青藏高原东缘几种树苗对增强紫外线-B和氮供应的响应

大气臭氧的损耗导致了地球表面具有生物学效应的紫外线-B(UV-B)辐射增强。同时，大气成分变化中除了UV-B辐射增强外，氮沉降是一个新近出现而又令人担忧的环境问题，其来源和分布正在迅速扩展到全球范围，并不断向陆地和水生生态系统沉降。本试验在四川省境内的中国科学院茂县生态站内进行，以云山、冷杉、色木槭和红椋子幼苗为模式植物，从生长形态、光合作用、抗氧化能力和矿质营养等方面研究了青藏高原东缘4种树苗对全球变化－增强UV-B辐射和氮供应(氮沉降)的响应。该试验为室外盆栽试验，包括四个处理：(1)大气UV-B辐射+无额外的氮供应(C)；(2)大气UV-B辐射+额外的氮供应(N)；(3)增强UV-B辐射+无额外的氮供应(UV-B)；(4)增强UV-B辐射+额外的氮供应(UV-B+N)。其目的：一方面有助于丰富我国对全球变化及区域响应研究的全面认识，进一步完善在全球气候变化条件下臭氧层削减和氮沉降对陆地生态系统影响的内容；另一方面，在一定程度上有助于我们更好的理解在全球变化下森林更新的早期过程。具体结果如下： 增强的UV-B辐射在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有显著的影响。UV-B辐射增强对幼苗的影响不仅与物种有关，而且，还与氮营养水平相关。总体表现为，高的UV-B辐射导致了色木槭和红椋子幼苗叶片的皱缩和卷曲，并降低了色木槭幼苗的叶片数和叶重，在额外的氮供应下，云杉、冷杉和红椋子的叶重也显著地降低了；色木槭和红椋子幼苗叶片的解剖结构受到了增强的UV-B辐射的影响，增强的UV-B辐射显著地降低了色木槭叶片的栅栏组织厚度，提高了红椋子叶片的厚度；增强的UV-B辐射显著地降低了4种幼苗的单株总生物量、植物地下部分的生长、总叶绿素含量 [Chl (a + b)]、净光合速率和最大量子产量(Fv/Fm)，提高了4种幼苗叶片的膜脂过氧化(MDA含量)，改变了植物体不同器官中的矿质元素含量；增强的UV-B辐射提高了冷杉、色木槭和红椋子叶片中的过氧化氢含量(H2O2)、超氧负离子(O2-)生成速率，在额外的氮供应下，云杉叶片中的活性氧含量也显著地提高了；在无额外的氮供应条件下，增强的UV-B辐射显著地提高了4种幼苗叶片中的UV-B吸收物质、脯氨酸含量和抗氧化酶的活性(SOD、POD、CAT、GR和APX)。在额外的氮供应条件下，UV-B辐射的增强却显著地降低了冷杉叶片中脯氨酸含量和红椋子叶片中UV-B吸收物质含量，但是，在4种幼苗叶片中，5种抗氧化酶的活性对UV-B辐射的增强没有明显的规律性，增强的UV-B辐射显著地提高了云杉叶片中的POD、SOD和GR的活性，提高了冷杉叶片中的POD和GR活性，提高了色木槭叶片中的POD、SOD和CAT活性和红椋子幼苗叶片中的POD和SOD活性。从这些结果可知，植物在遭受高的UV-B辐射导致的过氧化胁迫时，植物体内形成了一定的保护机制，但是，这种保护不能抵抗高的UV-B辐射对植物的伤害。 额外的氮供应在生长形态、光合、抗氧化能力、活性氧和矿质营养方面对4种幼苗都有一定的影响，不同幼苗对额外的氮供应响应不同，并且受到UV-B辐射水平的影响。在当地现有的UV-B辐射水平下，额外的氮供应显著地提高了幼苗的单株总生物量、植物地下部分的生长、Chl (a + b)、净光合速率(红椋子除外)、UV-B吸收物质(冷杉除外)、脯氨酸含量(红椋子除外)和部分抗氧化酶的活性，降低了H2O2的含量、O2-的生成速率和MDA含量(红椋子除外)，改变了植物体内部分矿质元素含量，显著地提高了云杉和冷杉叶片中的Fv/Fm。这些指标总体表明，在当地现有大气UV-B辐射水平下，额外的氮供应对植物的生长和发育是有利的。在增强的UV-B辐射水平下，4种幼苗的生长形态和光合大部分指标都没有受到额外氮供应的影响，额外的氮供应提高了红椋子幼苗的单株总生物量和Chl (a + b)含量，提高了冷杉和色木槭叶片中的活性氧含量和MDA含量，却降低了红椋子叶片中的活性氧含量；额外的氮供应也提高了云杉、色木槭和红椋子叶片中UV-B吸收物质和脯氨酸含量，降低了冷杉叶片中UV-B吸收物质和脯氨酸含量；在抗氧化酶活性方面，额外的氮供应降低了云杉、冷杉叶片中5种抗氧化酶的活性和红椋子叶片中POD和GR的活性，提高了色木槭叶片中的POD和SOD的活性；4种幼苗植物体内的矿质元素含量对额外的氮供应没有显著的规律性。从这些结果可知，在高的UV-B辐射下，额外的氮供应提高了云杉、冷杉和色木槭幼苗对高的UV-B辐射的敏感性，然而，额外的氮供应却促进了红椋子幼苗的生长，原因可能是，在高的UV-B辐射下，额外的氮供应增加了红椋子叶片的厚度、叶重和叶片数，降低了叶片中活性氧含量的结果。表明在高的UV-B辐射水平下，额外的氮供应降低了红椋子幼苗对高的UV-B辐射的敏感性。 在全球变化的趋势下，UV-B辐射增强和氮沉降可能同时存在，我们的研究表明，与大气UV-B辐射+无额外的氮供应处理相比，增强UV-B辐射+额外的氮供应处理显著地降低了幼苗的单株总生物量(红椋子除外)、Chl (a + b)、净光合速率、Fv/Fm(冷杉除外)和MDA含量(红椋子除外)，提高了活性氧含量 (云杉除外)、UV-B紫外吸收物质含量(冷杉除外)、脯氨酸含量和部分抗氧化酶的活性，改变了植物体不同器官中的矿质元素含量。结果表明，在当地现有条件下，全球变化(UV-B辐射增强和氮沉降)对云杉、冷杉和色木槭幼苗的生长是不利，尽管植物体内一些抗氧化性指标提高了，然而，却对红椋子幼苗的单株总生物量的累积没有显著的影响。 The depletion of the ozone led to the increase of ultraviolet-B (UV-B) with biological effects in the earth’s surface. At the same time, except for enhanced UV-B radiation, nitrogen deposition was an anxious environmental problem at present, rapidly expanding to the global scope and continuously depositing to land and aquatic ecosystem. The experiment was conducted in Maoxian Ecological Station of Chinese Academy of Sciences, Sichuan province, China. Picea asperata, Abies faxoniana, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings were selected as model plants to assess the effects of enhanced UV-B radiation and supplemental nitrogen supply on growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings in east Qinghai-Tibetan Plateau. The experiment was potted outdoor, including 4 treatments: (1) ambient UV-B without supplemental nitrogen (control, C); (2) ambient UV-B with supplemental nitrogen (N); (3) enhanced UV-B without supplemental nitrogen (UV-B); (4) enhanced UV-B with supplemental nitrogen (UV-B+N). One hand, it was helpful for enriching our country to comprehensive understanding of the researches in the global change and the region response, further