983 resultados para 237
Resumo:
从新几内亚核桃木的树皮中分离得到的吲哚类喹诺里西定生物碱10-Desbromoarborescidine A,因发现其具有阻滞钙离子通道的活性而倍受关注。10-Desbromoarborescidine A由A、B、C、D四个环组成,只有一个手性中心,是吲哚生物碱中结构较简单的一种,常作为此类生物碱全合成方法的模型化合物。但迄今为止,能高效而简便的实现手性10-Desbromoarborescidine A不对称全合成方法线路不多,大多数以不对称诱导的方式建立其手性中心,手性催化的方式仅有一例金属催化。从逆合成分析可知,Desbromoarborescidine A的全合成可以通过亚胺不对称催化还原进行关键的手性中心构建,而本课题组在之前的研究中通过手性有机小分子催化剂的发展,已将三氯硅烷氢转移还原亚胺发展成了一类简便实用、高效、高对映选择性并具有优良底物适应范围的不对称催化反应,我们希望以这一反应作为关键手段,发展一条Desbromoarborescidine A及其类似物不对称合成新路线。 根据我们设计的新路线,首先成功合成了其关键中间体,然后我们进行了关键的不对称催化尝试。用本实验室已有的高性能有机小分子催化剂虽得到了较好的对应选择性,但是产率很低。同时,为了验证整条线路的可行性,我们也用消旋的中间体进行拉通线路的尝试。但不幸的是,在脱除保护基时遇到了很大困难。尝试换不同的保护基,或改变脱保护基的顺序,都未能成功合成目标产物。究其原因可能是由于吲哚的特殊性造成的,吲哚类亚胺与常规的芳香亚胺有较大的差异,其NH基团无论保护还是不保护,对与其2位相联接的C=N双键均有很大的影响,导致其不对称催化还原难以进行。另外,由于所设计的还原产物含有处在吲哚苄位的胺基,稳定性较差,造成保护基脱除困难。 烯胺C-亚磺酰化反应是本课题组最近发现的一个新反应,之前未见文献报道。本研究对该反应进行了反应条件优化和底物扩展,发现带Cbz,Ac,COt-Bu,CO2Et,Bz等保护基的一系列环状和非环状烯胺在亚磺酸钠、DMAc和MeSiCl3的共同作用下能高效高产率生成β-胺基烯基亚砜类新化合物,为合成多官能团化的烯基亚砜新化合物提供了一条简便实用的途径。 The main constituent of Dracontomelum mangiferum B1, indoloquinolizidine alkaloid 10-Desbromoarborescidine A, has drawn great attention due to its calcium channel blocking activity. Its molecular structure is relatively simple compared with the other alkaloids of the same type, which has only one chiral center, albeit with four cycles A, B, C, and D. This compound is often used as a model target for exploring different strategies for the total synthesis of indole alkaloids. Nevertheless, so far there still lack practical and highly efficient methods for the asymmetric total synthesis of 10-Desbromoarborescidine A. Most of the current available methods rely on stoichiometric asymmetric synthesis for the construction of the chiral center. There is only one example reporting utilization of asymmetric catalysis, but with transition metal complex as the catalyst. Our retrosynthetic analysis shows that catalytic asymmetric reduction of imine could be used as the key step for the construction of the chiral center of Desbromoarborescidine A. Since in the previous studies our group has developed the asymmetric reduction of imines by trichlorosilane into a practical and highly efficient and enantioselective method using newly designed chiral organocatalysts, we hope to apply this method to develop a novel synthetic route for the total synthesis of Desbromoarborescidine A and its analogues in this study. According to the newly designed synthetic route, we first accomplished the synthesis of the key intermediates which was then examined for the critical asymmetric catalysis. The asymmetric reduction using the highly efficient organocatalysts, developed in our lab afforded high ee but poor yield. We tried different reaction conditions to improve the yield, but failed to get any good results. Simultaneously, to vertify the feasibility of the synthetic route we designed, we also tired to go through the route toward the racemic synthesis of Desbromoarborescidine A. But unfortunately, protection and deprotection proved to be big hurdles. All the different protection groups and different sequences of protection and deprotection we tried failed to get us through the designed synthetic sequence and furnish the final product. Most likely, the indole part is the culprit behind the failures.The NH group of the indole, no matter protected or not, may impact the catalytic asymmetric reduction of C-N double bond connected with 2-C. Additionally, the reduction product we designed contains an amino group in the β-position of the indole, which may cause problems due to its instability. C-sulfenylation of enamines is a novel reaction discovered recently by our group, which has not been seen before in the literature. In this study, optimization of the reaction conditions and exploration of the substrate scope were further undertaken for this reaction, which reveal that a series of enamines with N-Cbz, Ac, COt-Bu, CO2Et protection groups could all undergo smooth C-sulfinylations with the comined use of sodium benzene sulphinate, DAMc and MeSiCl3, efficiently furnishing the β-amino vinylsulfoxide products in high yield, affording a practical and highly efficient methods for synthesis of functional vinylsulfoxides.
Resumo:
近年来,随着对作物重茬障碍原因的深入研究,植物的化感作用越来越受到国内外众多学者的重视。花椒(Zanthoxy piperitum.)为芸香科植物,是一种收益早、用途广、价值高的经济树种,是川西干旱河谷地区的重要经济作物,其连作障碍也倍受关注,系统研究花椒化感作用将有助于理解和最终解决花椒连作障碍问题。本文首先通过萃取、层析等方法分离花椒主效化感成分;通过外加不同浓度的花椒叶水浸液研究了对土壤氮素养分循环的影响;研究了花椒叶水浸液对苜蓿生理生化、光合作用、氮素养分吸收的影响,并对外施氮肥对这种化感影响的缓解作用做了研究;研究了花椒化感潜力对全球变化——UV-B增强辐射的响应。主要研究结果如下: 1.用不同极性的有机溶剂对花椒叶水浸液浓缩浸膏萃取、柱层析,结合生物活性检测,分离得到主效化感作用组分的一种化感物质——对甲氧基苯酚。采用该物质纯品进行生物活性检测,证明其具有化感作用。 2.花椒叶水浸液处理土壤30天后,土壤硝态氮、铵态氮、无机氮(硝态氮+铵态氮)与对照相比,随着花椒叶水浸液浓度的增加呈现降低的趋势,其中土壤铵态氮含量显著降低,而硝态氮含量的变化则不显著,无机氮含量也显著降低。土壤脲酶和蛋白酶的活性与无机氮含量的变化趋势相同。随着花椒叶水浸液浓度的增加,氨化细菌数量显著降低,固氮菌的数量变化不显著,硝化细菌和反硝化细菌数量有减少的趋势。60天后,硝态氮含量、铵态氮含量、无机氮随水浸液浓度增加的变化趋势与30天时相似;随着花椒叶水浸液浓度的增加,氨化细菌、固氮菌的数量显著减少,硝化细菌数量、反硝化细菌数量仍呈减少趋势;土壤脲酶、蛋白酶活性与第30天的变化趋势相同。第60天与第30天的结果相比,相同水浸液浓度处理的硝态氮、铵态氮、无机氮均有下降的趋势,但除了25g.L-1水浸液处理的外,其它相同浓度的处理间差异均不显著;除了12.5 g.L的处理外土壤脲酶活性均呈增强的趋势;蛋白酶活性都有不同程度的增加;花椒叶水浸液处理的土壤硝化细菌和反硝化细菌数量呈增加趋势。 3.随着花椒叶水浸液浓度的增加,显著抑制了苜蓿根长、地上地下生物量、叶绿素含量、叶片中可溶性蛋白的含量,净光合速率。苜蓿体内四种抗氧化酶(POD、SOD、CAT、APX) 活性随着水浸液浓度的增加而降低,而丙二醛含量则增加。苜蓿氮初级同化相关酶硝酸还原酶(NR)、谷氨酰合成酶(GS)、谷氨酸脱氢酶(GDH)的活性随着水浸液浓度的增加受到不同程度的影响。总的来说,苜蓿硝酸还原酶、谷氨酰合成酶的活性受到抑制,而谷氨酸脱氢酶活性的变化则比较复杂,根呈先降低后增加的趋势,叶片则无显著变化。外施两种不同浓度的硝酸铵氮肥后,对12.5、25 g.L-1花椒叶水浸液处理的苜蓿化感作用有显著的缓解作用,表现在株高、生物量、光合作用等方面,大多达到与对照(0 g.L-1)未施氮肥无显著差异的水平,而对50 g.L-1水浸液处理的苜蓿幼苗,虽有一定的缓解作用,但这种作用均未达到与对照(0 g.L-1)未施氮肥时无显著差异的水平。 4. UV-B增强辐射处理花椒后,花椒的化感潜力显著增强。花椒叶片内UV-B吸收物质的含量和总酚含量均显著增加。 In recent years, with profound research on the reasons of continuous cropping obstacles, allelopathy received increasing attention to many scholars at home and abroad. Zanthoxy bungeanum as a Rutaceae plant is a high economic value species which gains early and uses widely. Zanthoxylum is an important economic crop in the arid valley of western Sichuan region, and its not even has received much concern for the continuous cropping obstacles. The systematic study of allelopathy of Zanthoxylum will contribute to the understanding and final settlement of this issue. The major allelopathic composition was separated through the extraction, chromatography combined with other methods. The impact on soil nutrient cycling was also studied through the addition of different concentrations of water extracts of Zanthoxylum. Furthermore, the effects of water extracts of Zanthoxylum leaves on alfalfa leaf physiological and biochemical indexes, photosynthesis, soil enzymes and nutrient uptake of nitrogen and the mitigation of allelopathy through using external fertilizer were studied to put forward scientific resolvent for Zanthoxylum continuous cropping obstacles .The response of allelopathic potential of Zanthoxylum to global change - UV-B enhanced radiation was studied . The main findings are as follows: 1. Through extraction with different polar organic solvents on concentrated water extract of Zanthoxylum leaf and then using column chromatography combined with detection of biological activity, one of the main allelopathic components- methoxy-phenol was isolated. The biological activity testing of the pure material of methoxy-phenol proved that it does have allelopathic potential. 2. Thirty days after treating soil with water extract of Zanthoxylum leaf, as compared with the control, the contents of soil nitrate, ammonium, nitrate plus ammonium nitrogen showed a trend of decrease with the increase of the concentration of water extract whereas the content of ammonium nitrogen showed a significant reduction, and the content of nitrate did not change significantly, the content of nitrate plus ammonium nitrogen also showed a significant (P <0.05) redction. The activity of soil urease and protease showed the same trend as the content of nitrate nitrogen plus ammonium nitrogen. With the increase in the concentration of water extract, the number of ammonification bacteria significantly reduced but nitrogen-fixing bacteria did not change significantly and there was a decreasing trend in the number of nitrifying bacteria and denitrifying bacteria. Sixty days after the treatment, with the increase in solution concentration of water extract of Zanthoxylum leaf, the content of nitrate、 ammonium nitrogen, nitrate plus ammonium nitrogen showed a similar change trend to 30 days’; the number of ammonification bacteria, nitrogen-fixing bacteria significantly reduced ; the number of nitrifying bacteria, denitrifying bacteria was still an downward trend; the activity of soil urease and protease showed the same trend as the 30th days’. Compared to the results of the 30th days’, the content of nitrate, ammonium, nitrate plus ammonium nitrogen showed a decrease trend between the treatment of same concentration, but there was no significant difference except the treatment of 25g.L-1 between the same concentration; the activity of soil urease showed enhanced trend except the treatment of 12.5 g.L-1; the activity of protease increased to varying degrees; the number of ammonification bacteria、 nitrifying bacteria and denitrifying bacteria were growing while nitrogen-fixing bacteria reduced.. 3. With the increase of the concentration of water extract of Zanthoxylum leaf, the water extract significantly inhibited the root length, aboveground biomass, content of chlorophyll and soluble protein in leaf and net photosynthetic rate. The activity of four antioxidant enzymes (POD, SOD, CAT, APX) reduced with the increase in concentration of the water extract but the content of MDA increased. The activity of enzymes related to primary nitrogen assimilation such nitrate reductase (NR), glutamyl synthetase (GS), glutamate dehydrogenase (GDH) were subject to different degrees with an increase in the concentration of water extracts. In general, the activity of nitrate reductase, glutamyl synthetase were inhibited, while change in the activity of glutamate dehydrogenase was more complex. The activity of glutamate dehydrogenase in leaf was first reduced and then increase,but did not change significantly in root. After using two external different concentrations of nitrogen fertilizer, there was a significant mitigation in inhibiton in plant height, biomass, photosynthesis, etc. in the treatment of 12.5,25 gL-1 of water extract of Zanthoxylum leaf, and most of these indexes showed no significant difference with the control (0 g.L-1, no external fertilizer was added) .Although there showed a certain degree of ease in the treatment of 50 g.L-1 , there was still a significant difference compared with the control (0 gL-1) in which no external fertilizer was used. 4.The allelopathic potential of Zanthoxylum positively responded to enhanced UV-B significantly. The content of UV-B absorbing compounds and the total phenol also significant increased.
