陆地森林植被植物细根对全球气候变化的响应研究进展

全球气候变化对陆地森林生态系统与生物多样性造成了较为明显的负面和潜在影响, 由此引发了各种生态环境问题。细根作为植物最活跃的组成部分之一, 在调节陆地森林生态系统碳平衡和养分循环的过程中发挥着重要作用。植物细根对全球气候变化的响应研究已成为当前全球变化背景下陆地生态系统关注的热门课题之一。全球气候变化是以温室气体(CO2、N2O)浓度持续上升、氮沉降加剧、全球气候变暖为主要特征。为此该文从以下几个方面对该领域的研究进展进行综述: (1)CO2浓度升高对植物细根的影响; (2)氮沉降增加对植物细根的影响; (3)温度升高和降水变化对植物细根的影响。最后, 进一步探讨了该领域研究仍存在的科学问题, 提出了未来研究展望。研究结果不仅为进一步研究全球气候变化对植物细根的影响提供重要的理论依据和参考, 也丰富了全球变化背景下根系生态学相关科学理论。     

华西雨屏区苦竹林土壤呼吸对模拟氮沉降的响应

2007年11月至2008年11月, 对华西雨屏区苦竹(Pleioblastus amarus)人工林进行了模拟氮沉降试验, 氮沉降水平分别为对照(CK, 0 g N·m^-2·a^-1)、低氮(5 g N·m^-2·a^-1)、中氮(15 g N·m^-2·a^-1)和高氮(30 g N·m^-2·a^-1)。每月下旬, 采用红外CO₂分析法测定土壤呼吸速率, 并定量地对各处理施氮(NH₄NO₃)。结果表明: 2008年试验地氮沉降量为8.241 g·m^-2, 超出该地区氮沉降临界负荷。在生长季节, 苦竹林根呼吸占总土壤呼吸的60%左右。模拟氮沉降促进了苦竹林土壤呼吸速率, 使苦竹林土壤每年向大气释放的CO₂增加了9.4%~28.6%。在大时间尺度上(如1 a), 土壤呼吸主要受温度的影响。2008年6~10月, 土壤呼吸速率24 h平均值均表现为: 对照<低氮<中氮<高氮。氮沉降处理1 a后, 土壤微生物呼吸速率和土壤微生物生物量碳、氮增加, 并且均与氮沉降量具有相同趋势。各处理土壤呼吸速率与10 cm土壤温度、月平均气温呈极显著指数正相关关系, 利用温度单因素模型可以解释土壤呼吸速率的大部分。模拟氮沉降使得土壤呼吸Q₁₀值增大, 表明氮沉降可能增强了土壤呼吸的温度敏感性。在氮沉降持续增加和全球气候变暖的背景下, 氮沉降和温度的共同作用可能使得苦竹林向大气中排放的CO₂增加。     

高寒草甸土壤呼吸及其组分对氮添加的响应

本研究比较了青藏高原高寒草甸土壤呼吸速率(Rs)、自养呼吸速率(Ra)和异养呼吸速率(Rh)随施氮梯度的变化,揭示土壤呼吸及其组分变化的主要影响因素,为评价未来氮沉降背景下高寒草甸土壤碳释放提供科学依据。于2014年在四川红原青藏高原高寒草甸建立长期氮素添加平台,采取完全随机区组试验设计,设置0(N0,对照)、2(N2)、4(N4)、8(N8)、16(N16)和32 g N·m^-2·a^-1(N32)6个水平氮素添加控制实验。于2020年生长季对Rs、Ra和Rh进行监测。结果表明：施氮显著降低了土壤呼吸及其组分(P<0.05),且Ra的下降幅度大于Rh,导致Rh/Rs随施氮水平逐渐上升;不同施氮处理下Ra和Rh与土壤温度均呈显著的指数正相关(P<0.05);施氮降低了Ra的温度敏感性(Q₁₀),但提高了Rh的Q₁₀值;土壤呼吸各组分与土壤湿度的关系均不显著,但土壤温度和土壤湿度双因子模型对Ra和Rh的解释度高于单因素模型。本研究揭示了高寒草甸土壤呼吸及其组分对氮添加的响应特征及机制,可为评...     