perfecting the effects of the depleted ozone layer and nitrogen deposition on land ecosystem under the global change; the other hand, it was favorable for us to better understanding of the early process of forest renews under the global change. The results were as follows: Enhanced UV-B radiation had significant effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits of 4 species seedlings. The effects of enhanced UV-B on plants were not only related with species, but also related with nitrogen nutrient level. Generally, the increase of UV-B radiation led to the shrinkage and curl of leaves in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, and reduced the number of leaf and leaf weight of Acer mono Maxim seedlings, under supplemental nitrogen supply, leaf weight of Picea asperata, Abies faxoniana and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings significantly also reduced; the anatomical features of leaf in Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings affected by enhanced UV-B radiation, the increase of UV-B radiation markedly reduced the palisade tissue thickness of Acer mono Maxim leaf and enhanced the leaf thickness of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings; the enhanced UV-B radiation significantly reduced total biomass per plant of 4 species seedlings, the growth of the underground parts, Chl (a + b), net photosynthetic rate and maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA content) and changed the content of mineral elements in different parts of plants; the enhanced UV-B radiation also increased the rate of superoxide radical (O2-) production and hydrogen peroxide (H2O2) content in leaves of Abies faxoniana, Acer mono Maxim, Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, under supplemental nitrogen supply, the reactive oxygen species in leaves of Picea asperata seedlings also significantly increased by enhanced UV-B radiation; under without supplemental nitrogen supply, enhanced UV-B radiation evidently induced an increase in UV-B absorbing compounds, proline content and the activities of antioxidant enzymes (SOD, POD, CAT, GR and APX) of leaves in 4 species seedlings. Under supplemental nitrogen supply, enhanced UV-B radiation induced a decrease in proline content of leaves in Abies faxoniana seedlings and UV-B absorbing compounds of leaves in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings, but, there were no obvious rules in the activities of five antioxidant enzymes of 4 species seedling leaves to enhanced UV-B radiation, enhanced UV-B radiation significantly increased the activities of POD, SOD and GR in Picea asperata leaves, the activities of POD and GR in Abies faxoniana leaves and the activities of POD, SOD and CAT in Acer mono Maxim leaves. The results indicated that some protective mechanism was formed when plants were exposed to enhanced UV-B radiation, but the protection could not counteract the harm of high UV-B radiation on plants. Supplemental nitrogen supply had some effects on 4 species seedlings in growth, morphological, photosynthesis, antioxidant and mineral nutrient traits. The response of 4 species seedlings was different to supplemental nitrogen supply, and was affected by UV-B levels. Under local ambient UV-B radiation, supplemental nitrogen supply significantly increased the total biomass per plant, the growth of underground parts, Chl (a + b), net photosynthetic rate (except for Acer mono Maxim seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and the activities of some