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.

遮荫与外源脱落酸喷施对粗枝云杉(Picea asperata Mast.)不同种群抗旱性的影响

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.

川西亚高山森林生态系统辐射传输研究

辐射传输研究是贯穿森林生态系统的纽带，太阳辐射为植物的生长发育提供光合能量、适宜的环境温度以及发育信息。一方面，气候变化使到达地面辐射能的质和量发生变化，影响到植被的生长发育，改变森林的结构，而森林结构的变化又会影响林冠内辐射能的分配和质量，这些变化会进一步影响到林下土壤温度，改变森林根系活性以及土壤营养转化的效率；连锁反应的结果有可能会使森林生态系统的生产力发生变化，改变碳素和氮素源库的调节方向，从而反馈影响地球气候系统。另一方面，人类作为生态系统的成员，必然需要森林生态系统为其提供更多的原材料和更好的生态服务功能，如何实现这些目标，就需要人类适度调整干预方式和频度，达到预期的目的。本文在建立适合于川西亚高山森林的叶面积测量技术、光照辐射模型和土壤温度变化模型的基础上，对川西亚高山地带森林生态系统的辐射传输特征进行了分析，并从森林结构的角度探讨了林分内的辐射分布以及对土壤温度的影响。主要成果如下： 1. 提出了一种照相法测量叶面积的方法。通过对摆放在平面上的叶片照相，利用投影变化，把非正射图像转化为正射图像，然后经过计算机图像处理得到每一片叶片的面积、周长、长度、宽度等信息。这种方法可使用户以任意方向和距离拍摄处于平面上的叶片，能同时处理大量的叶片，适于野外离体或活体叶片测量。叶片面积分辨率可调，分辨率可以与常用的激光叶面积仪相近甚至更高，而且叶片图像可以存档查询。 2. 提出一种模拟林内光照变化的模型。利用林冠半球照片，记录视点以上半球内的林冠构件空间分布，作为林冠子模型；天空辐射子模型采用国际照明委员会(CIE)的标准晴天和阴天以及插值模型。该模型能够模拟林下某一位点处的实时光斑变化。 3. 提出一种土壤温度变化模型。把土壤视为具有容量和阻力性质的结构，利用电阻和电容器件构建土壤能量分布模型。外界太阳辐射能经过植被以及其它一些能量分配器后进入土壤，其中有一部分转化为土壤势能，即土壤温度。土壤温度的变化类似于电池的充放电过程。在已知模型参数的情况下，可以从太阳辐射计算土壤温度的变化。在模型参数未知的情况下，通过输入和输出值推算模型的参数，而模型参数中的时间常数与土壤组成和含水量有关，这样就可以知道土壤水分的变化情况。 4. 从王朗亚高山森林典型样地林分结构的测量获得林地三维结构图、树冠形态、叶面积密度等参数，这些参数输入到Brunner (1998)开发的tRAYci 模型中计算出一段时间内林分任意位置处的光照值。与林下辐射计测量值以及半球照片计算结果的比较，该模型基本上能够满足对林分光环境了解的要求。 5. 从川西亚高山森林生产力的角度，探讨了森林生产力研究的方法以及川西地区的研究历史和成果，发现了其中的一些规律和问题，特别是在叶面积测量上，还没有使用标准的叶面积指数定义。综合来看，川西地区针叶林叶面积指数(单位土地面积上植物冠层总叶面积的一半) 应在4-5 之间。降雨丰富的华西雨屏带是川西地区森林生产力最高的地区，而向西北森林生产力逐渐降低。川西地区云冷杉林森林生产力平均约为600 gDM m-2 a-1，但是根据辐射能计算的潜在生产力则达到1800 gDM m-2 a-1。实际与潜在森林生产力的巨大差异说明其它因子对生产力的影响。 6. 王朗亚高山3 个典型森林林分中，白桦林样地(BF) 林下草本以糙野青茅、牛至、紫菀等喜阳性物种为主，林下透光度较高；冷杉林样地(FF) 林下透光度最低，以喜阴性物种水金凤、蟹甲草、囊瓣芹等为主；而云杉林样地(SF)林分林龄最大，林下透光度介于冷杉林和白桦林之间，草本层仍然以喜阴性物种东方草莓、紫花碎米芥、酢浆草等为主。冷杉林和云杉林的灌木层也很丰富，卫矛属、五加属、茶藨子属、忍冬属植物很丰富，而在白桦林则以栒摘要子属、榛子属、鹅耳枥属等植物为主。藓类植物在云杉林中最丰富，并且形成毯状层，其它两个林分则很稀少。3 个样地林分结构与林下光环境有很强的相关性，从光环境特征可以在一定程度上推测林分的结构。各样地单纯从乔木层材积推算的NPP 排列顺序为BF>FF>SF，与林下辐射透射率和林分年龄的顺序相同，暗示辐射对群落演替过程的驱动作用。 7. 用半球照相法测得BF、FF 和SF 3 个样地的有效叶面积指数以SF 样地最高，BF 最低。如果考虑针叶树叶片在小枝上的丛聚分布，利用北方针叶林的数值进行校正，则SF 样地LAI 显著增加(达到89%)，其它样地的LAI 基本不变甚至有所下降。校正后的数值与文献中地面测量的结果较相近，说明在使用半球照相法测量川西亚高山针叶林LAI 时必须加以校正。 8. 在3 个样地中，白桦、岷江冷杉和方枝柏种群为丛聚分布，紫果云杉在FF和SF 样地中基本上为随机分布。3 个物种出现丛聚分布的最短距离约为2m，在最短距离以内则为随机分布。最短距离可能与树冠大小有关，种子传播特征以及对光照的需求状况可能是造成这种分布格局类型的原因。 Radiative transfer plays a key role in forest ecosystems. Solar radiation providesenergy for photosynthesis, appropriate ambient temperature and development informationfor plants. However, quality and quantity of radiation reaching land surface are affected byweather and subsequently influence the growth and development of plants, which in turnchanges the budget of radiation in forest. Soil temperature changes with the variation ofradiation under forest canopy and influences the activity of roots and rate of nutrientturnover. Thus, any changes of radiation will induce chain reactions in the entireecosystem and display in the value of net primary productivity which will possibly shiftthe relationship between carbon source and sink at local or regional scale and feed back tothe global climate system. On the other hand, as a component of ecosystems, humanbeings of course need to demand more materials and better service from ecosystems. Forthese purpose, man must adapt their pattern and frequency of interference to ecosystems.This paper aims to research on the canopy structure, the radiation distribution and theirinfluence on soil temperature from the process of radiative transfer in subalpine forestecosystem of western