冬小麦旺盛生长期间CO2浓度升高对根际呼吸的影响

依托FACE（free air carbon dioxide enrichment）技术平台,利用阻断根法,采用H6400红外气体分析仪（IRGA）-田间原位测定的方法,研究了大气CO2浓度升高和不同氮肥水平对水稻／小麦轮作制中冬小麦旺盛生长期间根际呼吸的影响。结果表明,在整个测定期间,大气CO2浓度升高增强了根际呼吸速率,提高了根际呼吸排放量。在高N和低N处理中,高CO2浓度下的根际呼吸总排放量分别比Ambient极显著增加117．0％和90．8％。根际呼吸速率在孕穗初期达到最大值;使根际呼吸在土壤呼吸中的比重由24．5％（LN）～26．7（HN）提高到39．8％（LN）～47．1％（HN）。CO2浓度升高与氮肥用量对根际呼吸产生交互效应。表明大气CO2浓度升高将加快土壤向大气的CO2排放,结果将有助于评价未来高CO2浓度背景下农田生态系统土壤碳的固定潜力。     

杉木幼苗和伴生植物细根对土壤增温的生理生态响应

为揭示全球变暖背景下杉木人工林幼苗与其伴生的其它植物间的对土壤养分的竞争关系和适应性,本研究采用埋设加热电缆进行土壤增温(+5℃)技术,在福建省三明市陈大国有采育场内建立杉木(Cunninghamia lanceolata)幼苗试验小区,包括对照(NW)与增温(WNW)处理(均不除草)。采用内生长环法与土钻法相结合,测定增温对杉木幼苗及伴生的其他植物(主要为山油麻Helicteres angustifolia、东南野桐Mallotus lianus)等细根生物量、呼吸、形态、及根组织氮浓度的短期影响。结果表明,(1)增温显著降低了杉木1mm细根生物量,而显著增加了其他植物1mm细根生物量。增温显著提高了其他植物1mm细根的氮浓度,显著降低了其比根长(SRL)和比表面积(SRA);同时降低了比根呼吸(参比温度18℃,SRR_(18)),表明细根呼吸对增温产生了驯化现象。而增温对杉木细根的氮浓度没有显著影响,却显著提高了1mm细根比表面积;同时增温对杉木SRR_(18)没有显著影响,表明杉木细根呼吸没有产生驯化现象。(2)SRR_(18)与比根长间的关系受到增温的显著影响,但树种以及增温×树种的交互作用没有显著影响,表明杉木和其他植物细根竞争能力与维持成本间的平衡关系均受到增温的共同影响。综上结果显示,相较于杉木,伴生的其他植物在增温环境中对地下资源的竞争具有更强的优势,能通过增加细根生物量迅速抢夺吸收因增温而加速矿化的土壤养分,同时通过生理和形态的调整,减少根系单位质量的维持成本,从而提高其对全球变暖的适应性;而杉木在增温条件下面临其他植物的强烈竞争,细根生物量降低,处于不利地位,为了满足生长所需,需增大比根长和比根表面积,且因细根呼吸没有产生驯化现象,从而增加了细根单位质量的维持成本,说明杉木对全球变暖的适应性低于其他植物。该研究结果对于全球变暖下杉木人工林的管理具有重要意义。     

岷江冷杉根际土壤微生物对大气CO2浓度和温度升高的响应

应用自控、封闭、独立的生长室系统,研究了川西亚高山岷江冷杉根际土壤微生物数量对大气CO₂浓度升高 (环境CO₂浓度+350(±25)μmol·mol^-1,EC)和温度升高(环境温度+2.2(±0.5)℃,ET)及其CO₂浓度和温度同时升高 (ECT)的响应.结果表明,1)同对照(CK)相比,在6月、8月和10月,EC处理的根际细菌数量分别增加了35%、164%和312%,ET处理增加了30%、115%和209%,而EC和ET处理对根际放线菌和根际真菌数量影响不显著;ECT处理的根际放线菌数量分别增加了49%、50%和96%,根际真菌数量增加了151%、57%和48%,而ECT对根际细菌数量影响不显著.2)3种处理对非根际土壤微生物数量影响均不显著.3)在EC、ET和ECT处理下,微生物总数的根际效应明显,其R/S值分别为1.93、1.37和1.46(CK的R/S值为0.81).     