Resumo:
土壤微生物量、可溶性有机碳与氮虽然只占土壤有机碳、氮总量的较小部分,但可以在土壤全碳、氮变化之前反映土壤微小的变化,又直接参与土壤生物化学转化过程,因而在植被恢复过程中,较其它土壤理化性质等能够更好地指示土壤恢复情况。在青藏高原东缘存在大面积的次生人工林替代灌丛或采伐迹地,而关于这些人工林替代后的生态效果和生态过程的评估却十分缺乏,本研究通过评估岷江上游植被恢复重建过程中典型人工替代次生植被凋落物层与土壤碳、氮等养分大小,动态监测土壤微生物生物量、水溶性碳、氮等指标,结合温度与凋落物输入等影响土壤活性有机碳、氮因子的控制试验,系统分析不同人工替代次生植被土壤碳、氮等养分的差异原因,试图寻找低效人工林优化调控与持续管理技术,为区域生态公益林持续管理提供理论和技术依据。主要结论如下: 1. 通过对不同人工替代次生植被凋落物层和土壤碳、氮分析发现,油松和华山松人工林替代次生灌丛后土壤碳、氮含量较灌丛和阔叶人工林低,主要原因可能为凋落物质量(C/N)较差,而引起碳、氮等元素难以归还土壤。进而通过对不同人工替代次生植被凋落物层和土壤微生物生物量、水溶性有机碳、氮等指标的季节性动态模式的分析,发现各次生植被土壤微生物生物量C、N,P以及土壤水溶性碳、氮含量均呈明显季节性动态,呈现秋季明显大于其它季节,冬季最低,在表层土壤最为明显。 2. 油松、华山松人工林凋落物层和土壤水溶性有机碳(WDOC)、土壤水溶性有机氮(WDON)明显低于灌丛和连香树,土壤微生物生物量C、N也以油松和华山松人工林最低,而落叶类植被,如灌丛、连香树和落叶松之间没有明显差异,说明可利用底物的数量和质量差异是影响各次生植被凋落物分解和土壤微生物活性的主要原因。MBC/OC和MBN/ON能较好地指示土壤微生物活性的变化,MBC/OC凋落层总体以灌丛和连香树人工林最高,油松和华山松人工林最低;而土壤中MBC/OC连香树人工最高,华山松人工林最低。说明以油松和华山松为主的人工造林替代乡土阔叶灌丛造成土壤C、N等养分严重匮乏,微生物活性低下是影响其养分周转的主要原因。 3. 从各次生植被凋落物产生看,凋落物年归还量最大的为华山松人工林(5.1×103 kg ha-1),其次为落叶松人工林(4.8×103 kg ha-1),阔叶灌丛林地凋落物产生总量(4.4×103 kg ha-1)略大于油松人工林(4.2×103 kg ha-1),最小的为连香树人工林(3.6×103 kg ha-1);叶是凋落物的主体,落叶类树种月动态表现为单峰型,高峰主要在10-11月,如落叶松、连香树和灌丛林;常绿的松类月动态不明显,各月基本相同,最为明显地为油松林,华山松人工林略有二个小峰,分别出现在11月和5月。落叶阔叶灌丛的凋落物分解速率大于常绿针叶林,如油松和华山松。结合凋落物的产生量和分解速率,不同树种人工林替代次生阔叶灌丛后,人工油松和华山松林枯落物总贮量和厚度明显大于落叶松人工林、灌丛林和连香树人工林,说明以油松和华山松为主的人工造林替代乡土阔叶灌丛延缓了有机物向土壤的顺利归还,不利于土壤C、N等养分循环。 4. 通过控制地面凋落物和地下根系输入有机物对土壤碳、氮的影响研究发现,(1) 单独去除根系以及根系与地面凋落物同时去除处理1年后对表层(0-10cm)土壤WDOC均没有显著影响,而土壤WDON显著增加,油松人工林土壤微生物生物量C、N显著降低,人工落叶松林没有显著差异,说明油松人工林土壤微生物活性对地下碳输入的依赖大于其它次生植被,而落叶松土壤微生物活性对地下碳输入依赖性较小;去除地面凋落物,明显降低了落叶松人工林土壤WDOC,华山松和连香树土壤WDON均较对照显著减少,油松人工林土壤微生物量C较对照显著减少;双倍增加地面凋落物处理对土壤微生物生物量、WDOC和WDON没有明显地增加,相反,连香树、华山松和油松人工林土壤WDON较对照减少。说明油松人工林微生物活性不仅依赖于地下碳输入,而且对地上有机物输入的依赖性也较大;连香树、落叶松和华山松人工林土壤微生物生物量并没有因地面凋落物的去除减少可能与土壤总有机碳含量及活性均较高有关,而双倍增加地面凋落物反而降低了土壤微生物生物量,说明凋落物覆盖后改变了土壤微气候。 5. 碳矿化累积量与有机碳含量和活性有机碳含量之间存在显著地正相关关系。凋落物碳累积矿化量、矿化速率以连香树最高,油松和华山松人工林次之,落叶阔叶灌丛低于常绿针叶纯林,导致其差异的主要原因可能为凋落物产生的时间动态模式不一样,致使凋落物起始分解时间不一致。而土壤层有机碳矿化速率和矿化量以阔叶落叶灌丛和连香树最高,油松和华山松人工土壤最低,再次证实利用针叶纯林恢复植被阻碍了有机质周转与循环。 6. 凋落物累积矿化量与C/N值呈显著地相关关系,并随着温度的升高而明显增加,而土壤累积矿化量与C/N值没有显著相关关系,说明土壤有机碳质量(C/N)对温度的响应不十分明显。通过双指数模型对不同温度下碳矿化过程进行模拟和计算出活性有机碳与惰性有机碳比例,发现温度升高促进了惰性有机碳向活性有机碳的转化,增加了活性有机碳含量,说明温度升高可促进次生植被凋落物与土壤有机质的分解,进而可影响到林地碳源/汇关系的变化。 综上,通过对不同人工替代次生植被凋落物与土壤C、N大小、以及土壤微生物生物量、水溶性C、N等指标动态变化模式研究,结合温度与凋落物数量输入等影响土壤活性C、N因子的综合分析,以油松和华山松人工纯林对山地植被恢复,延缓或阻碍了有机质周转与循环,造成了土壤肥力退化。对现有低效人工纯林改造,应为地面大量有机物分解创造条件。 Although soil microbial biomass, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) are a small part of total soil organic carbon and nitrogen, they can directly participate in the process of soil biochemical translation and indicate the fine changes before changes of soil total organic carbon and nitrogen occur. So, they are good indexes to indicate soil restoration condition during the process of vegetation rehabilitation. There are large areas of secondary vegetations which substitute for indigenous shrubs in the eastern fringe of Qinghai-Tibet Plateau. However, it is not well known that the ecological effect and process after substitution by different secondary plantations. Based on comparison of soil organic and nitrogen contents in litter layer and soil under different secondary vegetations in upper reaches of Minjiang River, soil microbial biomass, DOC and DON in litter layer and soil were investigated in order to analyze the seasonal dynamic. Combining the effects of temperature, litter input and root exclusion on soil microbial biomass, DOC and DON, we also aim to understand the reason and mechanism of difference in soil carbon and nitrogen contents among different secondary vegetations. The study would contribute to comprehensively understanding C and N cycling processes and provide optimal control and sustainable technology of low-effect plantations in these regions. The results are as follows: (1) Organic carbon and nitrogen in litter layers and soil under different substitution plantations were investigated. The results showed that contents of soil organic carbon and nitrogen were lower in P. tabulaeformis (PT) and P. armandi Franch(PA) than those in native broad-leaf shrub and broad-leaf plantation. The low quality (C/N) of litter in PT and PA plantations caused carbon and nitrogen returning to soil difficultly. Seasonal dynamic of soil microbial carbon (MBC),-nitrogen (MBN),-phosphor (MBP), and WDOC and WDON showed similar pattern, which had the highest values in autumn and the lowest values in winter. (2) WDOC and WDON in litter layers and soil under PT and PA plantations were significantly lower than those in native broad-leaf shrub and Cercidiphyllum japonicum Sieb. et Zucc.(CJ). Soil MBC and MBN were also the lowest, while there were no significant differences among deciduous vegetations, i.e. native broad-leaf shrub, CJ and Larix kaempferi Lamb.(LK) plantation. The results suggested that difference in quantity and quality of available substance was main reason that affected the activity of microbe in soil and litter layer. MBC/OC and MBN/ON were good indexes to indicate the change of soil microbial activity. MBC/OC of litter had the highest value under native broad-leaf shrub and CJ plantation, and had the lowest value in PT and PA plantations, while MBC/OC of soil was the highest under CJ plantation, and was the lowest in PT and PA plantations. These results indicated that PT and PA plantations substituting for native broad-leaf shrub caused deficit of carbon and nitrogen in soil, low microbial activity was a main reason influencing the cycling and turnover of carbon and nitrogen in soil. (3) The annual litter fall production, composition, seasonal dynamic and decomposition of five typical secondary stands in upper reaches of Minjiang River were studied in this paper. The annual litter productions were: PA (5.1×103 kg ha-1), LK(4.8×103 kg ha-1), native broad-leaf shrub (4.4×103 kg ha-1), PT(4.2×103 kg ha-1),CJ(3.6×103 kg ha-1). The litter production of leaves in five secondary vegetations occupied a higher percentage in the annual total litter production than those of other components. The litterfall was mostly producted in the cool and dry period (October-November) for deciduous vegetations and relatively equably producted in every season for evergreen coniferous vegetations. The decomposition rate of leaf litter in the broad-leaf stand was higher than those in evergreen coniferous stand. Combined with annual litter fall production and decomposition rate of leaf litter, we found that stock and depth of litter layer were significantly larger in PT and PA plantations than those in native broad-leaf shrub, LK and CJ plantations. The results confirmed that PT and PA plantations substituting for native broad-leaf shrub delayed organic matter returning to soil and hindered cycling of carbon and nitrogen again. (4) We explored plant litter removal, double litter addition, root trenching, and combining root trenching and litter removal treatments to examine the effects of above- and belowground carbon inputs on soil microbial biomass, WDOC and WDON in four secondary plantations. During the experimental period from June 2007 to July 2008, 1 year after initiation of the treatments, WDOC in soil did not vary in root trenching, and combining root trenching and litter removal treatments, while WDON in soil significantly increased compared with CK treatment. Root trenching reduced soil MBC and MBN in PT plantation, while MBC and MBN in soil did not vary in LK plantation. The rasults implied that soil microbial activity was more dependent on belowground carbon input in PT plantation than those in other secondary plantations, on the contrary, soil microbial activity in LK plantation was not dependent on belowground carbon input. Plant litter removal significantly decreased soil WDOC in LK plantation, decreased WDON in PA and CJ plantations, and also significantly reduced soil MBC in PT plantation. However, double litter addition did not increase soil microbial biomass, WDOC and WDON, on the contrary, soil WDON in CJ, PA and PT plantations were decreased. These suggested that soil microbial activity was not only dependent on belowground carbon input, but also on aboveground organic material input. Double litter addition could change the microclimate and result in the decrease of soil microbial activity in CJ, PA and PT plantations. (5) We measured carbon mineralization in a 107 days incubation experiment in 5℃,15℃ and 25℃. Carbon cumulative mineralization was positively correlated with organic matter and labile organic carbon in litter layer and soil. Cumulative carbon mineralization and mineralization rate of litter layers in PT and PA plantations were higher than that in native broad-leaf shrub. This difference between native broad-leaf shrub and coniferous plantations in cumulative carbon mineralization and mineralization rate of litter layers could be attributed to the initiating time of decomposition due to the difference in seasonal dynamic of litter fall production between two types of secondary plantations. However, cumulative carbon mineralization and mineralization rate in soil were the highest in native broad-leaf shrub and CJ plantation, and were the lowest in PT and PA plantations. This also confirmed that PT and PA plantations substituting for native broad-leaf shrub hindered the cycling and turnover of organic matter again. (6) Carbon cumulative mineralization was positively correlated with C/N in litter layer and increased with temperature increasing, while carbon cumulative mineralization was not correlated with C/N in soil. This indicated that soil organic matter quality (C/N) was insensitive to temperature. Applying bi-exponential model, we computed the percent of labile and stable carbon in different temperature incubation and found that temperature increasing would accelerate the transform from stable carbon to labile carbon and increase the percentage of labile organic carbon. This illuminated that temperature incraesing could facilitate the decomposition of litter and soil organic matter in secondary vegetations and hence affect the relationship between carbon source and sink.
