RESPONSES OF PHENOLOGY AND GROWTH OF BETULA UTILIS AND ABIES FAXONIANA IN SUBALPINE TIMBERLINE ECOTONE TO SIMULATED GLOBAL WARMING, WESTERN SICHUAN, CHINA

 采用开顶式生长室(Open-top chamber, OTC)模拟增温对植被影响的研究方法, 研究了川西亚高山林线交错带糙皮桦(Betula utilis) 和岷江冷杉(Abies faxoniana)幼苗物候及生长特性对模拟增温的响应。结果表明, 温度升高使岷江冷杉幼苗芽开放时间显著提前(15.2 d); 糙 皮桦春季芽物候期变化不显著, 而落叶时间明显推迟(19.7 d), 叶寿命延长(22.8 d)。与对照(CK)相比, OTC内糙皮桦叶面积和岷江冷杉叶片长度及两者侧枝生长速率都显著加快。模拟增温对两物种基径相对生长速率都表现为正效应, 增温对两物种枝叶特性及分布格局表现为不同程度 的正效应、负效应或无影响。不同功能型两物种对模拟增温响应方式存在一定程度差异。     

增温对川西亚高山针叶林内不同光环境下红桦和岷江冷杉幼苗生长和生理的影响

 川西亚高山针叶林是青藏高原东部高寒林区的重要组成部分, 也是研究全球变化对森林生态系统影响的重要森林类型。开展亚高山针叶林不同树种对气候变暖响应差异的研究, 可为预测未来气候变暖背景下亚高山针叶林植被组成和森林动态提供科学依据。我们以川西亚高山针叶林两种主要树种——红桦(Betula albo-sinensis)和岷江冷杉(Abies faxoniana)为研究材料, 采用开顶式增温法(Open-top chamber, OTC)模拟气候变暖, 研究了增温对全光条件和林下低光环境中(约为全光的10%)生长的红桦和岷江冷杉幼苗生长和生理的影响。在人工林环境下, OTC使增温框内平均气温和地表温度分别升高了0.51和0.33 ℃; 而在林外空地处, OTC使二者分别升高了0.69和0.41 ℃。研究结果显示, 增温总体上促进了两种幼苗的生长和生理过程, 并促使幼苗将更多的生物量投入到其同化部位——叶, 使幼苗的根冠比(R/S)显著降低。增温通过增加叶片的光合色素含量和表观量子效率等光合参数, 促进了幼苗的光合过程和生长。然而, 增温对两种幼苗生长和生理的影响效果与植物种类及其所处的光环境有关。增温仅在林外全光条件下显著影响红桦幼苗的生长和生理过程。岷江冷杉幼苗对增温的响应与红桦相反, 即增温仅在林下低光环境下对岷江冷杉幼苗的生长和生理过程有明显促进作用。这种响应差异可能赋予这两种植物在未来气候变暖背景下面对外界环境变化时具有不同的适应能力和竞争优势, 从而对亚高山针叶林生态系统物种组成和森林动态产生潜在影响。     

长期模拟增温对岷江冷杉幼苗生长与生物量分配的影响

川西亚高山针叶林是青藏高原东部高寒林区的重要组成部分,也是研究全球变化对森林生态系统影响的重要组成。长期模拟增温对川西亚高山森林树木的生长、物质积累及其分配格局的影响至今鲜有报道。以川西亚高山针叶林优势种岷江冷杉(Abies faxoniana)幼苗为研究对象,采用控制环境生长室模拟增温的方法,研究了模拟增温对岷江冷杉幼苗生长、物质积累及其分配格局的影响。结果表明,模拟增温(2.2±0.2)℃处理65个月后,岷江冷杉幼苗基径、株高、单株叶面积和比叶面积(SLA)均显著增加,比叶重(LMA)显著下降。增温对岷江冷杉幼苗的茎、侧枝、叶和总生物量具有显著的促进作用,对根生物量没有显著影响。岷江冷杉幼苗的叶重比(LMR)下降、枝重比(SMR)增加、根重比(RMR)无显著变化。长期增温能显著促进岷江冷杉幼苗的生长和物质积累,改变生物量分配格局,促使叶片物质向茎转移,降低光合物质投入。     

模拟增温和功能群去除对岷江冷杉幼苗存活和生长的影响

通过模拟增温和功能群去除试验,研究不同功能群的草本植物在岷江冷杉幼苗更新定植过程的作用,分析其在温度升高条件下生理指标的变化.结果表明:增温提高了岷江冷杉幼苗的存活率和植物体内非结构性碳水化合物含量,但抑制了幼苗的生长和根冠比.岷江冷杉幼苗的非结构性碳水化合物含量,尤其是可溶性糖含量与存活率呈显著正相关.不增温处理中,草本植物降低了幼苗的存活率,促进幼苗的株高生长和地上生物量积累,禾本科植物和杂类草抑制了幼苗根系生长和地下生物量的积累;增温处理中,杂类草的存在提高了幼苗的存活率,莎草科则显著抑制了幼苗根系生长和地下生物量的积累,禾本科植物和杂类草抑制了幼苗的高生长和地上生物量积累.模拟增温提高了林线树种岷江冷杉幼苗的存活率,但使其面临来自草本植物更剧烈的竞争,抑制其生长.     

