosinensis seedlings should be attributed to initial stimulation on RGR and total leaf area resulted from elevated [CO2]. Under elevated [CO2], the extra carbon sequestered by B.albosinensis seedlings transferred into under-ground part because of increase in root biomass and R/S.
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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.
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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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随着全球气候变暖和温室效应加剧,干旱和荒漠化成为威胁人类生存和发展的主要 灾害,许多被子植物对干旱胁迫的生理、生态和生化响应已逐步得以报道,但很少有开 展干旱胁迫对雌雄异株植物的影响方面的研究。由于这类植物在长期进化过程中已经在 生长、性比、生殖格局、空间分布、资源配置和生物量分配等方面形成了明显的性别差 异,因此,干旱胁迫必将对其雌雄植株产生不同的生理生态影响。本研究以青杨为模式 植物,采用植物生态、生理及生物化学等研究方法,系统研究青杨雌雄植株在常温、增 温以及喷施外源脱落酸的条件下对干旱胁迫的响应,揭示其在生长形态、生物量分配、 光合作用、用水效率和生理生化等方面的性别间差异。主要研究结果如下: 1. 青杨雌雄植株对干旱胁迫的综合响应。 与较好水分条件相比,干旱胁迫显著降低了青杨雌雄植株的光合作用和生长发育, 影响了许多生理生化过程,并导致雌雄植株在生长发育、气体交换、用水效率、膜脂抗 氧化和抗氧化系统酶活性方面表现出显著的性别间差异。在较好水分条件下,雌雄植株 之间在株高、基径、生物量、净光合速率、蒸腾速率、用水效率以及丙二醛、脱落酸和 游离脯氨酸等生化物质含量方面均无显著差异。但在干旱胁迫下,雄株在生长发育、气 体交换、水分利用效率、膜脂过氧化保护和抗氧化系统酶活性方面均显著高于雌株,表 现出比雌株更高的株高、基径、叶面积、总叶片数、总生物量、总色素含量、类胡萝卜 素含量、净光合速率、蒸腾速率、羧化效率、光系统II最大光化学效率、内在水分利用 效率、碳同位素组分、过氧化氢酶和过氧化物酶活性等,而在CO2补偿点、比叶面积、 叶绿素a/b、丙二醛、脱落酸和超氧化物歧化酶活性等指标上显著低于雌株。与雌株相比, 雄株表现出更高的干旱胁迫适应能力,而雌株的生长发育和生理生化过程更易遭受干旱 胁迫的影响。 2. 干旱胁迫下的青杨雌雄植株对增温处理的综合响应 与环境温度相比,增温在干旱胁迫前后均显著促进了雌雄植株的生长发育、气体交 换,降低水分利用效率,影响生化物质含量,并促使青杨雌雄植株之间在干旱胁迫下表 现出显著的差异。在较好水分条件下,增温导致雌株的株高、基径、叶面积、总叶片数、 总生物量和超氧化物歧化酶活性显著高于雄株,而用水效率、丙二醛、脱落酸和游离脯 氨酸、抗坏血酸过氧化物酶和过氧化物酶活性低于雄株。在干旱胁迫下,增温将导致雄 株的株高、基径、叶面积、总生物量、净光合速率、蒸腾速率、气孔导度、总色素含量、 相对含水量、过氧化氢酶和抗坏血酸过氧化物酶活性等显著高于雌株,而光系统II 最大 光化学效率、内在水分利用效率、碳同位素组分、丙二醛、脱落酸、游离脯氨酸和超氧 化物歧化酶活性显著低于雌株。与雄株相比,水分较好条件下的增温有利于促进雌株的 生长发育,并在生理生态特征上优于雄株。而干旱胁迫下的增温则加剧了水分胁迫强度, 致使雌株的生长发育遭受比雄株更多的负面影响。 3. 干旱胁迫下的青杨雌雄植株对喷施外源脱落酸处理的综合响应 与对照相比,在干旱胁迫下喷施外源脱落酸可显著增加青杨雌雄植株的生长发育、 气体交换、降低水分利用效率,影响了生化物质含量,并导致青杨雌雄植株之间在干旱 胁迫下表现出显著的生理生态差异。在干旱胁迫下,喷施外源脱落酸致使雌株的株高、 叶面积、叶干重、细根干重、总生物量、净光合速率、蒸腾速率、气孔导度、光系统II 最大光化学效率、非光化学淬灭系数、相对含水量、总光合色素、类胡萝卜素、脱落酸、 超氧化物歧化酶和过氧化物酶活性的增加量显著高于雄株,而根重比、根冠比、细根/ 总根、比叶面积、内在水分利用效率、碳同位素组分、丙二醛、脯氨酸、过氧化氢酶和 抗坏血酸过氧化物酶活性等指标的减少量上显著低于雄株。与对照相比,干旱胁迫下的 喷施外源脱落酸则一定程度能减缓植株遭受胁迫的压力,促进植株生长和气体交换,减 少了植株体内的过剩自由基数量,并促使雌株的生长发育和光合能力显著提高,增强其 抗干旱胁迫能力。 With development of global warming and greenhouse effect, drought and desertification have been became main natural disasteres in resent years. Studies on ecophysiological responses of most angiosperm species to environmental stress have been reported, but little is known about dioecious plant responses to drought stress. Since significant differences on growth, survival, reproductive patterns, spatial distribution, as well as resource allocation between males and females of dioecious plant have been formed during evolutionary process, sexual different ecophysiological responses should be caused by drought stress. In this experiment, Populus cathayana Rehd. was used as model plant to study the sex-related responses to drought by using the ecological, physiological and biochemical methods under normal atmospheric temperature, elevated temperatures and exogenous abscisic acid (ABA) application treatment respectively, and to expose the sexual differences in growth, biomass allocation, photosynthesis, water use efficiency and some biochemical material contents in the males and females of dioecious plant. The results are follows: 1. A large set of parallel responses of males and females of P. cathayana to drought stress Compared with well-watered treatment, drought significantly decreased growth and photosynthesis of P. cathayana individuals, affected some physiological and biochemical processes, and induced males and females to exhibit obvious sexual