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花椒(Zanthoxylum bungeanum)是川西干旱河谷地区重要的经济作物,化感作用是花椒连作障碍的重要原因之一。花椒凋落物分解是影响花椒林地土壤肥力及花椒产量的重要因素,因此系统研究花椒化感作用是否对花椒凋落物的分解产生影响可以为解决花椒连作障碍导致的产量下降等问题提供科学的理论依据。本文通过室内模拟实验研究了花椒凋落物的四个分解动态以及分解后凋落物浸提液对花椒林地土壤性质的影响;通过野外盆栽实验研究了花椒凋落物浸提液对花椒幼苗的生长、花椒凋落物的质量及土壤性质的影响。最终从生理生化角度揭示了花椒的化感作用对凋落物分解的影响机理,为深入解决花椒连作障碍问题、对花椒凋落物采取有效的人工措施提供了科学的理论依据。主要的研究结果如下: 1. 室内分解实验证明,花椒凋落物在分解的60 d 内分解速率呈现由大到小的变化趋势,并且凋落物分解呈现明显的毒性动态。凋落物在分解的10 d、30 d 时,分解速率较大,30 d 以后分解速率显著降低。凋落物分解的10 d 左右酚酸释放量最大,此时凋落物的毒性最大,凋落物分解到10 d 以后,酚酸释放量逐渐减少,凋落物的毒性也逐渐减小。 2. 四个分解动态的花椒凋落物浸提液对土壤化学性质产生了显著的影响。花椒凋落物在分解的60 d 内,其浸提液使土壤pH值均显著的增加。分解0 d 的凋落物浸提液显著的降低了土壤铵态氮的含量,抑制了纤维素分解菌的生长;分解60 d 的凋落物浸提液显著的降低了土壤酚酸含量,增加了土壤有效磷的含量;分解30 d 和60 d 的凋落物浸提液均显著的促进了好气性纤维素分解菌的生长。这说明花椒凋落物在分解过程中呈现出明显的毒性动态:凋落物分解的初期毒性作用较大,随着分解的继续进行特别是在分解的30 d 以后,其毒性作用慢慢降低。 3. 花椒凋落物浸提液对花椒幼苗表现出明显的化感作用。不同浓度的浸提液对花椒幼苗地上及地下生物量、叶面积均产生了显著的抑制作用,并且随着浸提液浓度的升高抑制作用加强。凋落物浸提液对叶片厚度的影响较小,只有Y1对叶片厚度的生长抑制作用显著。 4. 花椒的化感作用改变了凋落物的质量,并对凋落物分解产生了显著的影响。对花椒幼苗用不同浓度的凋落物浸提液进行处理,Y1使凋落物有机碳含量、木质素含量、C/N、木质素/氮显著降低,纤维素含量显著升高;Y3使凋落物有机碳含量、木质素含量、C/N、木质素/氮显著升高。花椒凋落物质量的改变显著的影响了凋落物的分解,凋落物的分解速率大小依次为:Y1(10.15 a-1)> Y2(8.71 a-1)> CK(6.41 a-1)> Y3(5.08 a-1)。 5. 花椒的化感作用改变了土壤性质,并对凋落物分解产生了显著的影响。对花椒幼苗用不同浓度的凋落物浸提液处理的同时,也改变了土壤性质。不同浓度的凋落物浸提液显著的升高了土壤pH值、有机碳含量。各种浓度的凋落物浸提液对土壤多酚氧化酶的活性均起到了显著的促进作用。凋落物浸提液Y1对土壤纤维素分解酶的活性、细菌和真菌的生长也具有显著的促进作用。土壤性质的改变显著的影响了凋落物的分解,凋落物的分解速率大小依次为:Y1(10.30 a-1)>Y2(9.60 a-1)>CK(6.41 a-1)>Y3(6.29 a-1)。 6. 不论是凋落物质量发生改变还是土壤性质发生改变,在凋落物分解的整个过程中,C元素始终处于单调净释放的状态,并且C释放量与分解速率成显著的正相关,即凋落物分解越快,凋落物C释放量越大。凋落物分解过程中,均出现了酚酸大量释放的情况,并与凋落物分解速率成显著正相关。凋落物分解后的木质素含量、木质素/氮均增加,并且随着浸提液浓度的升高,凋落物木质素含量、木质素/氮升高。 Zanthoxylum bungeanum is an important economic crop in dry valley of the Minjiang river (Sichuan, Southwest China), but allelopathy is one of the important reasons for its continuous cropping. Zanthoxylum bungeanum litter decomposition affects Zanthoxylum bungeanum soil fertility and its output. So systemically investigate if allelopathy affects litter decomposition could provide the scientific methods to solve the problem of output fall caused by the continuous cropping. In this paper, the releasing dynamics of phenolic acid during Zanthoxylum bungeanum litter decomposition (0, 10, 30 and 60 days) and the effects of its aqueous extract on soil chemical properties were investigated via the laboratory study. Effects of Zanthoxylum bungeanum litter aqueous extract on the growth of young Zanthoxylum bungeanum seedlings, litter qualities and the soil qualities were investigated via the field study. Finally, we open out the action manner of Zanthoxylum bungeanum allelopathic effect on the litter decomposition, and provide the theoretical basis to solve the Zanthoxylum bungeanum continuous cropping. The main results showed that: 1. The laboratory litter decomposition experiment showed a trend of decomposition rate from large to small and an occurrence of phytotoxicity with clear dynamic patterns during Zanthoxylum bungeanum litter decomposition. The litter decomposition rate was larger at the tenth and 30th day during 60-day litter decomposition and gradually decreased after 30 days of litter decompostion. The releasing quantity of the litter phenolic acid was the highest at the tenth day, and here, the litter toxicity was the biggest. The releasing quantity of the litter phenolic acid gradually decreased after 10 days of litter decomposition, so the phytotoxicity of litter was gradually decreased with the litter decomposition. 2. The Zanthoxylum bungeanum litter aqueous extract after four decomposition stages had significantly effect on the soil chemical qualities. The pH value in soil was significantly increased in litter aqueous extract of four decomposition stages. The NH+4-N concentration was significantly decreased in soil amended with litter aqueous extract of 10-day decomposition which inhibited the growth of Aerobic cellulose-decomposer. The growth of soil Aerobic cellulose-decomposer was promoted by the litter aqueous extract of 30-day decomposition. Available phosphorus concentration was significantly increased and phenolic acid content was significantly decreased in soil amended with litter aqueous extract of 60-day decomposition which promoted the growth of Aerobic cellulose-decomposer. The study results showed an occurrence of phytotoxicity with clear dynamic patterns during Zanthoxylum bungeanum litter decomposition. The phytotoxicity of litter was the largest at the initial stage, but the phytotoxicity gradually decreased with the litter decomposition, especially after 30 days of decomposition. 3. The field study indicated that the Zanthoxylum bungeanum litter aqueous extract had significant allelopathic effects on the growth of young seedlings.Different concentration aqueous extract had signinficant inhibiting effects on biomass and leaf area of young seedlings. The inhibiting effect on the biomass strengthened with the litter aqueous extract concentration augment. Litter aqueous extracts had less effect on the leaf thickness, and only Y1 had significant inhibiting effect on the leaf thickness. 4. The Zanthoxylum bungeanum allelopathy had significant effect on the litter qualities and the litter decomposition. Treating the young Zanthoxylum bungeanum seedlings with different concentration of litter aqueous extracts, the leaf litter organic C, lignin, C/N and lignin/N all decreased and the cellulose content increased under Y1 treatment. The leaf litter organic C, lignin, C/N and lignin/N all increased under Y3 treatment. So the litter decomposition was significant affectded by the litter qualities, and the litter decomposition rate was Y1(10.15 a-1)> Y2(8.71 a-1) > CK(6.41 a-1) > Y3(5.08 a-1). 5. The Zanthoxylum bungeanum allelopathy had significant effect on the soil qualities and the litter decomposition. Treating the young Zanthoxylum bungeanum seedlings with different concentration of litter aqueous extracts, also changed the soil qualities. Different concentration of litter aqueous extracts had significant effects on the soil pH and organic C content. Every concentration of litter aqueous extracts accelerated the soil Polyphenol Oxidase activity and Y1 accelerated the soil Cellulase activity, the number of soil bacteria and fungi. So the litter decomposition was significant affected by the soil qualities, and the litter decomposition rate was Y1(10.30 a-1) > Y2 (9.60 a-1) >CK(6.41 a-1)>Y3(6.29 a-1)。 6. Whether the litter or soil qualities changed, the litter C element at the state of release at all times during the litter decomposition, and the release quantity increased with the decomposition rate augment. Litter released plentiful total penolics content during decomposition, and the release quantity had the positive correlation with the litter decomposition rate. The litter lignin content and the lignin/N all increased with the litter aqueous extracts concentration augment after litter decomposition.
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土壤线虫是土壤动物的主要功能类群之一,在土壤养分分解和循环中起到重要的作用。本研究通过施用两种形态氮肥,硝态氮(NO-3–N)和铵态氮(NH+4–N),对黄瓜整个生长期内根际土壤线虫的群落组成、结构及其多样性等的影响进行了比较研究。为增进土壤健康,提高土壤质量以及合理施用氮肥提供科学的理论依据。研究结果如下: 1. 氮肥处理后,不同浓度NO-3–N处理提高了根际土壤线虫数量,而NH+4–N处理(202.5 kg N/ hm2)抑制了线虫数量的增加。线虫群落结构相对较稳定,营养类群变化不大。且少量优势科/属会对土壤线虫群落特征起着至关重要的作用。 2. 在整个生长季节内,非寄生线虫的群体动态变化与寄生线虫的群体的动态变化具有相反的变化趋势。其中,NO-3–N处理和NH+4–N处理后植物寄生线虫出现频率的变化趋势相近,都是由高到低;非植物寄生线虫出现频率的变化趋势则都是由低到高。这说明非植物寄生线虫数量增长和空间占位对植物寄生线虫群体有一定的抑制作用。另外,也反映了适量氮肥在一定程度上能够减轻植物寄生线虫对黄瓜的危害。 3. 由多样性指数变化可知,NO-3–N和NH+4–N在低肥(67.5 kg N/hm2)和高肥(202.5 kg N/hm2)处理较中肥(135.0 kg N/hm2)处理,更不利于提高土壤线虫多样性地提高和线虫群落的稳定性。中肥不同形态氮肥处理与对照相比,H´指数在初花期和结果期显著增加了土壤线虫的H´指数,说明施入两种形态的氮肥能够提高线虫生物多样性程度。NH+4–N处理在初花期和结果期显著降低了土壤线虫种类的丰富性。土壤线虫生物多样性变化中,H´和SR指数在一定程度上能够反映施用无机氮肥对土壤线虫的多样性的影响,而J指数和λ指数效果不明显。NO-3–N和NH+4–N处理相比较,NO-3–N处理对黄瓜土壤线虫的多样性指数影响更大,促进了土壤线虫群体的多样化和种类的丰富度,更有利于提高土壤线虫的多样性,增加其稳定性。这些结果表明适量无机氮肥特别是NO-3–N的施用对黄瓜土壤线虫的生物多样性有一定的维护和提高作用。 4. 线虫数量与土壤质量指标的相关分析表明:线虫数量与有关土壤理化生指标,如土壤NO-3–N、NH+4–N、有机质含量等的正相关程度高,与总酚含量等显著负相关;与根际土壤微生物,细菌、真菌、放线菌数量等呈显著正相关。另外,线虫数量与土壤含水量未表现出显著相关关系。 Nematodes play a major role in decomposition and nutrient cycling in soil. Nematode community analyses are useful in assessing soil ecosystem status and function. The effects of two forms of mineral nitrogenous fertilizers (NO-3–N and NH+4–N) on nematode community composition, structure and diversity in rhizosphere of cucumber were investigated during different growing seasons of cucumber. Systematically research of effects of nitrogenous fertilizers could help to obtain better undstanding of a healthy soil and using nitrogenous fertilizers in reason. The main results are as follows: 1. The total numbers of nematode were more abundant in NH+4–N treatments than other teatments. However, NH+4–N teatment(202.5 kg N/hm2)dramatically inhibited it. All the tropic groups in the soil nematode communities were stable, and the dominant family or genus had an important function in the nematode community structure. 2. There was similar trend of the frequency of plant parasitic nematodes between NO-3–N and NH+4–N treatment, the similar trend of the frequency of non-plant parasitic nematodes was also found. But the frequency of plant parasitic nematodes exhibited a contrary trend to that of plant parasitic nematodes after different nitrogenous fertilizer treatments. The results showed that the increasing trend of the frequency and the niche of non-plant parasitic nematodes inhibited the plant parasitic nematodes, and indicated that right chemical fertilizers dosage could abate plant parasitic nematodes harm to cucumber. 3. The changes of the biodiversity index showed that the nitrogen treatment(135.0 kg N/hm2)promoted the stabilization of soil nematode diversity than other nitrogen treatments(67.5 kg N/hm2 and 202.5 kg N/hm2). In the treatment(135.0 kg N/hm2),The changes in nematode diversity between the control plots and treated plots were compared by the biodiversity index (H´, J, SR, λ). Among these tested index, H´ and SR were effective in reflecting the effects of different nitrogenous fertilizers on the diversity of soil nematodes. In comparison with the NH+4–N treatment, the NO-3–N treatment promoted the stabilization of soil nematode diversity. 4. Correlation coefficients between nematode abundance and soil quality indices indicated that the total numbers of nematode were affected positively by NO-3–N, NH+4–N and the organic matter, and negatively by total phenolic acids; the total num- bers of nematode had positive correlation with bacteria, fungi and actinomycetes nu- mbers. Soil water contents had only a weak negative influence on it.
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青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.