川西林线交错带岷江冷杉幼苗异龄叶形态对长期模拟增温的响应

全球气候变暖对高纬度、高海拔地区的植物形态产生强烈影响。川西林线交错带是青藏高原东部高寒生态系统的重要组成部分,对全球变化极度敏感。以川西林线交错带岷江冷杉(Abies faxoniana)幼苗异龄叶为对象,采用原位开顶式生长室(Open-top chamber,OTCs)模拟增温,研究了长期模拟增温下岷江冷杉幼苗异龄叶片叶长、叶厚等叶形态的响应,采用表型可塑性指数和变异系数对叶形态的可塑性进行分析。结果表明:增温限制岷江冷杉幼苗叶片的增长、增宽和叶面积、体积的扩大,使叶长、叶宽、叶面积、叶体积分别较对照减小12.77%、11.86%、17.76%、11.49%;增温促进叶片厚度的增加,较对照增加7.27%;除叶长宽比外,增温对其余叶形态均产生显著影响(P0.05)。叶形态对模拟增温的响应具有显著的年龄差异(P0.05)。温度、叶龄的交互效应对叶长和叶面积影响显著(P0.05),对叶宽和叶厚影响不显著(P0.05)。两种表型可塑性分析结果表明,除1 a叶叶长外,增温不同程度增大各叶形态可塑性。长期增温使冷杉幼苗叶片有旱生倾向且形态值更发散。研究提供了岷江冷杉幼苗叶片对长期增温的差异性响应证据,为评估青藏高原东缘优势植物对响应气候变暖提供了基础数据和理论参考。     

模拟增温对川西亚高山林线交错带绵穗柳生长、叶物候和叶性状的影响

采用开顶式生长室(open-top chamber, OTC)模拟增温的方法,研究了模拟增温对川西亚高 山林线交错带绵穗柳生长和叶性状的影响.结果表明：与对照样地相比,OTC内日平均气温(地上12 m)在植物生长季中增加2.9 ℃,而5 cm土壤温度仅增加0.4 ℃;温度升高使绵穗柳芽开放时间明显提前、落叶时间明显推迟,叶寿命延长;OTC内绵穗柳叶面积和侧枝生长速率明显加快,比叶面积明显增加,而叶氮浓度却显著下降;OTC内绵穗柳的气孔导度、净光合速率、光呼吸速率和暗呼吸速率总体上呈增加趋势.综上所述,绵穗柳适应增温效应的能力较强,在未来气候变化背景下,其分布的海拔高度有可能上升.     

模拟增温对川西亚高山林线交错带绵穗柳生长、叶物候和叶性状的影响

采用开顶式生长室(open-top chamber, OTC)模拟增温的方法,研究了模拟增温对川西亚高 山林线交错带绵穗柳生长和叶性状的影响.结果表明：与对照样地相比,OTC内日平均气温(地上12 m)在植物生长季中增加2.9 ℃,而5 cm土壤温度仅增加0.4 ℃;温度升高使绵穗柳芽开放时间明显提前、落叶时间明显推迟,叶寿命延长;OTC内绵穗柳叶面积和侧枝生长速率明显加快,比叶面积明显增加,而叶氮浓度却显著下降;OTC内绵穗柳的气孔导度、净光合速率、光呼吸速率和暗呼吸速率总体上呈增加趋势.综上所述,绵穗柳适应增温效应的能力较强,在未来气候变化背景下,其分布的海拔高度有可能上升.     