differences in growth, gas exchange, water use efficiency, lipid peroxidation protection and antioxidant defenses enzyme system. Under well-watered treatment, there were no significant sexual differences in height growth (HG), basal diameter (BD), dry matter accumulation (DMA), net photosynthesis rate (A), transpiration (E), water use efficiency (WUE), and malondialdehyde (MDA), abscisic acid (ABA) and praline (Pro). However, under drought stress, males were found to exhibit higher HG, BD, leaf area (LA), total leaf number (TLA), DMA, total chlorophyll contents (TC), carotenoids content (Caro), A, E, carboxylation efficiency (CE), the maximum efficiency of PSII (Fv/Fm), intrinsic water use efficiency (WUE ), carbon isotope composition (δ13C), catalase (CAT), peroxidase (POD) and lower CO2 compensation point (Γ), specific leaf area (SLA), chlorophyll a/b ratio (Chla/Chlb), MDA, ABA and superoxide dismutase (SOD) than females. The results suggest that males possess greater drought resistance than do females and females suffer more negative effect on growth and development, physiological and biochemical processes than males under drought stress. 2. A large set of parallel responses of drought-stressed males and females of P. cathayana to elevated temperatures Compared with environmental temperature, elevated temperature treatment significant increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents of P. cathayana individuals, and induced males and females to exhibit obvious differences under drought stress. Under good water condition, elevated temperature treatment caused females to show significant higher HG, BD, LA, TLN, DMA, SOD activity, and great lower WUE, MDA, ABA, Pro, ascorbate peroxidase (APX) and POD than do males. On contrary, under drought condition, elevated temperature treatment induced males to exhibit higher HG, BD, LA, DMA, A, E, stomatal conductance (gs), relative water content (RWC), CAT, APX activity but lower Fv/Fm, WUE, δ13C, MDA, ABA, Pro, SOD activity than do females. The results suggest that females will benefit from elevating temperature under good water condition by possessing better ecophysiological processes than that of males, but will suffer from greater negative effects than do males when grown under drought stress with elevated temperature treatment. 3. A large set of parallel responses of drought-stressed males and females of P. cathayana to exogenous ABA application Compared with controls, exogenous ABA application under drought greatly increased growth and gas exchange, decreased water use efficiency, changed some biochemical material contents in P. cathayana individuals, and induced males and females to exhibit obvious sexual differences under drought. Under drought stress, exogenous ABA application induced females to exhibit more increases in HG, LA, leaf weight (LW), fine root weight (FRW), DMA, A, E, g, Fv/Fm, non-photochemical quenching coefficient (qN), RWC, TC, Caro, ABA, SOD, POD s activity than males, but to show lower decreases in root/weight ratio (RWR), root mass/foliage area ratio (RF), fine root/total root ratio (FT), SLA, WUE, δ13C, MDA, Pro, CAT, APX than males. The results suggest that exogenous ABA application under drought stress will eliminate negative damages caused by drought stress at a certain extent,promote the growth and gas exchange of plant and decrease the number of superfluous 1O2 in plant cells of males and females of P. cathayana. Furthermore, exogenous ABA application promoted more drought resistance in females than in males by increasing more growth and photosynthetic capacity in females under drought stress.