antioxidant enzymes, and reduced H2O2 content, the rate of O2- production and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings) and changed the content of mineral elemental in different parts; supplemental nitrogen supply also evidently increased Fv/Fm in Picea asperata and Abies faxoniana seedlings. These results indicated that supplemental nitrogen supply was favorable for the growth of plants under local ambient UV-B radiation. Under enhanced UV-B radiation, mostly parameters in growth and morphology of 4 species seedlings were not affected by supplemental nitrogen supply. Supplemental nitrogen supply increased the total biomass per plant and Chl (a + b) of Swida hemsleyi (Schneid. et Wanger.) Sojak seedling, increased the reactive oxygen species and MDA content in Abies faxoniana and Acer mono Maxim leaves, and reduced the reactive oxygen species in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves; supplemental nitrogen supply also increased UV-B absorbing compounds and proline content in Picea asperata, Acer mono Maxim and Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, decreased UV-B absorbing compounds and proline content in Abies faxoniana leaves; in the activities of antioxidant enzymes, supplemental nitrogen supply significantly reduced the activities of antioxidant enzymes in Picea asperata and Abies faxoniana leaves and the activities of POD and GR in Swida hemsleyi (Schneid. et Wanger.) Sojak leaves, and increased the activities of POD and SOD in Acer mono Maxim leaves; the content of mineral elements in 4 species seedlings was no significantly rule to supplemental nitrogen supply. We knew from the results, under enhanced UV-B radiation, supplemental nitrogen supply made Picea asperata, Acer faxoniana and Acer mono Maxim seedlings more sensitivity to enhanced UV-B radiation, however, accelerated the growth of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings. The reason was probably that supplemental nitrogen supply increased the leaf thickness, leaf weight and leaf number, reduced the reactive oxygen content of leaf in Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings grown under high UV-B radiation. This showed that supplemental nitrogen supply reduced the sensitivity of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings to high UV-B radiation. Under the tendency of the global change, enhanced UV-B radiation and nitrogen deposition may probably coexist. The results showed, compared with the treatment of ambient UV-B radiation without supplemental nitrogen supply, the treatment of enhanced UV-B radiation with supplemental nitrogen supply significantly reduced the total biomass per plants (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), Chl (a + b), net photosynthetic rate, Fv/Fm and MDA content (except for Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings), and increased reactive oxygen content (except for Picea asperata seedlings), UV-B absorbing compounds (except for Abies faxoniana seedlings), proline content and part antioxidant enzymes, and changed the content of mineral elements of different parts. The results indicated that the global change (enhanced UV-B and nitrogen deposition) were not favorable for the growth of plants under local ambient UV-B radiation and nitrogen nutrient level,, though increased some antioxidant indexes, however, the treatment of enhanced UV-B with supplement nitrogen supply did not significantly affect on the biomass accumulation of Swida hemsleyi (Schneid. et Wanger.) Sojak seedlings.