Sichuan. The main results are: 1 Present a new photogrammetric method for leaf area. The main idea is to convertnon-vertically taken images of planar leaves to orthoimages through projectivetransformation. The resultant images are used to get leaf morphological parametersthrough image processing. This method enables users to take photos at almost anyorientation and distance if only the leaves are placed on same plane, and to processlarge quantity of leaves in a short time, which is suitable for field measurement. Theresolution of leaf area is adjustable to fit for special requirement. 2 A model using hemispherical photos combining with solar tracks and radiation courseis provided to simulate light variation in forest. The hemispherical photos of canopyrecord the real spatial distribution of each element of plants viewed from a point. Skyradiance is simulated with CIE standard clear sky or cloudy sky model. This modelcan be used to simulate real time light variation under canopy. 3 Present a soil temperature model. Soil could be regarded as a body of resistor andcapacitor. Some of the budget of solar radiation in soil body is transformed into soilpotential energy, the soil temperature. Variation of soil temperature is driven by solarradiation, vegetation, soil properties, etc. This model has two parameters, one of whichis time constant and is related to soil water content. The inversed model can be used tosimulate the variation of soil water. 4 By using model tRAYci developed by Brunner (1998), the 3-D distribution of light inthree subalpine forest stands of Wanglang Nature Reserve has been simulated andvalidated with value of radiometers in these stands. This model can basically satisfythe need for understanding light regimes of these stands. 5 Present some principles and questions of NPP (net primary of productivity) researchesin western Sichuan. The standard leaf area index (LAI) defined by Chen and Black(1997) has not been used in this region. Total leaf area and projected leaf area indexare still used in NPP researches which may differ around 1-fold in magnitude. Thestandard LAI which is a half of total leaf area above unit land area should be between4 and 5 for typical subalpine coniferous forest of western Sichuan concluded fromliteratures. The maximum forest NPP occurs in West China rain belt and decreasesnorthwestwards. Average NPP of spruce-fir forest in western Sichuan is about600gDM m-2 a-1, which is below the potential NPP of 1800gDM m-2 a-1 based onmeasured radiation in this region. The significant difference between potential and realNPP suggests that other factors influence the growth of stands. 6 In the three subalpine forest stands of Wanglang Nature Reserve, herbage layer ofAbstractbirch stand (BF) with age of 40 is dominated by heliophytes of Deyeuxia scabrescens,Origanum vulgare, Aster tongoloa etc.. However, both of the other two stands aredominated by shade tolerent species, such as Impatiens noli-tangere, Impatiensdicentra, Cacalia deltophylla and Pternopetalum tanakae etc. in fir stand (FF) withage of 180 and Fragaria orientalis, Cardamine tangutorum and Oxalis corniculata etc.in spruce stand (SF) with age of 330. Shrub species in the latter two stands arerelatively rich, typical dominant genera being Euonymus, Acanthopanax, Ribes andLonicera. Birch stand has relatively sparse shrubs dominated by genera of Cotoneaster,Corylus and Carpinus. Mosses are significant