郑州植物物候对气候变化的响应

利用统计和突变分析方法,对郑州气候（1956～2003年）和4种乔木物候（1986～2003年）的趋势变化特征进行了分析,并探讨了植物物候期与平均温度、日照的相关性以及对温度变化的响应趋势。分析发现：（1）郑州近50a来在冬、春季升温现象明显;日照在夏季下降最为显著,冬季其次,但在2～4月份历年呈弱上升趋势。（2）物候期变化趋势表现在展叶、开花、果熟期（除楝树外）呈提前趋势,落叶期略有推迟,绿叶期延长,特别是在20世纪90年代中后期,春季物候期（除垂柳外）提前10d左右。（3）平均温度是影响物候期最为显著的气候因子,温度每升高1℃,春季物候平均提前6d左右,绿叶期延长9．5～18．6d;物候期突变一般发生在温度突变之后。以上分析说明植物物候对气候变化响应比较敏感,通过分析和掌握气候和物候变化规律,了解其对当地植物物候的可能影响,可为农业生产、生态环境监测和评估等提供一些理论依据。     

落羽杉奇特的变态根——呼吸根

落羽杉为杉科落羽杉属古老“孑遗植物”,具有耐低温、耐盐碱、耐水淹、抗风、抗污染和抗病虫害等优良特性,既能生长于陆地,也可以生长在沼泽潮湿地带。因土壤潮湿,空气极为缺乏,会导致淹水根系严重缺氧。落羽杉为避免根系室息死亡．常于受窒息的根部生长出一批反向细胞,垂直向上生长,露出地面,形成千姿百态起呼吸作用的棕红色筒状膝根．即呼吸根。呼吸根是植物在缺氧胁迫下的一种生理反应,呼吸根内有许多发达气道,输送和贮存空气,起到一定的呼吸、通气、固着和贮藏养分等作用。所以落羽杉不怕水淹,在水中也能成长。     

滇西北纳帕海湖滨带优势植物茭草茎解剖结构对模拟增温的响应

高原湿地湖滨带植物对气候变暖表现出强烈的功能响应,是全球气候变化的主要现象之一。植物解剖性状直接关系到植物的生态功能,为探讨气候变暖对湿地植物茎解剖结构的影响,该研究利用开顶式生长室分析了模拟增温对滇西北纳帕海湿地湖滨带挺水植物茭草茎解剖结构的影响。结果表明:(1)茭草地上茎在增温4 ℃的范围内,主要通过增加表皮结构厚度以增加表皮失水来响应增温; 地下茎在增温2 ℃的轻度增温条件下与地上茎的响应策略相同,而在增温4 ℃时主要通过减小维管结构大小以降低气穴化风险来响应增温。(2)年最高温度和夜间积温是影响茭草茎解剖结构性状的关键因子,但该两个温度因子仅对地下茎筛管大小的影响达到显著水平(R²=0.838, P<0.01)。(3)内表皮细胞厚度是地上茎响应增温的最主要性状,并与温度因子呈显著正相关。地下茎导管和筛管大小是地下茎响应温度升高的主要性状,二者与温度变量呈负相关关系。综上表明,茭草地上茎和地下茎对增温响应策略存在差异,为揭示高原湿地植物应对气候变暖的响应规律以及生态适应策略提供了科学依据。基于当前气候变暖的背景,建议未来采用更科学的实验方法对更多高原湿地植物的生态响应过程及规律进一步深入研究。     

植物暗呼吸作用对大气CO2浓度升高的响应

植物暗呼吸作用对ＣＯ２浓度升高的响应目前存在两种截然相反的观点：一种认为暗呼吸作用将随着ＣＯ２浓度的升高而下降,可能的原因有胞间ＣＯ２浓度升高、呼吸酶活性改变及暗固定ＣＯ２作用的加强等直接原因;另一种认为暗呼吸作用将随ＣＯ２浓度的升高而提高,影响因素可归结为碳水化合物含量增加、高ＣＯ２浓度刺激其他呼吸途径和生长加快等间接原因。由于目前国际上在实验手段、材料及呼吸作用表达方式等方面的不一致性,这些观点尚难定论,需要更多的实验数据来进一步验证。     

Thermal acclimation and the dynamic response of plant respiration to temperature

Temperature-mediated changes in plant respiration (R) are now accepted as an important component of the biosphere's response to global climate change. Here we discuss the underlying mechanisms responsible for the dynamic response of plant respiration to short and long-term temperature changes. The Q(10) is often assumed to be 2.0 (i.e. R doubles per 10 degrees C rise in temperature); however, the Q(10) is not constant (e.g. it declines near-linearly with increasing temperature). The temperature dependence of Q(10) is linked to shifts in the control exerted by maximum enzyme activity at low temperature and substrate limitations at high temperature. In the long term, acclimation of R to temperature is common, in effect reducing the temperature sensitivity of R to changes in thermal environment, with the temperature during plant development setting the maximal thermal acclimation of R.     