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在人类活动导致全球变暖的前提下,由于全球气温的升高,地表水分加速向空中蒸发。从20世纪70年代至今,地球上严重干旱地区的面积几乎扩大了一倍。这一增长的一半可归因于气温升高而不是降雨量下降,因为实际上同期全球平均降水量还略有增长。干旱对陆地植物和农林生态系统产生深远影响,并已成为全球变化研究的一个重要方面。位于青藏高原东部的川西亚高山针叶林是研究气候变暖对陆地生态系统影响的重要森林类型。森林采伐迹地、人工林下和林窗环境作为目前该区人工造林和森林更新的重要生境,其截然不同的光环境对亚高山针叶林更新和森林动态有非常重要的影响。凋落物产生的化感物质可通过影响种子萌发和早期幼苗的定居而影响种群的建立和更新,而人工林和自然林物种以及更新速度的差异性也都受凋落物的影响。 云杉是川西亚高山针叶林群落的重要树种之一,在维持亚高山森林的景观格局和区域生态安全方面具有十分重要的作用,其自然更新能力及其影响机制一直是研究的热点问题。本试验以云杉种子和2年生幼苗为研究对象,从萌发、根尖形态、幼苗生长、光合作用、渗透调节和抗氧化能力等方面研究了不同光环境下水分亏缺和凋落物水浸液对云杉种子和幼苗生长的影响。旨在从更新的角度探讨亚高山针叶林自然更新的过程,其研究成果可在一定程度上为川西亚高山针叶林更新提供科学依据,同时也可为林业生产管理提供科学指导。主要研究结论如下: 水分亏缺在生长形态、光合作用、抗氧化能力、活性氧化对云杉幼苗都有显著影响。总体表现为,水分亏缺导致了云杉幼苗的高度、地径、单株总生物量降低,增加了地下部分的生长;水分亏缺显著降低了云杉叶片中相对含水量、光合色素、叶氮含量,净光合速率和最大量子产量(Fv/Fm),提高了幼苗叶片中膜脂过氧化产物(MDA)的含量;水分亏缺提高了幼苗叶片中过氧化氢(H2O2)含量,超氧荫离子(O2-)生成速率以及脯氨酸和抗氧化系统的活性(ASA, SOD, CAT, POD, APX和GR)。从这些结果可知,植物在遭受水分亏缺导致的伤害时,其自身会形成防御策略,并通过改变形态和生理方面的特性以减轻害。但是,这种自我保护机制依然不能抵抗严重水分亏缺对植物的伤害。 模拟林下低光照条件显著增加单株植物的地上部分生长,尤其是其叶片的比叶面积(叶面积/叶干重),同时其光合色素含量和叶片相对含水量也显著增加,这些改变直接导致植株光合速率和生物量的增加。同时,与高光照水平相比,低光照幼苗的膜脂过氧化产物(MDA)和活性氧物质均较低,显示出低光照比高光照水平对植物的更低的氧化伤害。尽管低光照也导致大部分抗氧化酶活性降低,但这正显示出植物遭受低的氧化伤害,更印证了前面的结论。 凋落物水浸液影响了云杉种子的萌发和根系的生长,更在形态、光合作用、抗氧化能力、活性氧物质以及叶氮水平上显著影响了云杉幼苗,其中,以人工纯林凋落物的影响更有强烈。具体表现在,种子萌发速率和萌发种子幼根的长度表现为对照>自然林处理>人工纯林;凋落物水浸液抑制种子分生区和伸长区的生长,人工林处理更降低了根毛区的生长,使根吸水分和养分困难。对2年生幼苗的影响主要表现在叶绿素含量、光合速率以及叶氮含量的降低;膜脂过氧化产物、活性氧物质和抗氧化酶系统的显著增加。同样的,人工纯林处理对云杉幼苗的影响显著于自然林处理。 在自然生态系统中,由于全球变暖气温升高导致的水分亏缺和森林凋落物都存在森林的砍伐迹地,林窗和林下环境中。我们的研究表明,与迹地或林窗强光照比较,林下的低光照环境由于为植物的生长营造了较为湿润的微环境,因此水分亏缺在林下对云杉幼苗造成的影响微弱。这可以从植物的形态、光合速率以及生物量积累,过氧化伤害和抗氧化酶系统表现出来。另一方面,凋落物水浸液在模拟林下低光照环境对植物的伤害也微弱于强光照环境,这与强光照环境高的水分散失导致环境水分亏缺有关;而人工纯林处理对云杉幼苗的伤害比对照和自然林处理显示出强烈的抑制作用。 Under the pre-condition of global warming resulted from intensive human activities, water in the earth’s surface rapidly evaporates due to the increase of global air temperature. From 1970s up to now, the area of serious drought in the world is almost twice as ever. This increase might be due to the increasing air temperature and not decreasing rainfall because global average rainfall in the corresponding period slightly is incremental. Drought will have profound impacts on terrestrial and agriculture-forest system and has also become the important issue of global change research. The subalpine coniferous forests in the eastern Qinghai-Tibet Plateau provide a natural laboratory for the studying the effects of global warming on terrestrial ecosystems. The light environment significantly differs among cutting blanks, forest gap and understory, which is particularly important for plant regeneration and forest dynamics in the subalpine coniferous forests. Picea asperata is one of the keystone species of subalpine coniferouis forests in western China, and it is very important in preserving landscape structure and regional ecological security of subalpine forests. The natural regeneration capacities and influence mechanism of Picea asperata are always the hot topics. In the present study, the short-term effects of two light levels (100% of full sunlight and 15% of full sunlight), two watering regimes (100% of field capacity and 30% of field capacity), two litter aqueous extracts (primitive forest and plantation aqueous extracts) on the seed germination, early growth and physiological traits of Picea asperata were determined in the laboratory and natural greenhouse. The present study was undertaken so as to give a better understanding of the regeneration progress affected by water deficit, low light and litter aqueous extracts. Our results could provide insights into the effects of climate warming on community composition and regeneration behavior for the subalpine coniferous forest ecosystem processes, and provide scientific direction for the forest production and management. Water deficit had significant effects on growth, morphological, physiological and biochemical traits of Picea asperata seedlings. Water deficit resulted in the decrease in height, basal diameter, total biomass and increase in under-ground development; water deficit significantly reduced the needle relative water content, photosynthetic pigments, needle nitrogen concentration, net photosynthetic rate and the maximum potential quantum yield of photosynthesis (Fv/Fm), and increased the degree of lipid peroxidation (MDA) in Picea asperata seedlings; water deficit also increased the rate of superoxide radical (O2-) production, hydrogen peroxide (H2O2) content, free proline content and the activities of antioxidant systems (ASA, SOD, POD, CAT, APX and GR) in Picea asperata seedlings. These results indicated that some protective mechanism was formed when plants suffered from drought stress, but the protection could not counteract the harm resulting from the serious drought stress on them. Low light in the understory significantly increased seedling above-ground development, especially the species leaf area (SLA), and photosynthetic pigments and relative needle content. These changes resulted in the increase in net photosynthetic rate and total biomass. Moreover, the lower MDA content and active oxygen species (AOS) (H2O2 and O2-) in low light seedlings suggested that low light had weaker oxidative damage as compared to high light. Lower antioxidant enzymes activities in low light seedlings indicated the weaker oxidative damage on Picea asperata seedlings than high light seedlings, which was correlative with the changes in MDA and AOS. Litter aqueous extracts affected seed germination and root system of Picea asperata seedlings. Significant changes in growth, photosynthesis, antioxidant activities, active oxygen species and leaf nitrogen concentration were also found in Picea asperata seedlings, and plantation treatment showed the stronger effects on these traits than those in control and primitive forest treatment. The present results indicated that seed germination and radicle length parameters in control were superior to those in primitive forest treatment, and those of primitive forest treatment were superior to plantation treatment; litter aqueous extracts inhibited the meristematic and elongation zone, and plantation treatment caused a decrease in root hairs so as to be difficult in absorbing water and nutrient in root system. On the other hand, litter aqueous extracts significantly decreased chlorophyll content, net photosynthetic rate and leaf nitrogen concentration of Picea asperata seedlings; MDA, AOS and antioxidant system activities were significantly increased in Picea asperata seedlings. Similarly, plantation treatment had more significant effect on Picea asperata seedlings as compared to primitive forest treatment. In the nature ecosystem, water deficit resulted from elevating air temperature and litter aqueous extract may probably coexist in the cutting blank, forest gap and understory. Our present study showed that water deficit had weaker effects on low light seedlings in the understory as compared to high light seedlings in the cutting blank and forest gap. The fact was confirmed from seedlings growth, gas exchange and biomass accumulation, peroxidation and antioxidant systems. This might be due to that low light-reduced leaf and air temperatures, vapour-pressure deficit, and the oxidative stresses can aggravate the impact of drought under higher light. On the other hand, litter aqueous extracts in the low light had weaker effects on the Picea asperata seedlings than those at high light level, which might be correlative to the water evapotranspiration under high light. Moreover, plantation litter aqueous extracts showed stronger inhibition for seed germination and seedling growth than control and primitive forest treatments.
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土壤可溶性有机物质(Dissolved organic matter,DOM)作为土壤有机质的活性组分,在陆地生态系统物质循环中扮演非常重要的角色。土壤DOM的主要成分可溶性有机碳(Dissolved organic carbon,DOC)和氮(Dissolved organic nitrogen,DON)参与C、N循环过程。为深入揭示全球C、N循环过程机制,在未开展DOC和DON的地区进行相关研究是有必要的。森林土壤(包括枯枝落叶层)DOC、DON动态及调控机理的研究是目前国际上森林生态系统C、N循环研究热点之一。本研究立足于暖温带岷江上游茂县地区人工林植被,对土壤DOC和DON的库容量,季节动态及其与其它养分之间的关系进行了系统研究,旨在了解DOC和DON在该区生态系统中的重要作用,并探讨作为DOM主要来源的叶凋落物对DOC和DON的动态影响,研究有助于更加详细地了解该区生态系统C和N循环过程。本论文主要研究结论如下: 1研究了岷江上游地区两大主要土壤类型(棕壤和褐土)不同植物群落下土壤的DOC和DON含量及特征,结果表明:DOC和DON在两种土壤类型中均有库容量存在,DOC在0-10cm和10-20cm土层的含量幅度分别111.96~159.95 mg kg-1和69.02~100.84 mg kg-1。DON在0-10cm和10-20cm土层的含量幅度分别11.88~23.08 mg kg-1和4.70~10.77 mg kg-1。游离氨基酸在0-10cm和10-20cm土层的含量幅度分别0.84~1.66 mg kg-1和0.39~0.73 mg kg-1。DOC、DON与土壤中的一些养分因子表现出了显著的相关关系,共同反映了土壤的状况和质量,在该区开展DOC和DON的系统研究是有必要的。 2 对油松与连香树林地土壤DOC、DON以及其它化学指标的季节动态进行了研究,结果表明:油松与连香树林地土壤DOC和DON的季节动态变化表现了类似的规律,DOC和DON的含量均以秋季最高。DOC和DON的季节动态变化主要受凋落物生物因素的影响,但其微生物活力的生物因素以及降雨、温度等非生物因素也是控制土壤DOC和DON含量的重要因素。土壤DON在土壤中的行为不同于矿质氮,其季节动态不同于NO3--N和NH4+-N的季节动态,在研究N循环过程中,应考虑DON的变化情况。 3 对油松与连香树林地分解层和表层土壤(0-10cm)氨基酸周转动态进行了研究,结果表明:油松林地和连香树林地均以分解层的氨基酸含量高于矿质表层土壤的含量。每个取样时期,油松林地内各层次的氨基酸含量高于连香树林地内相应层次的含量。两林地各层次无机氮含量均超过了氨基酸的含量,并且室内培养30天后无机N的含量仍然高于氨基酸的含量,所以可以认为该区立地条件下无论是在有机分解层还是矿质土层植物吸收利用的氮素仍是以无机N为主。 4 松林下松针凋落物易于累积,这与松针凋落物分解缓慢有关,从而导致松林内养分周转缓慢。通过用不同性质凋落物和灌丛地土壤构建微生态系统,比较油松、辐射松、连香树、灌丛虎榛子凋落物分解对C、N循环过程的影响,结果显示油松和辐射松针叶凋落物比连香树、虎榛子凋落物分解更慢,减缓了养分循环过程。然而将针叶凋落物与阔叶凋落物混合后,油松和辐射松针叶凋落物的分解加快,C、N元素的循环过程也加速。此结果表明在松林内维持具有高质量凋落物的灌丛植被或在松林内栽植一些阔叶树种如连香树对维持和增进松树人工林的土壤肥力有重大的作用。室内培养的结果还显示添加凋落物后土壤DOC和DON的含量显著增加,表明凋落物是土壤DOM的直接来源。然而不同物种凋落物处理下土壤DOC和DON的含量有所不同,并随时间发生改变。混合凋落物处理下土壤DOC和DON的含量均高于松针凋落物单独处理下土壤DOC和DON的含量。DON是一个主要的水溶性N库,随时间的变化趋势与无机N的变化趋势不同,在土壤N循环过程中起到了中间N库的作用。 As a labile fraction of soil organic matter, dissolved organic matter (DOM) plays a very important role in material cycling of terrestrial ecosystem. The turnover of DOM is now being considered as main components in nutrient cycling. DOM mainly includes dissolved organic carbon (DOC), -nitrogen (DON), -phosphorous (DOP) and –sulfur (DOS). Among these constituents, DOC and DON directly participate in C and N cycling. It is essential to study DOC and DON dynamics and their controlling factors in the areas where no related study has ever been carried out. Study about them can provide data supports on understanding the mechanism of the global C and N cycling. DOC and DON dynamics and their controlling factors have been focused on in the research of C and N cycling of forest ecosystems. Based on forest plantations of Maoxian, Minjiang River in warm temperate zone, soil DOC and DON pool size, their seasonal dynamics, and the correlation between DOC, DON and other nutrients were studied in order to understand the importance of DOC and DON in the study area. Soil DOC and DON dynamics induced by leaf litter decomposition were also studied. The study contributed to comprehensively understanding C and N cycling processes and providing baseline data for including DOC and DON into the indices system of evaluating nutrient conditions. The results were as follows: 1 Several different plant communities under brown soil and Cinnamon soil were chosen as sampling plots. The contents and features of soil DOC and DON were evaluated. The results showed that DOC and DON were present under the two soil types. DOC contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 111.96 mg kg-1to 159.95 mg kg-1, and 69.02 mg kg-1 to 100.84 mg kg-1. DON contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 11.88 mg kg-1to 23.08 mg kg-1, and 4.70 mg kg-1 to 10.77 mg kg-1. Free amino acid contents in the top soil (0-10 cm) and the subsoil (10-20 cm) respectively varied from 0.84 mg kg-1to 1.66 mg kg-1, and 0.39 mg kg-1 to 0.73 mg kg-1. Significant correlations were found between DOC, DON and some nutrient indices, which together reflected soil condition and quality. It was hence essential to study DOC and DON in the study area. 2 Seasonal dynamics of DOC, DON, inorganic N, microbial biomass C and N were studied under Pinus tabulaeformis and Cercidiphyllum japonicum plantation. The results indicated that seasonal dynamics of soil DOC and DON under the two plantations performed similar change pattern, with the highest values in autumn. The seasonal dynamics of soil DOC and DON were mainly influenced by the litterfall. However, biotic factors such as soil microbial activities and abiotic factors such as precipitation and temperature also controlled the dynamics of soil DOC and DON. The seasonal dynamic of DON was different from that of NO3--N and NH4+-N, which showed that the behavioral differences between DON and inorganic nitrogen. And hence, it was proposed to include DON into soil N cycling in the study area. 3 Amino acid dynamics in Oa and topsoil (0-10 cm) under P. tabulaeformis and C. japonicum plantation were studied. The results showed that amino acid content in Oa was significantly higher than that in mineral soil. At each sampling time, significantly higher amino acid contents were found in P. tabulaeformis plantation than in C. japonicum plantation. The content of inorganic nitrogen was much higher than the content of amino acid in each sampling layer at each sampling time. After a 30-days laboratory incubation the content of amino acid was still lower than the content of inorganic nitrogen. The results implicated that the form of N absorbed by plants in these study sites were mainly inorganic nitrogen. 4 Usually needle litter is more resistant to decomposition, which leads to needle litter accumulation in pure coniferous stands and slows down the rate of nutrient circulation. By constructing microcosms with local shrubland soil and containing the four single-species (P. tabulaeformis, P. radiata, C. japonicum, Ostryopsis davidiana) litters, the decomposition rates and related C and N dynamics of needle litters and broadleaved litters during the early stage were compared. The results showed that the decomposition rates of pine needles were lower than those of broadleaved litters, which descended C and N cycling processes. However, the presence of C. japonicum or O. davidiana litter into pine needles increased the decomposition rates of pine needles and also dramatically promoted C and N cycling processes. It should be appropriate for plantation managers to consider C. japonicum as an ameliorative species or remain O. davidiana in pine plantations to improve soil conditions and help maintain soil fertility. The laboratory incubation still showed that DOC and DON contents in all litter-amended treatments were significantly higher than no litter-amended treatment, which proved that litter could be a direct source of DOM in soils. Different species litters induced different soil DOC and DON contents, which correspondingly changed over time. DOC and DON contents in mixed litter treatments were higher than those in pine needle litter treatments. As a major soluble N pool, DON developed a different changing pattern over time compared with inorganic N and played a role of interim N pool in soil N cycling.