升温突变对川西不同坡向和海拔岷江冷杉径向生长的影响

坡向和海拔作为重要的地形因子决定着林木立地条件的水热分配，为揭示川西地区升温突变前后制约岷江冷杉(Abies fargesii var.faxoniana)径向生长的气候因子变化及其在坡向和海拔上的响应格局，通过树木年轮学方法构建了4个坡向(NE、N、NW和W)的3个海拔梯度(≥3650 m)共12个岷江冷杉的标准年表，采用皮尔逊相关分析和回归分析的方法分析了升温突变前后(1980年)限制岷江冷杉径向生长主要气候因子的变化及径向生长的变化趋势。结果表明：升温前，生长季(7月)低温和降水、前一年冬季(10—11月)最高温和平均温制约岷江冷杉的生长，而当年春季(3月)温度的升高和生长季前(5月)较多的降水不利于其生长。升温后，4个坡向的林线、东北坡中海拔，西坡中、低海拔岷江冷杉的径向生长与大部分月温度表现为正相关关系，且上述样点树木径向生长明显加速。1980年升温前，制约不同样点岷江冷杉径向生长的气候因子具有一致性——坡向和海拔对其径向生长与温度相关关系的干扰和影响较小。增温促进了各坡向林线岷江冷杉的径向生长，且偏阳坡在更大的海拔范围内对增温表现出正反馈。研究对未来气候变化背景下川西不同坡...     

川西米亚罗林区主要树木生长对气候响应的差异

为分析青藏高原东缘半湿润区不同树种树木生长对气候变化的响应规律,于川西米亚罗林区海拔3000 m左右(低海拔)采集铁杉、岷江冷杉、紫果云杉,海拔4000 m左右林线位置(高海拔)采集岷江冷杉、四川红杉,共计182棵树木年轮样芯,建立了不同树种的树轮宽度年表,对不同树种的年轮指数与各月气候因子进行相关分析.结果表明： 在低海拔处,树木生长与4、5月气温呈负相关,与4、5月降雨呈正相关,受到春季干旱胁迫的影响;但树种之间存在显著差异： 铁杉的生长受春季干旱胁迫影响最严重,岷江冷杉次之,紫果云杉所受影响很小.在高海拔处,树木生长主要受生长季温度的影响,岷江冷杉年轮指数与当年2、7月最低气温呈显著正相关,与上一年10月最高气温亦呈正相关;四川红杉年轮指数与5月最高气温呈显著正相关,但与2月均温、3月最低气温呈显著负相关.近几十年青藏高原东北缘气候有干暖化趋势,如果这种趋势持续发生,低海拔紫果云杉长势将超过铁杉和岷江冷杉;高海拔处的升温更有利于岷江冷杉的生长.     

高山林线土壤微生物群落结构对模拟增温的响应

研究土壤微生物群落结构对模拟增温的响应,对预测全球气候变化背景下土壤碳氮磷循环具有重要意义.采用开顶式生长室(OTC)模拟增温,研究了土壤有机质层和矿质土壤层真菌(F)、细菌(B)、革兰氏阳性菌(G⁺)和革兰氏阴性菌(G^-)PLFAs微生物量,以及真菌/细菌(F/B)和革兰氏阴性菌/革兰氏阳性菌(G^-/G⁺)比值对模拟增温的响应.结果表明: OTC模拟增温使空气温度增加0.87 ℃,土壤有机质层温度增加0.5 ℃,矿质土壤层温度增加0.23 ℃.土壤有机质层微生物群落组成比矿质土壤层对模拟增温的响应更敏感.细菌比真菌对模拟增温的响应更加敏感,模拟增温显著影响了土壤有机质层的F/B和G^-/G⁺比值,对矿质土壤层的所有PLFAs含量或比值均没有显著影响.微生物的PLFAs含量及真菌/细菌和G^-/G⁺比值总体呈现非生长季低于生长季前期和生长季后期.冗余分析表明,土壤中的碳含量(可溶性有机碳DOC 12.1%、凋落物可溶性碳DC 9.5%和全碳TC 3%)是微生物群落结构的决定性因素,可溶性组分(DOC和DC)对微生物群落结构的影响大于全量养分(全碳和总氮).     

Short-term responses of phenology, shoot growth and leaf traits of four alpine shrubs in a timberline ecotone to simulated global warming, Eastern Tibetan Plateau, China

The short-term effects of artificial warming on phenology, growth and leaf traits were investigated in four alpine shrubs using the open-top chamber (OTC) method in a timberline ecotone (3240 m a.s.l.) on the Eastern Tibetan Plateau. The OTC enhanced the mean air temperature by 2.9°C throughout the growing season. In contrast, only a slight difference (0.4°C) in the mean soil temperature was observed in the OTC compared with the control plots (CP). Spiraea mongolica , Potentilla fruticosa , Conicera hispida (deciduous shrubs) and Daphne retusa (evergreen shrub) showed earlier bud break, flowering and fruit coloring as well as longer flower longevity in the OTC than in the CP. All deciduous shrubs in the OTC had a longer leaf lifespan. Daphne retusa had higher leaf survival rates in the OTC. No significant differences in the total number of flowers and fruits were noticed for most species between the two treatments. Warming stimulated the shoot and leaf growth for most species. The specific leaf area tended to increase for many species in the OTC. However, the leaf nitrogen concentration tended to decrease in P. fruticosa and S. mongolica. The results obtained in the present study indicate that warming conditions can have strong impacts on alpine shrubs in a timberline ecotone.     