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近年来,随着对作物重茬(连年种植)障碍原因的深入研究,植物的化感作用越来越受到国内外众多学者的重视。而作为重要调料和药用植物的生姜,其连作障碍也备受关注,系统地研究生姜化感作用将有助于理解和最终解决生姜连作障碍问题。本文通过研究生姜不同部位、不同浓度的水浸液对与其间作的两个物种(大豆和四季葱)种子的萌发及幼苗生长的影响,从而证明生姜化感作用的存在;并通过温室盆栽实验研究了生姜的自毒作用(即研究生姜不同部位、不同浓度的水浸液对其幼苗的形态、生理生化、光合作用、土壤酶、土壤微生物多样性及土壤养分的影响),从而揭示生姜退化和衰老的机制,并为生姜筛选出合适的间作物种提供科学依据,对生姜连作障碍提出科学的解决方法。主要研究结果如下: 1. 与对照相比,生姜所有部位(根茎、茎、叶)、所有浓度(10、20、40、 80 g l-1)的水浸液均抑制了大豆种子和葱籽的萌发率、幼苗生长、水分吸收和脂肪酶活性,并且其抑制程度随着水浸液浓度的增加而增强,其生姜各部位水浸液抑制效应的强弱顺序为茎>叶>根茎。这一结果表明生姜根茎、茎、叶含有能够抑制大豆种子和葱籽种子萌发和幼苗生长的水溶性化感物质。根茎是生姜的主要收获部位,而生姜的残株(主要是茎和叶)应该从大田中处理掉以减轻其抑制效应。生姜水浸液中主要化感成分包括:根茎水浸液中主要是丁香酸和伞花内脂;茎水浸液中主要是阿魏酸,且其含量最高为73.4 ug/g;叶水浸液中除了阿魏酸,其他六种物质均检测出来,但含量较高的主要有丁香酸、伞花内脂和香豆酸。 2. 生姜茎和叶不同浓度的水浸液均显著抑制了生姜幼苗的株高、每株叶片数和叶面积,其抑制程度随着水浸液浓度的增加而有所增强,而生姜幼苗每株分枝数差异不显著;同时生姜水浸液也极大程度地影响了生姜幼苗的生物量(包括地下生物量、地上生物量和总生物量,均为鲜重)。在同一浓度下,茎水浸液对生姜幼苗形态指标及生物量指标均显示出最强的抑制作用,叶水浸液次之,根茎水浸液最弱。与对照相比,低浓度的生姜根茎水浸液提高了生姜幼苗叶片内四种抗氧化酶(SOD、POD、CAT、APX)活性,高浓度的根茎水浸液抑制了四种抗氧化酶活性,而茎和叶水浸液均随着浓度的增加而抑制了四种抗氧化酶活性,三种水浸液均随着浓度的增加降低了生姜幼苗叶片内叶绿素的含量,而增加了生姜幼苗叶片的相对电导率和丙二醛含量。同时,三种水浸液均随着浓度的增加降低了生姜幼苗的光合参数(包括胞间CO2浓度、气孔导度、蒸腾速率及净光合速率)。 3. 三种生姜水浸液对所测六种土壤酶活性均产生了不同程度的影响,其中影响最大的是酸性磷酸酶和蔗糖酶,在10 g l-1 时就达到了显著水平,并且所有酶均有随着水浸液浓度增加而增大的趋势;相同部位的水浸液随着浓度的增加,细菌和真菌的数量呈增加趋势,而放线菌的数量呈减少趋势;三种生姜水浸液均随着浓度的增加降低了土壤中有机质的含量,加剧了土壤中硝态氮含量的积累,根茎水浸液对土壤有效磷、速效钾和铵态氮均显示出低浓度提高其含量而高浓度降低其含量的趋势,而茎和叶水浸液则随着浓度的增加均降低了其含量。 4. 与生姜单作相比,所有间作系统均在旺盛生长期和收获期不同程度地提高了土壤酶活性,同时也增加了土壤细菌数量及土壤微生物总数但不显著;所有间作系统在旺盛生长期和收获期均不同程度地影响了土壤真菌及放线菌数量(增加或减少),所有间作系统间的多样性指数差异不显著,除了旺盛生长期四种作物(生姜-大豆-四季葱-大蒜)的间作模式显著降低了多样性指数,其值仅为生姜单作的33.18%;生姜与大豆间作不仅提高了19.6%的生姜产量而且获得了较好的经济效益,并且,所有间作系统均显著抑制了生姜姜瘟病的发生。 5. 不同栽培模式不同程度地影响了收获期生姜的株高、分枝数、根茎产量及内在品质。其中处理2显著地促进了生姜的分枝(10.5%),同时处理2、3和4也促进了生姜的生长(株高分别增加了15.0%、11.4%和14.0%),并且这三个处理提高了生姜的产量;处理2和3能有效提高生姜块茎中维生素C(分别较单作生姜显著提高了3.29%和4.05%)、处理3显著提高了可溶性糖(8.2%)、姜辣素(4.6%)和蛋白质等有益物质的含量,降低硝酸盐有害物质的含量(处理2显著降低了14.0%),改善了姜块的外观和内在品质。并且,生姜与大豆间作具有最高的纯收入和产投比,分别较生姜单作提高了24.80%和8.8%。Recently, allelopathy has been more and more paid attentions by national and foreign scholars with profound research on reasons of crop replanted (continuous planted) obstacle. Ginger rhizome is valuable all over the world either as a spice or herbal medicine and ginger replanted obstacle is also paid attentions. Systematic research on ginger allelopathy will contribute to understanding and ultimate solving problem of ginger replanted obstacle. The effects of ginger aqueous extracts with different parts and concentrations on seed germination and early seedling growth of soybean and chive were studied in this article to testify that ginger existed allelopathy. Furthermore, ginger autotoxicity was also studied by pot experiment in greenhouse (namely research on effects of ginger aqueous extracts with different parts and concentrations on morphological indexes, physiological and biochemical indexes, photosynthesis, soil enzymes, soil microbial diversity and soil nutrients) to reveal mechanism of ginger degeneration and senescence, provide scientific basis for selecting appropriate intercropping species and put forward scientific resolvent for ginger replanted obstacle. The main results were as follows: 1. All aqueous extracts at all concentrations inhibited seed germination, seedling growth, water uptake and lipase activity of soybean and chive compared with the control, and the degree of inhibition increased with the incremental extracts concentration. The degree of toxicity of different ginger plant parts can be classified in order of decreasing inhibition as stem>leaf>rhizome. The results of this study suggested that rhizome, stem and leaf of ginger contained water soluble allelochemicals which could inhibit seed germination and seedling growth of soybean and chive. The rhizome is the main harvested part of ginger. The residue (mainly stems and leaves) of the ginger plant should be removed from the field so as to diminish its inhibitory effect. The main allelopathic components of three kind of aqueous extracts were as follows: Rhizome extract chiefly contained syringic acid and vmbelliferone and stem extract mainly contained frulic acid whose content was the highest (73.4 ug/g). The other six substances were detected except of frulic acid, but only contents of syringic acid, vmbelliferone and p-coumaric acid were higher. 