only in spruce stand. The canopystructure controls the light regime of stand, which influence the composition of herblayers beneath the canopy. This light regime-community structure relationship can beused to infer the herb community from canopy structure. The NPP derived from timbervolume of arbor layer of the three stands decreases from BF to SF, which is in thesame order of transmitted total radiation under canopy and age of these stands,suggesting the driving effect of radiation in the succession of community. 7 The highest effective LAI of the three stands obtained by hemispherical photos is inplot SF and lowest in plot BF. After rectification of the clumping effect of leaves onshoot, the real LAI in plot SF increases significantly (89%) and approximate to theaverage LAI of coniferous forest in western Sichuan. Therefore, the LAI obtainedfrom hemispherical photos needs rectification for clumping effect. 8 Spatial distribution pattern for Betula platyphylla, Abies faxoniana and Sabinasaltuaria is clumpy, but Picea purpurea almost random in plot FF and SF. The shortestdistance for clumpy distribution for Betula platyphylla and Sabina saltuaria is 1.5m,and 2m for Abies faxoniana. And random pattern for these trees is exhibited within thisrange which almost coincides with the diameter of crown. Seed dispersalcharacteristics and light requirement may be the reason for different spatial pattern.

岷江干旱河谷三种蔷薇植物繁殖更新研究

繁殖更新是植物生活史的重要阶段，在退化生态系统中，植物繁殖更新能力往往较差，是植被恢复的限制环节，因而也成为恢复研究重点和核心。本研究选择岷江干旱河谷广泛分布的三种蔷薇：多苞蔷薇（R. multibracteata）、黄蔷薇（R. hugonis）和川滇蔷薇（R. soulieana）为研究对象，通过野外调查，在查明其生长、繁殖更新状况的基础上，采用控制和模拟实验，对种子和幼苗阶段进行了深入研究，综合分析更新潜力，并提出相对应的促进更新和植被恢复措施。主要结论如下： 1）三种蔷薇在岷江干旱河谷广泛分布，生长和繁殖状况良好，结实量大。各生长指标：株高、基径和冠幅，繁殖指标：结实数量、重量和单果重量都具有显著的空间差异性。基径对多苞蔷薇结实量影响最大；而冠幅对黄蔷薇结实量影响最大。海拔和纬度是对蔷薇生长和繁殖影响最大的环境因素，随着海拔和纬度的升高，植株生长更高大，结实量增加；坡度和坡向对其生长和繁殖也有一定影响，随着坡度 和坡向增加，蔷薇生长和结实受到抑制。 2）三种蔷薇在岷江干旱河谷更新现状不佳, 但更新潜力大。活力种子比率低，动物取食以及两年生幼苗的大量死亡是蔷薇更新的主要限制因素。多苞蔷薇和黄蔷薇的结实率低，川滇蔷薇较高。三种蔷薇种子产量大，但种子质量较差，更新具有充足的种源。三种蔷薇都能形成持久种子库，种子库中种子总量大，但有效种子少，黄蔷薇被动物啃食的比例很高，多苞蔷薇和川滇蔷薇也有一部分种子受到动物破坏。三种蔷薇幼苗库组成特征表现为，当年生幼苗所占比例很高，年龄较大幼苗所占比例小。 3）三种蔷薇都具有不同程度休眠，未经处理种子的发芽率极低。黄蔷薇休眠程度最深，为深度生理休眠；多苞蔷薇为中度生理休眠；川滇蔷薇为非深度生理休眠。三种蔷薇种子在形态上发育成熟，种皮具有透水性。蔷薇果果肉和瘦果中含有抑制物质，其浸泡液抑制了油菜种子萌发，果肉抑制作用更强，果肉和瘦果浸泡液的抑制程度分别为：川滇蔷薇>黄蔷薇>多苞蔷薇。切割和硫酸腐蚀提高了川滇蔷薇种子的发芽能力，而对多苞蔷薇和黄蔷薇没有影响。完全去除瘦果果皮和种皮提高了多苞蔷薇种子发芽率，但对黄蔷薇没有影响。赤霉素和烟水对蔷薇种子萌发没有促进作用。三种蔷薇打破休眠所需低温层积时间分别为：黄蔷薇>多苞蔷薇>川滇蔷薇。对于多苞蔷薇和川滇蔷薇，层积前对种子进行硫酸腐蚀或暖温层积能缩短低温层积时间，提高发芽率。对于多苞蔷薇，变温层积中暖温层积和低温层积具有一定的负补性，即延长暖温层积可以缩短种子萌发对低温层积的需要。 4）多苞蔷薇种子形态特征和种子休眠与萌发在不同海拔梯度间存在较大差异。种子采集时间、采集季节和干藏影响多苞蔷薇和川滇蔷薇的种子休眠。多苞蔷薇果实大小、种子大小和千粒重、种皮厚度随海拔升高而增加，而种子饱满率和活力随海拔升高而降低，种子休眠程度也随海拔升高而增加。种皮厚度与种子大小、千粒重成正相关关系，硫酸腐蚀后的种子经过不同时间的低温层积后，种子发芽率与种皮厚度、种子大小、千粒重、海拔成正相关关系。2006 年采集川滇蔷薇和多苞蔷薇种子休眠程度较2005 年低。种子休眠随种子年龄增加而减弱。高温和干旱能减轻多苞蔷薇和川滇蔷薇种子休眠。 5）三种蔷薇的生长和生物量积累在干旱胁迫条件下受到抑制，而生物量分配、叶片形态特征和水分利用特征等都发生了变化。三种蔷薇的根、茎、叶各器官生物量以及总生物量等在干旱胁迫下明显减小，叶片脱落数量增加。在干旱胁迫条件下，较多的生物量分配到地下部分，从而这使R/S 明显增加。比叶面积（SLA）和冠层面积比（LAR）对干旱胁迫的反应不敏感，仅有部分物种在干旱胁迫条件下发生了变化，并且其变化特点在不同年龄幼苗之间有一定差异。干旱胁迫对WUE 的影响在不同物种间存在差异。多苞蔷薇和黄蔷薇的WUE 随着干旱胁迫的增加而增大， 而川滇蔷薇的WUE 则随干旱胁迫增加而减小。在干旱胁迫条件下，多苞蔷薇和黄蔷薇叶片脱落量和生物量减小幅度较川滇蔷薇大，表明其抗旱能力较强。在干旱胁迫条件，三种蔷薇两年生幼苗的生物量减小幅度较当年生幼苗小，表明两年生幼苗的抗旱能力更强。 6）两种植被恢复措施中，幼苗移栽比播种具有更好的植被恢复效果。播种后，蔷薇种子的发芽率较高，但出苗率都很低，即使出苗，幼苗也几乎在一月内全部死亡。 三种微生境条件下（灌木、半灌木和裸地），种子出苗和幼苗成活没有差异。移栽幼苗总体死亡率都比较低，小于20％。特别是两年生幼苗死亡率更低，小于2％。移栽后的幼苗生长状况良好，在整个生长季中，各生长指标不断增加。生境对幼苗的存活率没有显著影响，但对于幼苗的生长和生物量积累有一定影响，裸地更有利于幼苗生长和生物量积累。与当年生幼苗相比，两年生幼苗具有更高的成活率。总之，三种蔷薇在干旱河谷分布广泛、生长繁殖状况良好，结实量大，具有丰富种源，繁殖更新潜力大，但繁殖更新状况不佳；种子散布后动物对种子的取食、种子的深度休眠过程、种子出苗以及当年生幼苗的存活和定居是更新的主要限制环节。