川西亚高山两种针叶树幼苗对增温和施氮的响应

人类活动引起全球大气中温室气体（CO2、CH4、NOx）浓度不断增加，致使地球表面温度在过去的100 年中已经增长了0.74 ± 0.18℃，预计到本世纪末将会增加1.1-6.4℃。此外，氮沉降也是当今社会的重要环境问题，随着经济发展的全球化, 高氮沉降也呈现出全球化趋势。全球气候变暖和氮沉降给陆地生态系统的地上、地下生物学和生物地球化学过程所带来巨大影响越来越引起人们的关注。 本文以川西亚高山针叶林的两个重要树种云杉和油松幼苗为研究对象，采用红外辐射增温（空气增温2.1℃，土壤增温2.6℃）和根部施氮（施氮量25 g N m-2yr-1）的方法，从生长形态、光合作用、抗氧化能力和矿质营养等方面研究这两种幼苗对气候变暖和氮沉降的响应。该实验为室外控制实验，包括四个处理：(1)不增温+不施氮（UU）；(2) 不增温+施氮（UF）；(3) 增温+不施氮（WU）；(4) 增温+施氮（WF）。本研究旨在从生理生化、物质代谢 、生长及形态等不同水平上研究模拟增温和施氮对两种树苗的联合效应，提高我们对全球变化下亚高山针叶林早期更新过程的理解，同时也为森林管理提供科学依据。具体研究结果如下： 单独增温处理显著提高了云杉和油松幼苗的地茎、叶重、茎重、根重以及总生物量；单独施氮处理也增加了两种幼苗的株高和总生物量。而增温和施氮联合作用对两种幼苗生长的影响并不相同，联合作用对云杉幼苗生长指标的正效应显著低于单独施氮处理，但是联合作用比单独增温或施氮更大程度的促进了油松幼苗生物量的积累。 单独增温和施氮都有利于提高云杉和油松叶片中叶绿素含量、净光合速率(A)、最大净光合速率(Amax)、表观量子效率（Φ）、最大光能转化效率（Fv/Fm）和量子产量（Y）。与对两种幼苗生长指标的影响相似，加氮和增温共同作用下油松幼苗的以上光合指标比在单独增温或施氮处理下有更大程度的提高；而联合作用下云杉幼苗叶绿素含量、净光合速率、最大净光合速率、表观量子效率、最大光能转化效率以及量子产量比单独施氮处理明显地降低。 增温和施氮都显著地降低了云杉和油松幼苗针叶组织中活性氧和丙二醛的积累。交互作用降低了云杉幼苗叶片的抗氧化酶活性、脯氨酸和ASA 的含量，却显著提高了油松幼苗SOD、POD、APX 等抗氧化酶的活性，并且对油松幼苗脯氨酸和ASA 积累的促进作用比单一因子更加明显。因此，增温和施氮共同作用下油松幼苗叶片中O2-产生速率、H2O2 及MDA 含量明显降低，而云杉叶片中只有O2-产生速率出现降低趋势。 增温和施氮都降低了云杉体内的P、Ca、Mg 元素的含量，增加了Cu、Zn、Mn 在各器官内的积累。对油松幼苗而言，增温和加氮单独作用也显著降低了Ca 含量增加了Cu、Zn、Mn 的积累，但是不同于云杉幼苗的是P、Mg 也显著增加。增温和施氮联合作用对云杉幼苗体内元素的影响与单一施氮处理或增温处理相似，不同的是比单一因子作用更为明显降低了P、Ca、Mg 含量，增加了植株中N、Cu、Zn、Mn 的含量，但是油松矿质元素含量在联合作用下并没有产生类似于云杉幼苗的双因子叠加效应。 总之，尽管单独增温或者施氮都有利于云杉和油松幼苗生长指标、光合能力以及抗氧化能力的提高。但是，增温和施氮对云杉幼苗生长生理的促进效应非但没有在交互作用下有更大的提高，反而低于单独氮处理。与此不同的是，增温和施氮联合作用比单因子作用更有利于油松幼苗生长及生理指标的提高。 With the continued increase in atmospheric concentrations of greenhouse gases (CO2、CH4、NOx), the mean global surface temperature has increased by about 0.74 ± 0.18℃ over the past century and is predicted to rise by as much as 6.4℃ during this century. Besides global warming, nitrogen deposition is another serious environmental problem caused by human activities, and high nitrogen load has become globalization as a result of global economy development. Global climate warming and nitrogen deposition have induced dramatic alternations in above - and below- ground biology and biogeochemistry process in terrestrial ecosystems, and more and more attention has been invited to those problems. This experiment mainly studies two important species Picea asperata and Pinus tabulaeformis in subalpine coniferous forest of western Sichuan, China. Infared heaters are induced to increase both air and soil temperature by 2.1℃ and 2.6 ℃, respectively. Ammonium nitrate solution (for a total equivalent to 25 g N m-2 year-1) is added to soil surface. There are four treatments in this study: (1) unwarmed unfertilized (UU); (2) unwarmed fertilized (UF); (3) warmed unfertilized (WU); (4) warmed fertilized (WF). This study is conducted to determine the influences of experimental warming and nitrogen fertilization on physiolchemistry, nutrition metabolism, growth and morphology in the two coniferous species seedlings. The current study is favorable for increasing our understanding on the early phase of regeneration behavior in subalpine coniferous forest, and it also provide scientific direction for forest management under future global changes. The results are as follows: Artificial warming alone significantly increased basal diameter, leaf mass, stem mass, root mass and total biomass for Picea asperata and Pinus tabulaeformis seedlings, and single nitrogen fertilization are also favorable for growth of the two species and stimulate plant hight and total biomass. The two species seedlings respond differently to the combination of elevated temperature and nitrogen addition. Warming combined with nitrogen fertilization weakens the positive effects of nitrogen addition for growth of Picea asperata seedlings. However, the combination of elevated temperature and nitrogen fertilization further increase biomass accumulation of Pinus tabulaeformis seedlings. Both elevated temperature alone and nitrogen fertilization alone can increase photosynthetic pigments contents, net photosynthetic rate (A), maximum net photosynthetic rate (Amax), apparent quantity yield (Φ), maximum photochemical efficiency of