2. Stem and leaf aqueous extracts of ginger with different concentrations significantly inhibited plant height, leaf numbers per plant and leaf area, and the degree of inhibition increased with the incremental extracts concentration. However, tiller number per plant of ginger seedling showed no significant difference. At the same time, ginger aqueous extracts also influenced biomass including under-ground biomass, above-ground biomass and total biomass (fresh weight) to a large extent. Under the same concentration, stem aqueous extract showed the mostly inhibitory effect on morphological indexes and biomass indexes of ginger seedling. Rhizome aqueous extract showed the leastly inhibitory effect and leaf aqueous extract was intervenient. Enhanced concentration of ginger aqueous extracts significantly reduced total chlorophyll content, accompanying with increases in memberane permeability (REL) and lipid peroxidation (MDA). Compared with the control, rhizome ginger aqueous extract of lower concentration (10 g l-1) increased the activities of major antioxidant enzymes (superoxide dismutase, SOD; peroxidase, POD; catalase, CAT; ascorbate peroxidase, APX) of ginger leaf tissue and higher concentration inhibited the activities of four antioxidant enzymes. However, stem and leaf aqueous extract inhibited the activities of four antioxidant enzymes with increase in concentration. Meanwhile, enhanced concentration of ginger aqueous extracts significantly reduced photo-parameters of ginger seedling (including CO2 concentration, stoma conductivity, net photosynthesis rate and transpiration rate). 3. Rhizome, stem and leaf ginger aqueous extract showed different effect on six soil enzyme activities, and acid phosphatase and invertase showed significant effect when aqueous extract concentration got 10 g l-1. Furthermore, six soil enzyme activities increased with increase in aqueous extract concentration. Bcterial and fungi number tended to increase while antinomyces tented to decrease with the increase in aqueous extract concentration of identical part. Ginger aqueous extracts reduced soil organic matter content with increased concentration, accompanying with NO3-—N accumulation in soil. Rhizome aqueous extract showed the same tendency for available P, available K and NH4+—N, namely lower concentration increased their contents in soil and higher concentration reduced their contents. While stem and leaf aqueous extracts reduced their contents with the increamental concentration. 4. All intercropping systems increased soil enzyme activities to different extent both at VGS and at HS compared to solo ginger. All intercropping systems increased the colony numbers of soil bacteria and total of soil microbe but not significantly either at VGS or at HS. All intercropping systems increased the colony numbers of soil fungi and actinomytes to a different extent (increase or decrease) both at VGS and at HS. For DI, difference between all cultivation patterns and S-G was not significant either at VGS or at HS except that G-S-C-G whose value was only 33.18% of S-G at VGS significantly decreased. G-S not only increased ginger yield by 19.6% but also obtained better economic benefit. Furthermore, all intercropping systems significantly inhibited occurrence of bacterial wilt of ginger. 5. Different cultivated pattern influenced plant height, tiller numbers, rhizome yields and intrinsic quality of ginger. Treatment 2 significantly facilitated tiller occurring (10.5%). Treatment 2, 3 and 4 promoted ginger growth (plant height respectively increased 15.0%、11.4% and 14.0%) and enhanced rhizome yields. Treatment 2 and 3 effectively increased vitamin C content (significantly increased 3.29% and 4.05% compared to solo ginger). Treatment 3 significantly increased contents of beneficial substances such as soluble sugar (8.2%), gingerols (4.6%) and protein. Treatment 2 significantly decreased contents of deleterious substance namely nitrate (14.0%) and improved appearance and intrinsic quality of ginger rhizome. Furthermore, treatment 2 (ginger/soybean intercropping) could obtain better economic benefit and showed the highest net income and ratio of benefit and cost whose values respectively increased by 24.80% and 8.8% compared to solo ginger.