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川西亚高山针叶林是四川森林的主体,是长江上游重要的生态屏障。云杉作为亚高山针叶林人工更新的主要树种,已经在该地区形成了大面积的人工纯林。目前,许多云杉人工林分已经进入更新成熟龄,而这些人工林的持续更新却成为日益凸现的问题。探讨这些云杉人工林的自我更新潜力及云杉种子种群更新特点,可为培育后续森林资源提供科学依据。 本文以川西米亚罗亚高山60a云杉人工林为研究对象,并以该区域内相对稳定的植被群落——天然林为对照,采用种子收集器、土壤种子库筛选、室内外种子萌发实验及野外幼苗调查等方法,从异质性微生境的角度研究了种子雨下落之后,不同微生境对种子库、种子萌发、幼苗建立及分布这一前期更新过程的影响,得出如下结果: 1、通过对川西亚高山60a云杉人工林和天然林6年内种子雨雨量、形态特征、散步动态等的持续观测和综合比较可以发现,云杉林结实特点由于林木自身的特征存在着巨大的变动,2002年和2006年两个种子结实大年内,60a人工林种子雨强度分别达到1088.2 ± 52.3粒/m2和704.3 ± 48.9粒/m2,远大于天然林579.9 ± 28.9粒/m2 和507.5± 30.7粒/m2;且云杉林结实质量优于天然林。60a人工林结实量大,种子质量也最好,相对天然林来说对种群的天然更新以及群落的演替都有最大的贡献潜力。应该说,在川西亚高山云杉人工林的天然更新过程中,种源不是影响天然更新的因素。在种子结实大年里,达到更新成熟的云杉人工林有着优于该地区相对稳定植被群落——天然林的种源优势。至少在种子结实大年,种子供应不是该区域人工林天然更新的限制因子。 2、相对于天然林种子库,人工林种子库在种子萌发前能够有较多的有活力种子。虽然这其中有种子雨输入量有差别的因素存在,但两种林分种子库对种子的保存率的不同才是造成这种差异的主要因素。在人工林中,不同地被类型形成的微生境显著地影响了种子库中种子的密度、垂直分布。有地被物存在的微生境能够将种子雨的大部分截留在地被层中,成为幼苗出现的主要场所;同时不同的地被物对种子的保存情况存在显著的差异,苔藓和凋落物层能都较好地保持其中的种子,到种子萌发前,这两种种子库类型能分别为天然更新提供366.1粒/m2和302粒/m2的有效种子。从这点来看,林下地被物上的种子库能够为天然更新萌发阶段提供数量可观的物质基础。 3、种子的萌发和幼苗的定居是天然更新过程中种子库向幼苗库转化的关键环节。总的说来,米亚罗人工林区60a云杉人工林种子向幼苗的转化率十分低下,凋落物、苔藓、草本、裸地四种主要地被物以及天然林内种子/幼苗的转化率分别仅为2.22%、2.14%、0.57%、0.67%、1.05%。这种低的转化率成为云杉林天然更新的限制性因子。但在现有更新条件下,微生境对这一环节仍然显示出十分显著的影响,表现为凋落物和苔藓对现有更新的新幼苗的保存率高于其它类型及天然林。苔藓和凋落物在种子萌发,幼苗保存,以及幼苗分布上都要优于其它地被物类型;另外,微地形对天然更新过程的影响也很显著,凹立地上更适宜于种子的汇集、萌发和定居。 Subalpine coniferous forests dominate most parts of the forested areas in western Sichuan, and they are important ecological barriers in the upper reaches of the Yangtze River. Picea asperata is one of the keystone spruce species for re-afforestation after felling of the natural forests and there have been a total of ca. 13 000 ha of plantations dominate by this species established. Nowadays, many P. asperata plantations have reached reproductive maturity. However, continued regeneration becomes to an important problem in these plantations. Understanding their self-regeneration potential and the regeneration characteristics of seed populations in spruce plantations of these plantations can have some insights on the management of these plantations and the establishment of following forest resources. A subalpine 60a P. asperata plantation distributed in Miyaluo artificial forest area was studied in this paper, at the same time. Synchronously, a 150a natural spruce forest was studied as comparison. Using seed collecting traps, sieving method for soil seed bank, seed germination experiments and seedling investigations in the field, the effects of heterogeneous microsites on early natural regeneration processes after seed rain were studied, which included seed banks, seed germination, seedling establishment and distribution. The main results are as follows. 1. Through a 6-year long term investigation of seed rain intensities, characteristic, dispersal dynamics of 60a P. asperata plantation, we could concluded that the seed setting properties of 60a P. asperata plantation have a great variation for the characteristics trees. In the mast seed year of 2002 and 2006, the seed rain intensities of plantation was 1088.2 ± 52.3 seeds/m2 and 704.3 ± 48.9 seeds/m2 respectively, which were much greater than that of natural spruce forest (579.9 ± 28.9 seeds/m2 in 2002, and 507.5± 30.7 seeds/m2 in 2006). Furthermore, the quality of seed rain in P. asperata plantation was better than that of natural spruce forest. Contrasting with natural spruce forest, 60a P. asperata plantation has a greater potential on natural regeneration of P. asperata population and succession of community for the reason of greater seed rain intensities and better seed quality. We can confirm that seed source was not a limiting factor which influences the natural regeneration progress of P. asperata population distributed in subalpine mountain zone, at least in the mast seed year. 2. Contrasting with natural spruce forest, P. asperata population had more viable seeds in seed bank before the germination. Although the difference of seed rain intensities of two forests has effect on this phenomenon, the difference of seed conservation ability in two forests was the main factor. In the P. asperata plantation, the seed densities and seed vertical distribution pattern were significant effected by the microsites, which posed by different ground cover types. In other word, Microsite with ground covers can obstruct most seeds and keep them in ground cover layer from seed rain, and these ground covers would be the main site for seed occurrence. There was a significant difference about seeds conservation ability among these ground covers. Litter and moss could better conserve P. asperata seeds which distributed in this two covers. Seed banks exist in litter and moss ground cover types could respectively provide 302seed/m2 and 366.1seed/m2 for natural regeneration before the seed germination. From this point of view, we could conclude that ground covers can ensure considerable numbers of seeds for the germination process. 3. Seed germination and seed establishment are key steps that the seeds invert to seedlings in natural generation process. In sum, the seed/seeding transform rate in 60a P. asperata plantation distributed in Miyaluo artificial forest area is very low. the seed/seeding transform rates in litter, moss, herb, soil surface and natural spruce forest were 2.22%、2.14%、0.57%、0.67%、1.05%, respectively. The low transform rate become to a limiting factor of P. asperata natural regeneration process. However, under the existing conditions of natural regeneration, microsit still had significant effect on this transform. The states of Seed germination, new seedling conservation, and older seedling distribution in litter and moss were better than in any other ground cover type or natural spruce forest. In addition, the micro-relief has significant effect on natural regeneration process. Concave site was more suitable for seed collection, seed germination and seedling distribution.