水分是影响结实、种子休眠解除和萌发，幼苗存活和定居的最主要的限制因素。在植被恢复中，应在种子成熟季节大量采集种子，在室内打破休眠后进行人工播种，培育两年生幼苗，通过幼苗移栽方式进行植被恢复。川滇蔷薇应栽种在相对湿润的过渡区，而多苞蔷薇和黄蔷薇可以应用于核心区植被恢复。 Regeneration is an important phase in plant life cycle. It has been a key component of ecological restoration in degradation ecosystem in which plants commonly has poor regeneration. In this paper, we investigated the natural growth, propagation and regeneration status of native three rose species, Rosa multibracteata, R. hugonis and R. soulieana, and analysis the limitation in seed germination and seedling establishment stages. Advice on facilitating the use of these plants in restoration based on the results has been proposed. The results were as follows: 1) Three rose plants widely distributed in the dry valley of the Minjiang River, and made a good performance in growth and propagation. There were significant spatial differences in each growth parameter, such as ramet height, basal diameter, crown diameter and propagation parameters including hip number of a clump, hip mass of a clump and a hip mass. Basa diameter was the most important growth parameter influencing fruit number for R. multibracteata and crown diameter was for R. hugoni. Altitude and latitude had the greatest effect on the growth and propagation of rose plants among environmental conditions. Each parameter of growth and propagation increased with the increase of altitude and latitude. In addition, the increase of slope and aspect limited the growth and propagation. 2) Three rose plants had poor natural regeneration, but great regeneration potential. Low seed viability, predation and higher mortality of current year old seedlings were the limitation in regeneration. R. multibracteata and R. hugonis had higher fruiting rates than R. souliean. All three plants produced a great number of seeds, while their viability was poor. Three rose plants had persistent seed banks, with high total seed number but very low viable seed density. Predation was most severe in R. hugonis, and it also existed to some degree in R. multibracteata and R. soulieana. The seedling age-structure was characteristic of current-year seedlings predominating and few older seedlings were observed. 3) Three rose seeds were dormant and untreated seeds germinated with very low germination percentages. The rose seeds had morphological mature embryos, and achenes were permeable. Some inhabit substances existed in hips and achenes for the extracts of hips and achenes inhibited germination of Brassica campestris. The inhibition effect of the extracts of three rose hip and achenes was R. soulieana>R. hugonis>R. multibracteata. Mechanical and H2SO4 scarification increased R. soulieana germination but had no effect on germination of R. hugonis and R. multibracteata seeds. Full removal of pericarp and testa improved the germination of R. multibracteata but did not affect R. hugonis germination. GA3 and smoke water had no positive effect on rose seed germination. The periods of cold stratification required to released seed dormancy was R. hugonis > R. soulieana >R. multibracteata. H2SO4 scarification and warm stratification shortened cold stratification to release dormancy for R. soulieana and R. multibracteata. Warm stratification had complementary effect for cold stratification, i.e. the longer warm stratification seeds received, the shorter cold stratification were required to obtain the same germination percentage. Three rose seeds had different kinds of dormancy; R. hugonis has deep physiological dormancy, R. multibracteata with intermediate physiological dormancy and R. souliean non-deep physiological dormancy. 