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光是植物赖以生存的重要环境因子,但是植物在获得光的同时不可避免的会受到紫外辐射的伤害。尤其是近年来,人类向大气中排放的大量氮氧化合物和氟氯烃类化合物(CFC’s)引起臭氧分子的分解,导致到达地球表面的紫外辐射增加,特别是UV-B辐射增强。而另一方面,植物对UV-B辐射反应的敏感性在种间和品种间存在差异,主要受植物基因型,生态型和生活型的控制。本项目分别以粗枝云杉和青杨组杨树为模式植物,从形态和生理生化方面分别研究了来自不同水分背景下的粗枝云杉种群和来自不同UV-B背景下的青杨种群在增强UV-B下的反应及其反应差异,并探讨了干旱、喷施外源脱落酸(ABA)对它们抗UV-B能力的影响。研究成果可为生态系统的恢复与重建提供理论依据和科学指导。主要研究结果如下: 1. 粗枝云杉的两个种群,湿润种群(来自四川黑水)和干旱种群(来自甘肃迭部)在水分良好和干旱状况下表现出对增强UV-B的不同响应。同时,干旱对粗枝云杉抗UV-B能力的影响也得到研究:两种胁迫共同作用时,干旱表现出在一定程度上减弱了增强UV-B对粗枝云杉的生理特性的影响。 干旱胁迫显著降低了两个粗枝云杉种群的光合同化速率(A), 气孔导度(gs)和PSII的有效光量子产量(Y), 同时,提高了非光化学猝灭效率(qN)和超氧化物歧化酶(SOD)的活性。与湿润种群相比,干旱种群抗旱性更强,表现为干旱种群拥有更高的SOD和干旱进一步加剧了UV-B的胁迫效应。 本研究中,干旱胁迫单独作用时,显著降低了青杨两个种群的生物量积累和气体交换,具体包括A、gs、蒸腾速率(E)和光合氮利用效率(PNUE),提高了两个种群的瞬时水分利用效率(WUEi)、长期水分利用效率(WUET)、碳同位素组分(δ13C)和氮含量(N)。同时,UV吸收物质和ABA含量也得到积累。另一方面,增强UV-B对青杨两个种群各个指标的影响,同干旱所引起的效应有着相似的趋势。同低海拔种群相比,高海拔种群有着更强的抗旱和抗UV-B能力,具体表现在高海拔种群有着更多的生物量积累,更强的气体交换和水分利用效率及更高水平的ABA和UV吸收物质含量。相比干旱诱导的生物量积累和气体交换的降低,在干旱和增强UV-B两个胁迫同时作用于青杨时,这种降低表现的更为明显。显著的干旱和UV-B的交互作用还表现在WUEi, WUET, δ13C, 可溶性蛋白含量, UV吸收物质含量, ABA, 叶片和茎中的N含量以及C/N比中。 3. 经过一个生长季的试验观察,增强UV-B、外源ABA及两因子共同作用对青杨的生物量积累、气体交换、内源ABA和UV吸收物质含量、抗氧化系统以及碳、氮含量和碳/氮比均产生显著影响。本试验中,青杨的两个种群分别来自中国西南部的不同海拔地区,高海拔种群来自青海大通而低海拔种群来自四川九寨。外源ABA的胁迫为直接喷施ABA到青杨叶片,而增强UV-B胁迫是利用平方波系统分别保证青杨苗暴露于外界UV-B强度和两倍于外界UV-B强度下。 研究结果显示,增强UV-B显著的降低了两个青杨种群的株高、基茎、总叶面积和总生物量等生长指标,同时也导致其A、gs、E和叶片中碳含量的减少。而显著增加了SOD和过氧化物酶(GPx)活性水平,诱导了过氧化氢(H2O2)和MDA的显著增加,促进了UV吸收物质和不同器官中内源ABA含量的显著积累。另一方面,外源ABA引起了青杨光合同化速率的下降,SOD和GPx酶活性的增强,H2O2 和 MDA含量也表现出显著增加,同时,内源ABA含量得到显著累积。同低海拔种群相比,高海拔种群具有更加抗UV-B和外源ABA的特性。显著的UV-B和ABA的交互作用表现在A, E, SOD和GPx活性,以及叶片和根部的内源ABA等一系列指标中。在所有胁迫下,叶片中的碳和氮含量同其在茎和根中的含量显著相关,另外,叶片和茎中的氮含量同茎中的碳含量显著相关。 Sunlight is an indispensable environment factor for plants survival and development. Meanwhile, photosynthetic organisms need sunlight and are thus, inevitably, exposed to UV radiation. Especially for recent years, ultraviolet radiation, especially UV-B reaching the Earth’s surface increased because of depletion of ozone layer resulted from emission of NxO and CFC’s from human activities. On the other hand, the sensitivity of plants to UV-B radiation depends on the species, developmental stage and experimental conditions. In this experiment, two populations of Picea asperata Mast from different water background and two populations of Populus cathayana Rehder from different altitude background were selected as model plants to assess the effects of enhanced UV-B radiation. Morphological and physiological traits induced by enhanced UV-B in each plant species were observed and the different responses were discussed, furthermore the influences of drought and exogenous ABA on responses induced by enhanced UV-B were studied. The study could provide a strong theoretical evidence and scientific direction for the afforestation and rehabilitation of ecosystem. The results are as follows: 1. Different responses of two contrasting Picea asperata Mast. populations to enhanced ultraviolet-B (UV-B) radiation under well-watered and drought conditions were investigated. And the effects of enhanced UV-B on tolerance of drought were also observed in our study that the UV-B exposure may have alleviated some of the damage induced by drought. Two contrasting populations, originating from a wet and dry climate region in China, respectively, were employed in our study. Drought significantly decreased CO2 assimilation rate (A), stomatal conductance (gs) and effective PSII quantum yield (Y), while it significantly increased non-photochemical quenching (qN) and the activity of superoxide dismutase (SOD) in both populations. Compared with the wet climate population, the dry climate population was more acclimated to drought stress and showed much higher activities of SOD and ascorbate peroxidase (APX), and much lower levels of malondialdehyde (MDA) and electrolyte leakage. On the other hand, enhanced UV-B radiation also induced a significant decrease in the chlorophyll (Chl) content in both populations under well-watered conditions, and a significant increase in UV-absorbing compounds in the wet climate population. After one growing season of exposure to different UV-B levels and watering regimes, the increases in MDA and electrolyte leakage, as induced by drought, were less pronounced under the combination of UV-B and drought. In addition, an additive effect of drought and UV-B on A and gs was observed in the wet climate population, and on the activity of APX and qN in the dry climate population. 