photosystem II (Fv/Fm) and effective quantum yield (Y). Similarly with growth parameters, the combination of warming and nitrogen addition induced more increment of these above photosynthetic parameters for Pinus tabulaeformis seedlings. However, these photosynthetic parameters of Picea asperata seedlings under the combination of warming and nitrogen addition are lower than those under nitrogen fertilization alone. The levels of active oxygen species (AOS) and malodiadehyde (MDA) in needles of the two coniferous species seedling are obviously decreased by experimental warming or additional nitrogen. Warming combined with nitrogen fertilizer reduces the activities of SOD, CAT and APX, and the contents of proline and ASA of Picea asperata seedlings, but the combination significantly increases activities of these antioxidant enzymes in needlels of Pinus tabulaeformis seedlings and further improves the accumulation of proline and ASA compared to either artificial warming or nitrogen addition. Therefore, the rate of O2 - production, the contents of H2O2 and MDA in needles of Pinus tabulaeformis seedlings are remarkably reduced by the combination of warming and nitrogen addition, but the combination only significantly decreased the rate of O2 - production of Picea asperata seedlings. Elevated temperature or nitrogen fertilization decrease the contents of P, Ca, Mg but increase Cu, Zn, Mn contents for Picea asperata seedlings. For Pinus tabulaeformis seedlings, elevated temperature alone and nitrogen fertilization alone decreased Ca, but increased P, Mg, Cu, Zn, Mn contents. The effects of the combination of warming and nitrogen addition on these element contents in needles of Picea asperata seedlings are added or multiplied the effects of warming and nitrogen addition alone, resulting in less contens of P, Ca, Mg and more contents of Cu, Zn, Mn than either elevated temperature or nitrogen fertilization. Howere, these adding or multipluing single-factor effects on contents of these elements are not observed in the case of Pinus tabulaeformis seedlings. In conclusion, growth parameters, photosynthetic capacities and antioxidant abilities of Picea tasperata and Pinus abulaeformis seedlings are improved by experimental warming or nitrogen fertilization. Interestingly, the positive effects of warming and nitrogen addition on growth and physiological performances are not multiplied by the combination of elevated temperature and nitrogen fertilization, even dempened for Picea asperata seedlings. However, for Pinus tabulaeformis seedlings, growth and physiological performances are further improved by the combination.

Field studies on the environmental factors in controlling microcystin production in the subtropical shallow lakes of the Yangtze River

Microcystin (MC) problem made more and more care about in China, intercellular MC (Int-MC) and cellular MC (Cel-MC) were important contents to reflect the producing-MC ability by cyanobacteria and by lakes. To study the correlations between Int-MC, Cel-MC concentration and biological and environmental factors, eight cyanobacterial blooming lakes were studied in the middle and lower reaches of the Yangtze River. Microcystin-RR (MC-RR) and Microcystin-LR (MC-LR) were the primary toxin variants in our data. From the linear correlations between MC and environmental factors, cellular-YR had significant correlation with most of chemical factors except total nitrogen (TN) and the ratio of total nitrogen and total phosphorus (TN/TP), most intracellular MC analogues had significant correlations with total dissolved nitrogen (TDN), ammonium (NH4+), nitrite (NO2-), TP, total dissolved phosphorus (TDP), Microcystis. From the canonal correspondence analysis, Int-MC concentrations were closely related with the chemical and biological factors, such as TP, total organic carbon (TOC), chlorophyll a (Chl a), Microcystis biomass, et al. While Cel-MC contents, especially Cel-RR and Cel-LR, were closely related with light environmental in the lakes such as water depth and transparence.