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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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九寨沟湖泊湿地在维持九寨沟的生态平衡中起着重要的作用,在旅游产业的发展下,湿地生态系统及生物多样性面临着较大的威胁。尽管九寨沟湿地具有重要的生态价值,但目前对其研究尚比较薄弱。湿地植物群落和植物地理研究可以为湿地资源的可持续利用和监测保护提供科学依据。作者从2004年8月到2007年11月对九寨沟湿地的植物物种组成、地理分布、优势植物群落的结构、生长动态、湿地土壤种子库进行了调查研究。主要结果如下: 1. 九寨沟湿地物种组成、地理分布特点及湿地植物群落特点 九寨沟湿地共有苔藓植物8科13属16种,维管植物为48科107属199种。九寨沟湿地植物的地理成份较为丰富,维管植物在科级水平上有7种地理分布型(变型),在属级水平上有13种地理分布型(变型), 在种级水平上共有29种地理分布型(变型)。九寨沟湿地植物以温带成份和我国特有成份为主,同时兼有热带、亚热带成份和环极—高山成份。九寨沟湿地植物的分布表现出明显的垂直地带性和水平地带性。湿地植物群落可划分21个群落类型,不同植物群落类型的物种多样性及物种组成存在较大的差异。九寨沟湿地植物的物种多样性和群落多样性以及较高的生产力特征,是维持其湿地生态景观多样性和稳定性的基础。 2. 土壤、水环境、海拔等对湿地植物的分布及生物多样性的影响 九寨沟湿地土壤、水等环境因子存在较大的差异。帕米尔苔草和宽叶香蒲等群落的凋落物较多,土壤有机碳、土壤总磷较高,可能是九寨沟湿地的重要土壤碳库。 九寨沟湿地植物沿水环境梯度的分布规律表现为:沉水植物(轮藻—篦齿眼子菜,水苦荬,杉叶藻)——挺水植物(水木贼,芦苇,宽叶香蒲)——湿生草本(苔草、节节草、披散木贼)——湿生灌木(柳灌丛,小檗灌丛)等。海拔也影响湿地植物的物种组成。 水深对物种多样性有影响,水深与物种丰富度负相关。随着水深的增加,水木贼、芦苇、杉叶藻、宽叶香蒲等群落的物种多样性下降;在长期淹水和季节性淹水的地方,水木贼群落物种多样性存在显著差异。土壤总氮与水木贼群落物种丰富度正相关。 3. 土壤营养元素、水环境对植物生长的影响 水深影响湿地植物生物量的分配。芦苇无性系分株在47 cm水深的环境中单株平均生物量最大;在干滩地中(地面水深0 cm),叶生物量百分比最大,而茎生物量百分比最小,茎的生物量百分比和生长速率随水深的增加而增加;在较干的滩地生境中,开花率、花序的生物量百分比明显大于水较深的生境。 水深与水木贼地上生物量负相关,但水木贼地上生物量在长期淹水和季节性淹水的地方没有显著的差异。在水浅的地方,杉叶藻、水木贼、芦苇等植物群落中,其他伴生物种的生物量占样方总生物量的百分比较大。 土壤有机碳、土壤总氮、土壤总磷等对湿地植物生物量的影响比较大:宽叶香蒲地上生物量与土壤总磷正相关;水木贼地上生物量与土壤总氮正相关;杉叶藻地上生物量与土壤有机碳正相关。 水深、土壤营养成分对湿地植物高度、密度等有影响。水木贼的平均高度在季节性淹水的地方比长期淹水的地方低,平均密度在长期淹水的地方比季节性淹水的地方低;除了5月份,其他观察月份水木贼的密度都与水深负相关,同时与土壤有机碳正相关。另外,芦苇密度与土壤有机碳含量正相关,宽叶香蒲密度与水深负相关,帕米尔苔草高度与土壤有机碳负相关。 4. 优势植物群落的动态变化 在优势植物群落中,优势种的高度、密度、盖度、生物量等在群落中占绝对优势。除五花海,水木贼群落的物种组成、高度、生物量在两年间没有显著的变化。芦苇群落的物种丰富度在近两年有所增加。 湿地植物生长表现为明显的季节动态,生长的峰值大多在7月-8月。优势植物群落的物候与水文周期有关。湿地植物群落的物种组成和密度,可以作为对湿地监测和保护的生物指示。 5. 九寨沟湿地土壤种子库特征及其在湿地生物多样性恢复中的作用 水深和现存植被物种丰富度可以解释湿地土壤种子库的变化。水深可以解释表层物种丰富度45%的变化。现存植被物种丰富度可以分别解释10 cm土层、2-5 cm土层及5-10 cm土层土壤种子库45%、48%和25%的变化。 湿地土壤种子库的密度为0-15945粒m-2, 种子库中共发现23个物种。现存植被优势物种和种子库优势物种不同。各层土壤种子库密度和物种丰富度并不存在显著的差异,但第二层土壤种子库密度最大。海拔、现存植被优势种盖度、土壤总磷、土壤总氮、土壤有机碳对湿地土壤种子库的密度和垂直结构没有影响。土壤种子库物种丰富度小于地上植被物种丰富度。湿地土壤种子库与地上植被的相关性不大。在浅水区域,湿地土壤种子库在湿地植被恢复中有一定作用。但在深水区域,保护现存植被更重要。 The lakeshore wetlands are valuable ecological units of the Jiuzhaigou lakes. Pressure for travel industry development pose a continuing and severe threat to the biodiversity-support function of the wetland system. Despite the ecological importance of wetlands in Jiuzhaigou, they are so far poorly studied. Both general plant communties and biogeographical studies are needed in order to attain basis for sustainable use the wetland resources and adequate protection of these areas. The present study was undertaken to examine aquatic plants distribution and the species compositon, structure and growth dynamics of their communities with variations of environmental factors along altitudes, water depth and soil properities gradients in Jiuzhaigou. Analysis of field survey data collected during August 2004 and November 2007 in lakeshore wetlands in Jiuzhaigou National Nature Reserve, Sichuan, China. The results were as following: (i) Species composition and biogeography in wetland vegetation 8 families, 13 genus, 16 species of moss and 48 families, 107 genus and 199 species of vascular plants in Jiuzhaigou wetlands were found. The floristic compositions were abundunt. Ten geographical distribution types at family level, 13 geographical distributions types at generic level and 29 geographical distribution types at specific level in vascular plants were found. Most species in Jiuzhaigou wetlands are temperate elements and Chinese endemic elements, with a few of tropical and subtropical and some circumarctic elements. And the plant distributions show clear vertical and horizontal patterns. There were 21 major wetland plant community types. Species composition and species richness in different plant communities are different. The species diversity and plant community diversity and their high biomass are the basis for the diversity and stability of wetland landscapes in Jiuzhaigou. (ii) Water depth, soil nutrients and altitudes influence on the species diversity and plant distribution. Total phosphorous and organic cabon in soil were higher in C. pamiernensis and T. latifolia communities, where are important cabon reservoirs in Jiuzhaigou wetlands. Along gradients of water depth, among populations of the dominant plant species present: submerged macrophytes (Chara vulgaris, Potagemonton pectinatus, Veronica anagalis-aquatica,Hippuris vulgaris), emergent macrophytes (Equisetum fluviatile, Phragamites australis, Typha latifolia), helophytes (Carex pamirensis )and shrubs (Salix sp., Berberis sp. ). Altitudes influence on the assemblage of plant communities. Water depth negatively correlated with species richness. Specie richness showed differences between permanently flooded sites and seasonally flooded sites in E. fluvatile communities. And total nitrogen in soil was negatively correlated with species richness in E. fluviatile communities. Altitudes show no significant influence on species richness, but in fact, through our analyses, they do have influence on the assemblage of wetland plants. (iii) Water depth, soil nutrients influence on the plant growth Water depth influences the biomass allocation in Phragmities australis. The average aboveground biomass of a single ramet (4.2 g) was the largest in the habitat with water level 47 cm above the soil surface. At the habitat with water level under soil surface 15 cm (-15 cm), the leaf biomass percentage (of the total ramet biomass) was the largest (46.1%), and the height and percentage of ramose ramets ( with branches on stem )(of the total ramets in a plot) were found obviously different. The deeper in water, the larger the biomass percentage and growth rate of stems were. The flowering rate and biomass of panicles were greater in shallow water than those in deep water. Water depth negatively correlated with aboveground biomass of E. fluviatile. However, above-ground biomass of E. fluviatile showed no significant difference between permanently flooded sites and seasonally flooded sites. But in shallow water, more biomasses of accompanying species were found in dominant plant communities such as H. vulgaris communities, E. fluviatile communities and P. australis communities. Water depth, soil nutrients influence on shoot density and shoot length of wetland plants. The shoot density of E. fluviatile was correlated to water depth in all growth months. Annual average density was significantly lower at permanently flooded sites than at seasonally flooded sites. But the annual average shoot length was significantly lower at seasonally flooded sites than at permanently flooded sites. (iv) Growth dynamics of dominant communities in Jiuzhaigou wetland The shoot length and shoot density, coverage and biomass of domiant species were dominated in plant communities. The species composition increased in P. australis communities in recent two years. The species richness in E. fluviatile communities showed no difference between 2005 and 2007. The above-ground biomass and shoot density in Five-flower Lake from July 2005 to July 2007 were significantly different, while in other sites, the differences were not significant. Shoot height, shoot density and above-ground biomass showed significant seasonal changes in all sites. Growth dynamics correlated with the cycle of water levels in lakes. Most plants growth parameters peaked at July or August. The biomass of T. latifolia peaked in August. But the shoot length of T. latifolia in deeper water peaked in July. The shoot length of E. fluviatile increased significantly from May to August except in seasonally flooded sites in Arrow-bamboo Lake. The species composition of communities and shoot density can be used as bioindicators in Jiuzhaigou wetland. (v) Soil seed bank in Jiuzhaigou wetland and its role in vegetation restoration Seed density in all soil layer samples was negatively correlated to water depth. Water depth can explain 45% variance of species richness in surface layer in sediment. Species richness in extant vegetation can explain 45%, 48%, 25% variance of species richness in total 10 cm and in 2-5 cm and 5-10 cm layer sediment respectively. Mean seed densities in wetlands ranged from 0 to 15945 m–2. A total of 23 species germinated in seed bank. The dominant species in seed bank and extant vegetation showed great difference. The total number of species and seedlings that germinated in different layers was not significantly different. But the second layer had the greatest seed density. In shallow water, seed bank can contribute to vegetation restoration, while in deeper water, protection of extant vegetation may be a better strategy.
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LUCC是全球变化研究的核心主题之一,也是社会经济可持续发展的关键问题。改革开放后四川的社会经济发展非常快,在各种因素的驱动下,土地利用/覆盖发生了深刻变化。目前四川省缺乏基于实际调查数据的、全域性的、具有连续时间序列的LUCC和驱动力分析及土地可持续利用研究成果,这对我们从全局上把握全省土地利用现状、发展变化趋势,利用土地政策参与宏观调控,实现长期可持续发展目标,建设资源节约型、环境友好型社会极为不利。本研究针对这一问题,选取全川八大土地利用类型作为研究对象,研究了全省1996年到2006年的土地利用/覆盖格局和变化情况,分析了不同尺度的驱动因素,对全省农用地和建设用地的集约利用状况、潜力进行了分析评价,并提出相应的对策措施。 1.1996年-2006年10年来整个省域的土地利用/覆盖格局变化。 (1)1996年-2006年全省的土地利用/覆盖格局 1996年,全省是一个以农用地为主的土地利用/覆盖格局,林地和牧草地属于优势覆盖类型(合占69.17%),居民点及工矿用地和交通用地合占只有3%左右。 2000年的LUCC格局较为明显的特点是耕地所占比重下降0.4个百分点,水域和未利用土地所占比重有所下降,牧草地保持不变,其余地类所占比重有所上升。 与2000年相比,2004年林草地的优势格局进一步得到强化(合占比重达到70.23%)。耕地面积占幅员面积的比重下降0.83个百分点,略有下降的有未利用土地、水域和牧草地。值得关注的是在“退耕还林还草”的大背景下,牧草地占幅员面积的比重下降0.04个百分点。 到2006年,仍为林草地为主导优势的格局,二者合占上升0.15%。在城市化快速推进的背景下,居民点及工矿用地中的城市用地和建制镇用地占比重超过15%,农村居民点占比重降至76%。交通用地中农村道路占比重降至57.8%,公路用地占比重升至37.5%。五个地貌区的土地利用/覆盖格局与全省的变化基本一致。值得关注的是盆西平原区的交通用地上升幅度和盆地丘陵区的未利用土地的开发利用力度明显大于其它地貌区。 (2)1996-2006年10年间土地利用/覆盖格局的变化 1996-2000年4年间,耕地、水域和未利用地三个地类下降,年均减少0.75、0.19和0.32个百分点。其中耕地年均减少49229.0公顷,约一半流向林地,13.77%流向园地,约20%流向建设用地。另外5个地类面积增长,增长绝对量最大的是林地,年均增长40063.7公顷,交通用地增幅最大,4年年均增长达1.95%。 2001-2004年是西部大开发逐步推进、“退耕还林还草”项目全面展开和土地整理深入实施的关键期,LUCC更为深刻。耕地、未利用地、水域和牧草地四个地类面积下降,其余地类按增长幅度依次是园地、交通用地、居民点及工矿用地和林地。耕地加速下降,年均降幅达到1.59%,其减少去向主要是林地(占66.75%)和园地(占19.84%),其增加来源主要是未利用地、园地和水域。交通用地的增幅最大,为3.96%,其增加主要来源于耕地、未利用土地和林地,分别占49.96%、16.63%和13.09%。居民点及工矿用地增长幅度为3.12%。 从1996年到2006年的10年间,耕地、未利用地、水域和牧草地下降幅度分别为10.36%、3.61%、1.34%和0.26%。园地增幅达23.61%。绝对面积增长最大的则是林地,达630733.3公顷。交通用地和居民点及工矿用地增幅也较大,分别为15.00%和9.31%。 10年间年均总变化量为310326.6公顷,2000年-2004年之间变化最大(为356865.8公顷),高于平均变化量,而1996-2000年间和2004-2006年间都小于平均变化量。 (3)10年间不同地貌区的LUCC变化 盆西平原区的特点是园地大幅上升达77%,居民点及工矿用地和交通用地也大幅上升,耕地、未利用地下降幅度大,该区耕地、水域、未利用地的减少强度和园地、居民点及工矿用地、交通用地的上升强度均居五区第一;盆地丘陵区的特点是牧草地下降幅度大,为-36.89%,交通用地、园地和林地上升幅度较大,该区耕地减少、未利用地减少、林地增加、居民点及工矿用地和交通用地增加的变化强度均居五区相应地类增减的第二位;盆周山地区的特点是耕地减少较多,交通用地和园地增长较大,该区林地变化强度居各区第一位,牧草地和水域变化强度居各区第二位,耕地、居民点及工矿用地和未利用地居各区第三位;川西南山地区的特点是园地、耕地、交通用地和居民点及工矿用地变化幅度大,另外四个地类变化较小。该区减少的牧草地占全省牧草地减少的97.91%,变化强度居各个地貌区的第一位,园地相对变化强度居五区的第二位;川西北高山高原区的特点是耕地大幅下降、园地大幅上升,交通用地升幅也较大,其余地类变化不大。值得注意的是,该区牧草地和水域面积增加,与全省该地类的变化相反。其余地类的相对变化强度均是五个地貌区中最小的。 用变化强度分值考量变化强度,盆西平原区的变化强度最大,盆地丘陵区和盆周山地区的变化强度相当,川西北高山高原区的变化强度则要小得多。 (4)1996年及2006年全省土地利用/覆盖格局的景观生态学分析 全省是以自然景观占优势(占约70%)、农业景观为补充、建设用地景观居于从属地位的土地利用景观格局。景观多样性和均匀度不高。到2006年,全省总的景观格局并无大的改变。总体情况是随着时间的推移和人类活动的加强,区内景观优势度上升、多样性和均匀度变小。但斑块数减少,斑块面积和斑块孔隙度有所增大。斑块的形状指数和分维数均有所下降,表明受人为干扰有加剧的趋势。反映景观格局结构的破碎度指数有轻微下降。景观指数的变化表明全省土地利用有缓慢集中、规模聚集的趋势。 (5)三大生态建设工程对土地利用/覆盖变化的影响 1996-2006年间LUCC与三大生态建设工程实施的耦合分析,发现退耕工程对耕地、林地、牧草地等地类覆盖变化的影响最大,天保工程次之,长防工程最小。 2.四川省LUCC驱动力分析 (1)总体分析: 从整体上分析,人为因素对区域整体LUCC的影响从1996年的63.32%增加到2006年的66.99%,变得日益强烈。同时人为因素影响强度表现出明显的区域差异,地势平缓、经济区位条件好的区域其人为影响强度明显较高。 政策体制转变下的经济高速增长、快速的城市化、工业化过程和生态建设是四川省LUCC宏观尺度的驱动因素。区域的LUCC主要受到了由内向外(从城市到乡村)和由外向内(从山顶向平地)两种作用力的共同推动。局部尺度上,如距离交通线、水利线、中心城市的远近,地形凸起、大型独立项目落址、重污染项目的阻隔等,甚至一些乡规民俗等因素也会成为LUCC的驱动影响因素。在较小的尺度上,人类个体行为选择对LUCC的影响也是存在的。 根据驱动因子的特性作者将其划分为驱变、阻变、良性、惰性因子等类型。 (2)分地貌区的驱动因子分析 各地貌区都存在城市化、工业化、生态工程实施、自然灾害等驱动因子,但主次不一。对于盆西平原和盆地丘陵区,城市化、工业化是前两位的因子,而对另外三个地貌区,生态工程实施和产业结构调整则成为第第一、二位的驱动因子。 (3)分地类的驱动因子分析(以坡耕地为例) 分坡度的耕地变化分析发现,耕地减少主要集中在2°以下的平地、15°-25°和25°以上三个坡度级,是其它坡度级耕地减幅的三倍左右。这表明耕地减少受城市化进程和“退耕还林还草”工程驱动影响尤为巨大。 3.土地利用格局优化、集约利用评价和可持续利用及对策研究 (1)土地利用格局优化的战略选择及调整预测 土地利用格局调整的战略是农业生产用地、建设用地和生态及其他用地占幅员的比重分别稳定在13%、7%和80%左右,重点是三大类别内部二级和三级地类的合理调整。 (2)全省土地集约利用评价 全省农用地利用集约度为0.46,总体上集约度不高,处于较适度利用阶段。建设用地利用集约度为0.38,处于较适度利用阶段。集约利用提升空间较大。 农用地的潜力主要在于加强土地保育、完善利用制度、提高单产。城市建设用地的包括存量潜力、强度潜力、结构潜力,空间很大。农村居民点整理潜力可以逐步挖掘。 (3)新增建设用地集约利用的统筹安排 据测算,到2020年,四川省城市建设用地需求量在463850-492360hm2之间,城镇各业新增建设用地规模为361276.79hm2,占用耕地200565.94 hm2。2004-2020年间四川省农村居民点整理潜力33.86万hm2。农村居民点建设用地需求量为70.57万公顷。 (4)土地集约利用措施与坡耕地可持续利用战略 提出了土地集约利用的措施。在对坡耕地生态系统结构与功能分析的基础上,提出坡耕地可持续利用战略与生态恢复战略,并从技术和政策层面提出了坡耕地合理利用和生态退耕的措施和建议。 LUCC is one of the key questions of global change and sustainable development of society. After the opening and reform of China, the society and economy of Sichuan Province developed very fast ,the land-use/cover changed very strong droved by many factors .But nowadays we have no constant spatial-temporal study and driving force analysis about the whole province based on investigation. And it is lack of land sustainable utilization study based on correlative study. So we choose all the land resource in Sichuan, combine RS and GIS and field investigation, and take statistic-mathematic means and system analysis, to study the LUCC patterns and different scale driving force of different physiognomy regions, land cover types and periods; to analyze the current situation and potential of land resource intensive utilization, and gave out corresponding measurements. We found that forest and grassland are the dominant cover types of Sichuan provincial land –use/cover pattern, and becoming more and more stronger from 1996 to 2006,the natural landscape is the metric and occupy 70%,the diversity and evenness index are not high; the totally change quantity from 2000 to 2004 is the biggest; cultivated land especially steep cultivated land ,garden plot, forestry land ,settlement and industry land and traffic land changed relative stronger; among five physiognomy regions ,the changing intensity of PEN XI PING YUAN QU is the biggest, CHUAN XI BEI GAO SHAN GAO QU is smallest; under the background of policy system changing, the fast developing of economy, fast urbanization and industrialization and ecology construction are the macro-scale driving force of Sichuan provincial LUCC; to compare the impacts of “TUI GENG GONG CHENG” on LUCC especially to cultivated land ,forestry land and grassland is strongest, “TIAN BAO GONG CHENG ” is stronger,“ HANG FANG GONG CHENG” is smallest; the intensive utilization level of farmland and construction land of whole province is relative moderation, there is huge potential to excavate and fulfill the increasing demand of construction land;we must take synthetic measurements to accelerate the sustainable utilization of land resource, including administrative, economical ,technological and ecological policies.
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混农季节性放牧(agropastoral transhumance)通过作物种植和畜牧生产相结合的方式对不同海拔高度带上的资源进行相互补充利用,在亚洲兴都库什地区、青藏高原、横断山、东部及南部非洲、南美安第斯地区等具有悠久的历史。这种传统的生计系统几千年以来一直是居住在该地区的人类社会和自然生态系统相互作用的主要形式之一。这种传统的资源利用方式与山地自然植被以及特殊的山地人类文化和社会特征具有密切的协同演变关系。认识和理解这一关系,是山地生态学和人类学的核心科学问题之一。近年来,山地生态系统的多重功能性及动态演变对山区社会经济可持续发展的重要意义受到人们的不断关注。本文通过对云南省德钦县的12个自然村的混农季节性放牧以及对云南德钦、四川壤塘等山地植被格局特别是高海拔地带植被格局的的详细调查,探讨青藏高原东缘地区混农季节性放牧的主要特征、系统构成及相互关系,及其在全球变化、经济全球化和市场化及现代化过程中的变化趋势,分析混农季节性放牧与高山林线格局及生态系统的互动关系,旨在探讨山地地区人类活动与自然生态系统之间的互动关系,从而为山区社会经济可持续发展、环境建设和生物多样性保护等国家战略提供理论依据。 调查结果表明,混农季节性放牧是一种适应青藏高原东部高山峡谷地区环境因子及自然资源呈明显的垂直分布、资源数量稀少而时空分布异质性极高的生存环境的一种传统经济形式。这种传统的畜牧业的主要生产目的仍然是提供当地基本生存所需的产品,饲养牲口的种类和数量取决于农户的当地需求并且受资源的限制,因而维持在比较低的水平的。分布在不同海拔高度的放牧资源在一年中被牲口利用的时间也不同,互为补充,共同构成混农季节性放牧的资源基础。根据各社区永久居住点的位置和该村的土地资源特别是牧草地资源的分布范围,牲口迁移的距离和格局有较大的差异。。天然牧场仍然是最主要的畜牧业生产资源。混农季节性放牧中的农业系统和牧业系统互为补充,共同构成调查地区完整的的生计系统,农耕活动为放牧活动提供精饲料如粮食等和冬季饲料如秸秆, 其数量往往成为家庭畜牧业生产规模的主要决定因子之一。 通过对牲口数量和结构、牲口的时空迁移格局、牧业活动在整个经济活动中的相对重要性以及牧业活动和作物种植的关系方面的研究分析,混农季节性放牧在近几十年发生了深刻的变化。主要表现在牲口数量总体下降,牲口组成发生变化,牲口移动性降低、牧业活动的经济重要性下降以及牧业活动和种植活动之间的相互依存度降低等。上述变化的根本驱动力是发生在当地、地区及全球尺度上的环境、政治、社会经济、技术和文化等的变化,从而造成当地群众畜牧生产目标、土地利用和劳动力的分布等发生了变化。当地生计系统发生的改变可能会带来对方面而深刻的政治、社会经济、文化和生态影响。 混农季节性放牧这种古老的传统生计策略面临着许多挑战,如冬季饲料短缺、草场退化、缺乏市场竞争力、经济重要性降低、对年轻人缺乏吸引力、国家缺乏专门的政策指导等。与此同时,经济全球化、市场经济、新技术的应用、替代生计机会的增加、国家对于山地生态系统的作用的重新定位等也为传统生计系统转型、实现社会与生态共赢创造了机遇。 混农季节性放牧活动对亚高山及树线交错带生态系统系统的互动方式主要体现在以下几个方面:(1)牲口啃食、践踏等影响森林群落更新,改变森林群落的组成和结构,从而影响森林群落的演替进程和植被格局。林线边缘是搭建夏棚的首选地点,因此林线及树线交错地带就成了牲口活动的主要场所之一;(2)利用火烧开辟、维持和改良高山牧场; 3)在亚高山火灾迹地的放牧活动能够阻止火烧迹地的顺向演替; 4)牧民在林线附近获取建材和薪材等活动影响高山林线附近森林的结构和功能。 在调查区域,梅里雪山、白马雪山、甲午雪山的林线海拔高度在4200-4300m之间; 四川雅江、理塘一线,林线位置多在4300-4400m;四川壤塘二林场一带的林线主体在4100-4200m,在个别地区达到4300m; 在贡嘎山的南坡和东坡一带,林线位置在3600-3700m;而在四川松潘一带,林线位置主体在3700-3800米左右。树线高度的分布趋势和林线一致。混农季节性放牧及其有关人类利用活动使研究地区很多地方高山林线降低、树线交错带宽变窄或消失。在研究地区,总体情况是,阳坡和半阳坡(南坡、西南坡等)的林线和树线比阴坡和半阴坡(北坡、东北坡等)低,变化幅度达20-200m。这种差异主要是为了开辟牧场而人为清除了南向坡自然林线及其以上的植被从而使林线位置下降所致。在南坡自然林线保留得比较好的地方,林线和树线依然可以达到甚至超过北坡林线和树线的高度。放牧活动抑制了高山林线带火烧迹地的天然更新,从而使林线位置保持在目前的位置。 放牧活动对高山林线带森林群落更新的影响是显著的。自然林线内的乔木个体密度特别是新生苗和幼苗的密度大大高于非自然林线。没有放牧的自然林线及树线交错带内的I级个体(新生苗)密度达到725-2917株/公顷,而与之相对的处理样地内I级个体的密度只有0-228株/公顷;II级个体(高度10-50cm)也表现出类似的趋势,在没有放牧的自然林线及树线交错带样方内,其密度达到550-5208株/,而在放牧处理样方内只有14-321株/公顷。在非自然林线带样地内,在有正常放牧的样地内,完全缺乏I级个体。 从相对比例来看,没有放牧的样方内的I、II级个体在全部个体中所占的比例显著高于有放牧活动的样方。放牧使林线交错带的乔木幼苗数量显著减少,从而影响林线及树线交错带森林群落的天然更新过程。林线和树线交错带的灌木对乔木幼苗具有重要的保护作用,能够为树线树种如冷杉等幼苗的定居体提供有利的微气候环境,同时保护苗免受牲口的啃食和践踏。火烧以后接着进行放牧能够100%地抑制高山林线带的幼苗更新。 高山牧场放牧强度降低、使用时间缩短而低海拔地带放牧强度增加是研究地区混农季节性放牧系统的一个显著变化。这种变化也必然会引起各海拔带上的生态系统的变化。放牧强度的降低、生产性用火的停止将导致原来通过人工火烧而降低并通过进一步的火烧和放牧活动来维持的林线及其以上地带的灌木盖度和高度的增加,从而为林线森林群落的扩张创造条件。 青藏高原东部高山峡谷地区是我国重要的山地生态系统,在我国的生物多样性保护、生态环境建设、社会经济可持续发展战略中具有举足轻重的作用。正确认识人类特别是当地传统的生计系统与生态环境系统的互动关系是实现上述战略目标的前提。决策者必须以综合、系统的的视角协调促进社会经济可持续发展、保护生物及文化多样性和维持人、牲口和生态系统之间的平衡的多重目标。 Agropastoral transhumance, which makes a complementary exploitation of the natural resources at different altitudinal belts through a combination of migratory animal husbandry and crop cultivation, has a long history in Hindu-Kush Himalaya, Tibet Plateau, Hengduan Ranges, eastern and southern Africa and the Andes region of south America.For millennia, this traditional livelihood strategy has been one of the main forms of interaction between human societies inhabiting in these regions and their natural ecocystems. A close co-evolutionary relationship has been developed between this indigenous resources management systems and the mountain vegetation systems on the one hand and a unique set of cultural values and social features on the other. Understanding this relationship has been one of the core scientific issues in mountain ecology and anthropology. In recent years, the importance of the multiple functions of the mountain ecosystems and their dynamic changes in the sustainable socio-economic development of the mountain regions has gained increasing attention. This paper, which is based on a detailed study on the agropastoral practices of the 12 natural villages in Deqin County of Yunnan, and the mountainnn vegetation patterns in Deqin of Yunnan and Rangtang County of Sichuan, intends to reveal the major characteristics, system composition and the inter-relations of the subsystems of the agropastoral transhumance in Eastern Tibetan Plateau as well as the trends of changes of the system within the context of global changes, economic globalization and modernity process of China and analyze the relations between agropastoral transhumance and alpine ecosystem, ao as to understand the interactions between human activities and natural ecosystems of the mountains and provide theoretical basis for the national strategies in eocioeconomic development, environmental reconstruction and biodiversity conservation in the mountain regions. Results of the survey indicate that agropastoral transhumance in the investigated area is a traditional economic form that is highly adapted to the eastern Tibet Plateau where the topography features high peaks and deep gorges and where the highly variable environmental parameters and scanty natural resources exhibit a distinct vertical spectrum of distribution and great temporal and spatial heterogeneity. The main objective of pastoral management is still aimed at the production of basic goods and services of local people and thus the type and size of animals raised for each household mainly depend on local needs and are limited by the availability of natural resources. The scale of production is relatively low. Pastoral resources at different altidudinal belts are complementarily used at different seasons of a year and thus form the resources basis for agropastoral production of the study area. Migration distances and patterns vary with the location of the permanent settlements, the elevational distribution range of the resources of the villages concerned. Natural pastures (rangelands) are the main fodder resources and sumplement feedings only account for less than 5% of the total fodder consumption. Crop cultivation and pastoral activities support each other to form a complete livelihood system. The ability of the farmig lands (crop cultivation) to provide the pastoral activities with concentrates and sumplements often becomes a main factor limiting the scale of livestock production at household level. Agropastoral transhumance is experiencing drastic changes in recent decades as is reflected in the size and composition of animals, the seasonal migration pattern, the relative importance of pastoralism in the household economy and the interplays of agricultural and pastoral elements of the system. In general, there is a decline in animal population and mobility, a shift in animal composition to meet new needs arising from changed macro-economic situation, a decrease in the relative importance in the household economy and an increasing decoupling of agro&pastoral relations. The fundamental divers of these changes can be traced to environmental, social, economic, technological and cultural changes from local to global levels and such changes have further caused local changes in livestock management objectives, land use and distribution of labor forces. Changes in local