2. The significant effects of drought, enhanced UV-B radiation and their combination on Populus cathayana Rehd. growth and physiological traits were investigated in two populations, originating from high and low altitudes in south-west China. Our results showed that UV-B acts as an important signal allowing P. cathayana seedlings to respond to drought and that the combination of drought and UV-B may cause synergistically detrimental effects on plant growth in both populations. In both populations, drought significantly decreased biomass accumulation and gas exchange parameters, including A, gs, E and photosynthetic nitrogen use efficiency (PNUE). However, instantaneous water use efficiency (WUEi), transpiration efficiency (WUET), carbon isotope composition (δ13C) and nitrogen (N) content, as well as the accumulation of soluble protein, UV-absorbing compounds and abscisic acid (ABA) were significantly increased by drought. On the other hand, cuttings from both populations, when kept under enhanced UV-B radiation conditions, showed very similar changes in all above-mentioned parameters, as induced by drought. Compared with the low altitude population, the high altitude population was more tolerant to drought and enhanced UV-B, as indicated by the higher level of biomass accumulation, gas exchange, water-use efficiency, ABA concentration and UV-absorbing compounds. After one growing season of exposure to different UV-B levels and watering regimes, the decrease in biomass accumulation and gas exchange, induced by drought, was more pronounced under the combination of UV-B and drought. Significant interactions between drought and UV-B were observed in WUEi, WUET, δ13C, soluble protein, UV-absorbing compounds, ABA and in the leaf and stem N, as well as in the leaf and stem C/N ratio. 3. During one growing season, significant effects induced by enhanced UV-B radiation, exogenous ABA and their combination on biomass accumulation, gas exchange, endogenous ABA and UV-absorbing compounds concentrations, antioxidant system as well as carbon (C) content, nitrogen (N) content and C/N ratio were investigated in two contrasting Populus cathayana populations, originating from high and low altitudes in south-west China. Exogenous ABA was sprayed to the leaves and enhanced UV-B treatment was using a square-wave system to make the seedlings under ambient (1×) or twice ambient (2×) doses of biologically effective UV-B radiation (UV-BBE). Enhanced UV-B radiation significantly decreased height, basal diameter, total leaf area, total biomass, A, gs, E and carbon (C) content in leaves, and significantly increased activities of SOD and guaiacol peroxidase (GPx), hydrogen peroxide (H2O2) and malonaldehyde (MDA) content as well as the accumulation of UV-absorbing compounds and endogenous ABA concentrations among different organs in both populations. In contrast, exogenous ABA showed significant decrease in A and significant increases in activities of SOD and GPx, H2O2, MDA content and the endogenous ABA concentrations. Compared with the low altitude population, the high altitude population was more tolerant to enhanced UV-B and exogenous ABA. Significant interactions between UV-B and ABA were observed in A, E, activities of SOD and GPx, as well as in endogenous ABA in leaves and roots of both populations. Across all treatments, C and N content in leaves was strongly correlated with those were in stems and roots, respectively. Additionally, leaf and stem N content were significant correlated with stem C content.