Effect of ecosystem warming on boreal black spruce bud burst and shoot growth

The boreal forest is predicted to experience the greatest warming of any forest biome during the next 50–100 years, but the effects of warming on vegetation phenology are not well known. The objectives of this study were to (1) examine the effects of whole ecosystem warming on bud burst and annual shoot growth of black spruce trees in northern Manitoba, Canada and (2) correlate bud burst to cumulative degree-days (CDD). The experimental design was a complete randomized block design that consisted of four replicated blocks. Each replicate block contained four treatments: soil warming only (heated outside, HO), soil and air warming (heated inside, HI), control outside (no chamber, no heating, CO), and inside a chamber maintained at ambient conditions (no soil or air warming, control inside, CI). Bud burst was measured during the first and second years of the experiment, starting in 2004, and annual shoot growth was measured for the first 3 years (2004–2006) of the study. On average, shoot bud burst occurred 11 and 9 days earlier in 2004 and 2005, respectively, for HI than for other treatments. However, mean CDD required for bud burst for HI was within the standard deviation of CO for both years. In year 1 of the treatments, shoot bud burst occurred earlier for HI than other treatments (CI, CO, HO), but final shoot length of HI trees was less than in CO trees. In the second year of warming, final shoot length was not different for HI than CO. By the third year of warming final shoot length was significantly greater for HI than all other treatments. Empirical results from this study suggest that soil and air warming causes an earlier bud burst for all years of observation and greater shoot lengths by the third season of warming. A longer growing season and greater annual shoot growth should increase carbon uptake by boreal black spruce trees in a warmer climate.     

Dynamic and complex microclimate responses to warming and grazing manipulations

Synthesis efforts that identify patterns of ecosystem response to a suite of warming manipulations can make important contributions to climate change science. However, cross‐study comparisons are impeded by the paucity of detailed analyses of how passive warming and other manipulations affect microclimate. Here we document the independent and combined effects of a common passive warming manipulation, open‐top chambers (OTCs), and a simulated widespread land use, clipping, on microclimate on the Tibetan Plateau. OTCs consistently elevated growing season averaged mean daily air temperature by 1.0–2.0°C, maximum daily air temperature by 2.1–7.3°C and the diurnal air temperature range by 1.9–6.5°C, with mixed effects on minimum daily air temperature, and mean daily soil temperature and moisture. These OTC effects on microclimate differ from reported effects of a common active warming method, infrared heating, which has more consistent effects on soil than on air temperature. There were significant interannual and intragrowing season differences in OTC effects on microclimate. For example, while OTCs had mixed effects on growing season averaged soil temperatures, OTCs consistently elevated soil temperature by approximately 1.0°C early in the growing season. Nonadditive interactions between OTCs and clipping were also present: OTCs in clipped plots generally elevated air and soil temperatures more than OTCs in nonclipped plots. Moreover, site factors dynamically interacted with microclimate and with the efficacy of the OTC manipulations. These findings highlight the need to understand differential microclimate effects between warming methods, within warming method across ecosystem sites, within warming method crossed with other treatments, and within sites over various timescales. Methods, sites and scales are potential explanatory variables and covariables in climate warming experiments. Consideration of this variability among and between experimental warming studies will lead to greater understanding and better prediction of ecosystem response to anthropogenic climate warming.     

增温对川西北亚高山高寒草甸植物群落碳、氮含量的影响

采用开顶式生长室（OTC）模拟增温实验,研究了川西北亚高山草甸植物群落碳、氮含量对温度升高的响应。由于OTC的增温作用,在整个生长季内,地温（15 cm）、地表温度和气温(30 cm)的平均值在OTC内比对照样地分别高0.28、0.46和1.4℃,OTC内土壤相对含水量也明显减少,低于对照样地5.49%。受增温及土壤含水量减少的影响,一年后,植物群落的生物量积累和碳、氮含量发生了明显的改变。除10月份OTC内地上鲜体生物量略高于对照样地外,OTC内地上鲜体生物量和根系生物量与对照样地相比,都出现了不同程度的减少;OTC内植物群落地上活体的碳浓度在整个生长季高于对照样地,而氮浓度低于对照样地;OTC内植物群落地下活根的碳浓度在整个生长季高于对照样地,并且在8月份统计检验显著,而氮浓度却低于对照样地;OTC内植物碳库在整个生长季较对照样地有不同程度的增加,增幅范围为0.90%～5.65%,而OTC内植物氮库较对照样地有不同程度的减少,减幅范围0.40%～1.28%。