livelihood systems could have profound political, socioeconomic, cultural and ecological conseuquences. Agropastoral transhumance, as an age-old traditional livelihood strategy, is facing multifacet challenges, such as winter fodder shortage, rangeland degradation, lack of market competitiveness, decrease in economic importance, lack of appreciation among the young generation and adequate policies from the government. At the same time, economic globalization, market economy, intrdoctution of new technologies, increase of alternative income generating opportunities and the national re-oreitation of policies on mountain ecosystems have all brought about new opportunities for the transformation of the traditional livelihood system and the synchronized development of local society and the environment. Agropastoral transhumance interacts with the ecosystems at the timberline and treeline ecotone mainly through the following aspects: 1)Animal browsing and stamping affect the regeneration process of the forest communities and alters the composition and structure of the forest which in turn affect the succession process and vegetation pattern of the forest communities. Forest edges are the priority locations for summer houses and therefore the timeline and treeline area becomes the major venues of aninal activities; (2)herders create, maintain and improve pastures through burning that remove the forest communities at the timeline and treeline ecotone; 3)immediate grazing on the fire sites can significantly prevent the fire sites from perogressive succession; and 4)herders harvesting of construction timber and firewoods affects the structure and functions of the forest communities at the timberline and treeline zone. Timberline position in the survey region shows geographical variations. It is around 4200-4300m in Meilixueshan, Baimaxueshan and Jiawuxueshan in Northwest of Yunnan and rises to 4300-4400m in Yajiang County and Litang County of Sichuan. In Rangtang of Sichuan, it is between 4100-4200m, though reaching 4300m in localized sites. In the southern and eastern slopes of Gongga Mountain, the timberline is only between 3600m and 3700m and in Songpan County at the upper reach of the Minjiang River the timberline is around 3700-3800m.Treeline pattern follows similar trend. In many places, agropastoral transhumance and related human activities have lowered the timberline and treeline and narrowed or removed the treeline ecotone. In the area of survey, generally speaking, timberlines and treelines are lower on the southern slopes than on the northern slopes, with a difference between 20 and 200m. This is mainly because that the use of fires to crerate pastures has removed the forest vegetation at the previous timberline and above. In fact, in many places, well-preserved forests on the south slopes have even high timberline position that the corresponding northern slopes. At subalpine zone, grazing activities could have prohibited the natural regeneration of many forest fire sites and maintained the forest position at the present level. Grazing has a significant impact on the regernation process of forest communities at the timberline zone. Natural timberline and treeline ecotone has much higher density of treeline species individuals especially the emergents and seedlings than the timberlines that are maintained by human activities. In natural timberline and treelien ecotone without grazing interference, the density of the I Class seedlings (less than 10cm in height) ranges 725-2917 /hm2; while that in the treatment plots (with grazing disturbance) is only 0-228//hm2;II Class seedlings (10-50cm)exhibit similar density trends, reaching 550-5208//hm2 in natural timberline without grazing but only 14-321//hm2 in the plots with grazing treatment. In the man-created timberlines, there is no I Class seedling at all in plots with normal grazing activities. In relative terms, in plots without grazing activities, the propotion of I Class and II Class seedlings is much higher than that in plots with grazing. Grazing activities have significantly reduced the number of seedlings in the timberline ane treeline ecotone, and thus affect the natural regeneration process of the forests. Shrubs at the timberline and treeline ecotone can effectively protect the seedlings from severe climate and animal tramping, thus increasing the survival rate of the seedlings. Grazing following fires can completely inhibit forest regeneration process at timberline. Changes in agropastoral transhumance will have great impact on the timberline and treeline pattern of the studied area. The decrease in grazing intensity on alpine pastrues and the cessation of the use of fires will result an increase in the cover and height of shrubs above the present human-maintained treeline, which will create further condition for the expansion of timberline forest communities. Eastern Tibet Plateau harbors some most important mountain ecosystems of China that are of vital importance to the country’s strategy in biodiversity conservation, environmental construction and sustainable sociaoeconomic development. A proper knowledge of the interactions between traditional livelihood systems and the ecosystems in the region is a precondition to the realization of the above strategic goals. Therefore, the decision-makers must have a holistic and systemic perspective so as to integrate the multiple objectives of promoting sustainable socioeconomic development, conserving biological and cultural diversity and maintaining the balances among people, animal population and the ecosystems.
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随着工业化的发展,大气中二氧化碳的浓度(CO2)预测从现在的平均350μmol·mol-1升高到2030年的570μmol·mol-1,其增温作用将持续多个世纪。植被在大气二氧化碳减排以及调控区域水热状况过程中起重要作用,而其机理过程目前十分不清楚。本实验应用自控、封闭、独立生长室,研究了CO2浓度和温度升高对红桦根、茎、叶和枝可溶性蛋白含量和分配的影响,从蛋白水平上来解释川西亚地区的建群种-红桦对CO2升高和温度升高及其交互作用的响应规律,为全球气候变暖川西亚高山的植被保护和恢复提供理论依据。研究结果表明: 1. CO2浓度升高增加了可溶性蛋白的总量,改变了可溶性蛋白分配模式,即,可溶性蛋白分配到根的比例增加,分配到茎、枝、叶的比例减少。可能意味:在CO2浓度升高条件下,红桦根系的生长和营养物质吸收功能将会增强。 2. CO2浓度升高增加了根和茎的清蛋白含量,降低了叶片的清蛋白含量,叶片的球蛋白含量、醇溶蛋白含量和谷蛋白含量均增加。表明CO2浓度升高增加了清蛋白在根中积累,球蛋白、醇溶蛋白和谷蛋白大量在叶片中积累;前人研究所指出的CO2浓度升高使植物叶片可溶性蛋白的含量降低可能仅仅是由于清蛋白含量的降低造成的。 3. 温度升高使红桦幼苗整株所含可溶性蛋白总量增加,但可溶性蛋白总量的分配因红桦幼苗器官的不同而异。温度升高下根、茎、叶和枝的分配量分别占总可溶性蛋白的27.74%、35.57%、23.00%、13.68%,即茎>根>叶>枝。对照的根茎叶枝的分配量分别占总可溶性蛋白的21.01%、41.41%、23.08%、14.50%,即茎>叶>根>枝。表明温度升高使可溶性蛋白分配到根的比例增加,有利于根的可溶性蛋白的积累,增强了根吸收水分和矿质营养的能力,从而有利于根系的生长。 4. 温度升高处理下清蛋白和球蛋白在根中含量升高,在茎、叶和枝中含量下降,但没有达到显著水平;醇溶蛋白在根和叶中含量显著增加;谷蛋白在茎中的含量显著降低。表明温度升高增加清蛋白和球蛋白在红桦幼苗根部的积累,也有利于根和叶醇溶蛋白的积累,但不利于谷蛋白在茎的积累;温度升高条件下叶片可溶性蛋白升高是醇溶蛋白在叶片中积累的结果。 5. CO2浓度和温度同时升高条件下红桦幼苗的可溶性蛋白总量增加很少,只有分配到茎的可溶性蛋白比例增加,并且对可溶性蛋白分配规律没有影响。CO2和温度同时升高下红桦幼苗枝的可溶性蛋白含量的降低是可溶性蛋白总量的降低而不是碳水化合物稀释的结果,并且CO2和温度同时升高对红桦幼苗的生长没有明显的促进作用。 6. CO2和温度同时升高处理对可溶性蛋白含量有显著影响。清蛋白含量在根、茎、叶和枝中均降低,球蛋白含量在根中显著降低,醇溶蛋白含量在根、茎、叶和枝中均降低,谷蛋白含量在根中显著降低。表明CO2浓度和温度同时升高对根的影响显著,即降低了根的可溶性蛋白含量,可能导致根的吸收能力下降。 7. 因此,CO2和温度同时升高对可溶性蛋白影响不能简单地通过CO2和温度单因子影响机理来解释。 It is well known that atmospheric CO2 concentration and temperature are increasing as a consequence of human activities. Atmospheric CO2 concentration are predicted to increase from 350μmol·mol-1 now to 570μmol·mol-1 2030. And temperature will continue to increase for several centuries as a result of CO2 enrichment. Vegetation play a key role in reducing atmospheric CO2 and adapting and controlling warter and energy process in a certain region, while the underlying mechanism are not clear, yet. Betula albo-sinensis, as the dominating tree species of subalpine dark coniferous forest in west Sichuan province, play an important role in determing structure and function of forest ecosystem. In our study, effects of elevated atmospheric CO2 concentration (ambient±350±25μmol·mol-1), increased temperature (ambient±2.0±0.5℃) and their combination on contents and allocation of soluble protein were studied in independent and enclosed-top chamber system under high-frigid conditions. Chambers with ambient CO2 concentration and temperature are taken as control. The results are as the following, 1) Elevated atmospheric CO2 increased the accumulation of total weight of soluble protein in whole plant and changed allocation of soluble protein in red birch by increasing its allocation to roots and reducing its allocation to stem. This caused much more accumulation of soluble protein in roots which might help to prompt growth, development and nutrient absorption ability of roots. 2) Treatment EC increased content of albumin in roots and stems, reduced the content of albumin in leaves, and increased the content of globulin, promalin and glutenin in leaves. That is to say EC increased the accumulation of albumin in roots and accumulation of globulin, promalin and glutenin in leaves. The reduced soluble protein contents in plant leaves by EC, as reported by former researchers, are mainly resulted from the reduced content of albumin in leaves. 3) Elevated temperature increased the total of soluble proteins, but its allocation was dependent on organs. In treatment ET, roots, stems, leaves and branches take 27.74%, 35.57%, 23.00% and 13.68% of total weight of soluble protein. In treatment CK, roots, stems, leaves and branches take 21.01%, 41.41%, 23.08% and 14.50%. Elevated temperature changed allocation of soluble proteins in that it stimulated soluble proteins accumulation in roots and improved the uptake of water in roots. 4) Treatment ET increased the content of albumin and globulin in roots, and reduced the content of albumin and globulin in stems, leaves and branches. The content of promalin in roots and leaves was increased significantly, and the content of glutenin in stems was reduced significant. This suggested that ET stimulated the accumulation of albumin and globulin in roots and accumulation of promalin in leaves and roots; that treatment ET increased content of soluble protein in leaves was mainly resulted from the increased promalin content in leaves. 5) Regarding treatment ETC, the total of weight of soluble proteins increased, but not significantly; but increased in stems. So the combination of elevated atmospheric CO2 and temperature had not changed the allocation of soluble proteins in red birch seedling and reduced soluble proteins in branches were not the result of increased carbohydrate. 6) Treatment ETC reduced the content of albumin and promalin in roots, stems, leaves and branches, reduced the content of globulin and glutenin in roots significantly. That is to say elevated atmospheric CO2 and temperature reduced the content of soluble proteins in roots significantly which might help to prompt growth, development and nutrient absorption ability of roots. 7) The effects of elevated atmospheric CO2 and temperature on soluble protein cannot be simply interpreted through their mechanism that obtained when they were imposed on plant separately.
Resumo:
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.