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雌雄异株植物对环境的不同响应一直是一个有趣而新颖的研究领域,由于雌雄个体不同的繁殖成本及不同的生存策略,使得雌雄植株在生长、存活、生殖格局、空间分布、资源配置等方面已经表现出明显的不同,在生理和分子水平上也表现出明显的性别间差异。干旱是制约农林业发展的环境因子之一,叶锈病是对杨树危害最严重的病害之一,由于长期进化的结果,不同性别的植物必然对生物和非生物胁迫有着不同的响应。本文以雌雄异株的青杨为模式植物,研究雌雄间在生理、生化、亚细胞结构和蛋白质水平上对生物和非生物胁迫的差异响应。主要研究结果如下: (1) 青杨雌雄植株对锈病胁迫的生理生化差异响应 在正常的对照组中,雄株叶片比雌株叶片有着较高的活性氧自由基产生速率、较高的SOD、POD、PPO 和较低的CAT 活性;在锈病感染的早期阶段, SOD、POD、CAT 活性、活性氧自由基产生速率、H2O2 含量、膜脂过氧化程度和细胞膜的电渗率在雌雄株中都增加,而PPO 仅在雄株中增加明显,APX 仅在雌株中增加明显,并且雌株比雄株有着更严重的锈病感染程度、细胞膜的伤害程度和光合系统II 的破坏程度,雌株有更多的净光合速率、气孔导度和叶绿素a 含量的降低,在同工酶变化上,雌雄间对锈病也显示出不同的表达模式。结果显示,雄株比雌株对锈病有着更好的抗性和更有效的ROS 清除系统。 (2) 青杨雌雄植株对干旱胁迫的生理生化及亚细胞结构的差异响应 与较好水分条件相比,干旱下雄株比雌株有着更高的A-Ci 响应参数,如Rubisco 最大羧化速率、光呼吸速率、暗呼吸速率和最大电子传递速率等。干旱显著地增加了膜脂过氧化程度和游离脯氨酸含量,并且雄株比雌株表现出较低的膜脂过氧化程度,较高的总蛋白和游离脯氨酸含量。无论是中度干旱还是极度干旱,除了CAT 外,雄株比雌株表现为较强的抗氧化酶活性,在同工酶谱带上,雌雄间表现出不同的变化模式,并且有些条带是干旱影响应的,而有些条带是性别特异性的,这些性别特异性条带能够作为鉴定性别快速而准确的标记。干旱显著地影响了线粒体、叶绿体和细胞壁的结构,尤其在中度干旱胁迫下,雄株线粒体和叶绿体比雌株呈现出较好的完整性,并且雄株细胞壁要比雌株更厚。因此, 雄株比雌株表现出更强的干旱忍耐性和更高效的抗氧化酶系统。 (3) 青杨雌雄植株对干旱胁迫的蛋白质组差异响应 用双相电泳检测到雌雄间近1000 个蛋白点,通过对比发现对照组雌雄间有54 个差异蛋白点,干旱下雌雄间有108 个差异点,其中102 个被质谱成功鉴定。对照组雌雄间的差异蛋白主要集中在与光合作用相关蛋白、抗氧化酶、胁迫防御蛋白和一些调节基因表达的蛋白;干旱胁迫下雌雄间差异蛋白明显增多,主要有参与信号转导、调节基因表达、蛋白质加工、转录产物的转录翻译后修饰的调节性蛋白蛋白和参与氧化还原平衡、抗胁迫、细胞壁合成、光合作用、能量代谢、氨基酸代谢和脂肪酸代谢等的功能性蛋白。干旱下这些蛋白的表达量在雌雄中有的表现出相同的表达模式,如干旱下雌雄株中Rubisco 激活酶、小热激蛋白等表达都增加,而有的表现出相反的表达模式,如Rubisco 大亚基的降解片段、羰酸酯酶等在雄株中表达量上调而在雌株中却是下调。因此,雌雄间在蛋白质水平上对干旱胁迫响应的差异是显著的,也是复杂的。 It is an interesting and novel topic that dioecious plants possess different responses to environmental stress. As for the different productive cost and different survive strategy, different sexual plants have shown obviously morphological, physiological and molecular differences. Drought is one of the most worldwidely important environmental stress factors that limit plant growth and ecosystem productivity. Rust disease is one of the economically important diseases in many trees. As a result of the long evolutionary process, male and female plants should show different responses to abiotic and biotic stress. In this paper, using a dioeious tree of Populus cathayana Rehd as a model, we study the sexual differences to drought and rust disease stress in physiological, biochemical, sub-cellular and proteomics levels. The main results are follows: (1) The sexual differences in physiology and biochemistry of poplar to rust disease In controls, males showed higher production of superoxide radicals, higher activities of SOD, POD, PPO and lower CAT activity. Under rust disease, the activities of antioxidant, the content of ROS and the degree of cellular member destroyed were increased in both sexes, except for PPO in diseased males and APX in diseased females. However, females showed more seriously disease severity and cellular member and PS II destroyed degrees. Net photosynthesis rate, transpiration rate and chlorophyll a content were decreased more in diseased females than in males. There were also some different changes inantioxidant isozymes under rust disease. The results suggested that male poplar possessed a more effectively antioxidant system and were more resistant to rut disease than females. (2) The sexual differences in physiology and biochemistry of poplar to drought stress Under drought stress, there were higher rates of RuBP-saturated CO2 assimilation, dark respiration, photorespiratory release of oxygen, the max electron transportrate in CO2-saturated and carboxylation efficiency in males than in females. And males showed lower TBARS and higher proline content. Except for CAT, the activities of other antioxidants were higher in males than in females. Meanwhile, there were obviously differences in isozyme changes between teo sexes. Drought stress obviously destroyed the integralities of chloroplasts and mitochondria and the sexual differences in sub-cellular level were obviously under the moderate water stress. Male cell walls were more sensitive to drought stress than did female. The results suggested males were more resistant to drought stress. (3) The sexual differences in proteomics of poplar to drought stress By 2-D and MS analysis, we identified 102 different protein spots between males and females. Under control conditions, the different proteins were mainly in photosynthesis related proteins, antioxidants, stress response proteins and some gene expression related proteins. Under drought stress, the different proteins were focused on (i) regulated proteins such as signaling conduction, kinase, HSP, gene expressional regulation and protein modification, (ii) functional proteins such as photosynthesis, energy metabolism, antioxidant, redox, stress response, lipid metabolism and amino acid metabolism. Some protein showed the same expressional pattern, while some showed contrary expressional pattern. Thus, the results suggested that sexual differences in proteomics were significant and complex.