4）The seeds traits and dormancy of R. multibracteata showed significant difference across altitudes. Year and season of seed collection had significant effect on seed dormancy for both R. souliean and R. multibracteata. Hip size, seed size, seed weight, seed coat thickness and seed dormancy level increased with the increase of the altitude. There were positive relations between seed coat thickness with seed size and seed weight. Germination percentage of seeds treated with H2SO4 scarification following different periods of cold stratification showed positive relation with seed coat thickness, seed size, seed weight and altitude. Seeds of R. souliean and R. multibracteata collected in 2006 had low dormancy level than those collected in 2005. Seed dormancy decreased with increasing seeds age. High temperature and drought were associated with low dormancy level. 5) Seedling growth, the total dry mass and their components of seedlings were reduced, while leaf senescence accelerated under drought stress. More biomass allocation to root system resulted in higher R/S ratio under drought. Water-use efficiency (WUE) of R. multibracteata and R. hugonis increased, while it declined for R. soulieana under drought stress. R. soulieana seedlings had poor drought-resistance capacity it had more senescent leaves, and its reduction of biomass was stronger than two other rose plants under drought. The reduction degree of one year old seedlings under drought stress was slighter than that of current year seedlings. Therefore, one year old seedling was more drought-resistent compared to current year seedlings. 6）Planting seedlings may have better effect in comparison with direct seeding. Most seeds germinated after seeding, but seedling emergence was very low. More than 80 % seedlings from direct seeding died within a months after emergence. Seedling emergence and survival rate did not show difference among microhabitats. Mortality rates of seedlings artificially planted in microhabitats were general lower than 20 %, and the mortality rate of one year old seedlings was lower than 2 %. Each grow parameter including plant height, leaf number and branch number continually increased after planting. Microhabitat type had effect on the growth parameter and biomass production, but it did not influence the seedling survival. Bare land tended to facilitate seedling growth. One year old seedlings had higher survival rate than current year seedlings. In conculsion, the three rose had wide distribution in the dry valley of the Minjiang River. They produced many seeds and had tolerance to drought stress to some degree. But they had poor regeneration in habitats may be caused by predation, seed dormancy，and high mortality in current year seedlings. We recommend that rose plants should be utilized in restoration by planting two-year old seedlings in spring. A large quantity of seeds should be collected artificially in autumn, release seed dormancy in room, and then cultivate two-year old seedlings by seeding in particular container. R. soulieana seedling probably be planted in transition area, and R. multibracteata and R. hugonis can be used in core area of the dry valley of the Minjiang River.