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生长、含水量变化和水分张力变化都能引起树木直径的变化,树木直径变化表现为长期的不可逆生长和短期的昼夜变化。因为木质部、韧皮部和树皮的不同功能,它们膨胀和收缩的速度也不相同。木质部直径的变化与木质部液流相关,而韧皮部和树皮直径的变化不仅和木质部液流相关,还与韧皮部储存水以及木质部与韧皮部间的水分交换有关。
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变水处理与短期遮荫对棉花水分利用效率的影响研究表明 ,在充分供水 -水分胁迫 -复水的变水处理过程中 ,短期遮荫使净光合速率、蒸腾速率、水分利用效率降低 ;光照 -遮荫 -光照期间 ,总的水分利用效率以水分胁迫的最高 ,充分供水的次之 ,复水处理最低 ,其中遮荫 40 %的水分利用效率高于遮荫 75%的处理 .回归分析表明 ,水分利用效率与光合速率表现出极显著正相关 ,两者回归直线的斜率随水分胁迫而增大 ,而水分利用效率与蒸腾速率的关系较弱 ,多数处理两者相关达不到显著水平
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通过对干湿交替环境下春小麦、马铃薯、大豆和玉米等作物的产量、水分利用效率及光合作用、蒸腾作用、气孔导度等生理变化的研究表明 :( 1 )春小麦和马铃薯在干湿交替环境下可获得与充分供水相当的产量而它们的水分利用效率却显著提高 ,大豆减产幅度较大 ,玉米减产严重 ,其水分利用效率显著低于全湿处理 ;( 2 )浇水后各作物的光合速率、蒸腾速率和气孔导度都有所增加 ,但不同作物增加的幅度不同 ,就是同一作物各指标的增幅也不同 ;( 3)干湿交替环境下同化物的运输模式有利于春小麦籽粒的充实和马铃薯块茎的膨大 ,而不利于玉米产量的形成 ;( 4 )产量不仅决定于营养生长阶段 ,更主要决定于生殖生长阶段。此外 ,还就干湿交替过程中若干生理变化和经济产量形成机制作了初步探讨。
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本文在Eagleson随机动力学水平衡模型的基础上 ,改进了原模型中蒸腾、蒸发量等项的计算方法 ;并根据黄土区降水分布特点 ,将全年分为 1 6个时间长度不等的降水时段 ,分别统计降水特征参数 ,利用质量守衡定律和逐步校正演算法 ,可计算出水文生态系统平衡体系水均衡要素的定量分配和活动层土壤含水量的年动态变化 ,代表了原模型中确定土壤含水量的简单统计法 ,从而可提高模型计算精度和实用性 .改进后的随机动力学水平衡模型更适用地黄土高原的特定水文生态系统
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在盆栽条件下研究了干湿交替对玉米生长速率、叶片水势、渗透势、气孔导度、相对生长速率和耗水量的影响。结果表明玉米在 3~ 7叶期经历土壤水分缓慢亏缺 ,再进行复水的干湿交替后玉米叶片渗透调节能力明显增加 ,叶片生长表现出补偿效应 ,每次干湿交替后生长速率迅速下降的叶水势趋于下降 ,气孔导度对土壤水分变化非常敏感 ,并在干旱—复水过程中具有后效作用 ,蒸腾耗水量随干—湿交替而具有下降趋势 ,初步证明可在节水灌溉条件下人为控制不同生育时期的供水时间形成干湿交替 ,促进渗透调节能力增强和补偿生长来实现作物高产、高效、优质的目的
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通过对不同水分处理下油菜不同叶位叶片光合速率(Pn)、蒸腾速率(E)和水分利用效率(WUE)的测定表明,油菜主茎叶片的光合速率、蒸腾速率和水分利用效率随叶位的降低而降低;干旱胁迫下油菜水分利用效率显著提高,光合速率变化不明显,但上部、下部叶间光合速率差异大。充分供水时叶片的蒸腾速率提高,水分利用效率则降低,从上到下叶片水分利用效率、光合速率降低不剧烈,而蒸腾速率表现为上部和下部叶高,而中部叶低。据测定,干旱处理下油菜单株生物量为37.45g,充分供水时为38.76g,干旱导致油菜生物量降低;水分胁迫能提高作物水分利用效率;在充分供水条件下叶片蒸腾速率较高,这可能是气孔行为的结果。