Soil respiration under climate warming: differential response of heterotrophic and autotrophic respiration

Despite decades of research, how climate warming alters the global flux of soil respiration is still poorly characterized. Here, we use meta‐analysis to synthesize 202 soil respiration datasets from 50 ecosystem warming experiments across multiple terrestrial ecosystems. We found that, on average, warming by 2 °C increased soil respiration by 12% during the early warming years, but warming‐induced drought partially offset this effect. More significantly, the two components of soil respiration, heterotrophic respiration and autotrophic respiration showed distinct responses. The warming effect on autotrophic respiration was not statistically detectable during the early warming years, but nonetheless decreased with treatment duration. In contrast, warming by 2 °C increased heterotrophic respiration by an average of 21%, and this stimulation remained stable over the warming duration. This result challenged the assumption that microbial activity would acclimate to the rising temperature. Together, our findings demonstrate that distinguishing heterotrophic respiration and autotrophic respiration would allow us better understand and predict the long‐term response of soil respiration to warming. The dependence of soil respiration on soil moisture condition also underscores the importance of incorporating warming‐induced soil hydrological changes when modeling soil respiration under climate change.     

川西亚高山针叶林土壤呼吸速率与不同土层温度的关系

采用密闭气室红外CO2分析法（IRGA）,在野外连续定位测定了川西亚高山冷杉原始林的土壤呼吸,并对其不同土层（0、5、10、15和20cm）的温度进行了同步测定．在此基础上,分析了土壤呼吸的日、季节动态变化,及其与不同土层温度的关系和土壤呼吸Q10值变化．结果表明：冷杉原始林土壤呼吸呈现明显的日变化和季节变化．土壤呼吸的日最高值出现在12：00-14：00,最低值出现在8：00—10：00,与土壤表面温度的日变化一致;土壤呼吸的季节变化表明：7—8月的土壤呼吸高于9-11月,与不同土层温度季节变化规律一致;土壤呼吸与不同土层温度呈显著的指数增长关系,土壤呼吸速率与土壤15cm深处温度的相关性明显高于其它土层;利用Q10模型计算0～20cm各土层的Q10值分别为2．36、4．75、4．90、6．27和5．46,表明海拔高、温度低的环境条件下,土壤呼吸的Q10值偏高．     

Thermal acclimation of shoot respiration in an Arctic woody plant species subjected to 22 years of warming and altered nutrient supply

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long‐term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short‐term temperature (T) response and sensitivity of leaf respiration (R), the high‐T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q₁₀); overall, Q₁₀ values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high‐resolution measurements of R across a range of measuring Ts (25–70 °C) yielded insights into the T at which maximal rates of R occurred (T_max). Although growth temperature did not affect T_max, N+P fertilization increased T_max values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.     

氮添加对沙质草地微生物呼吸与根系呼吸的影响

土壤呼吸可以细化为根系呼吸和微生物呼吸,二者对氮添加的响应有所不同.本文以科尔沁沙质草地为研究对象,探讨氮添加对土壤CO₂排放的影响,并细化为微生物呼吸和根系呼吸的响应特征.结果表明: 在观测期(5—10月),土壤呼吸、微生物呼吸月动态均呈先升高后降低的趋势;微生物呼吸是土壤呼吸的主要贡献者,占82.6%;观测期内根系呼吸贡献率随月份而变化,根系呼吸贡献率两个峰值分别出现在5月(占49.4%)和8月(占41.9%),6个月的平均贡献率为17.4%;在10 ℃条件下,根系呼吸较微生物呼吸对氮添加的响应更为敏感,微生物呼吸速率在氮添加后降低了3.9%,而根系呼吸降低了17.7%;氮添加提高了土壤呼吸、微生物呼吸温度敏感性Q₁₀值,也提高了二者对土壤水分变化的敏感程度.     

Thermal acclimation of leaf and root respiration: an investigation comparing inherently fast- and slow-growing plant species

We investigated the extent to which leaf and root respiration (R) differ in their response to short‐ and long‐term changes in temperature in several contrasting plant species (herbs, grasses, shrubs and trees) that differ in inherent relative growth rate (RGR, increase in mass per unit starting mass and time). Two experiments were conducted using hydroponically grown plants. In the long‐term (LT) acclimation experiment, 16 species were grown at constant 18, 23 and 28 °C. In the short‐term (ST) acclimation experiment, 9 of those species were grown at 25/20 °C (day/night) and then shifted to a 15/10 °C for 7 days. Short‐term Q₁₀ values (proportional change in R per 10 °C) and the degree of acclimation to longer‐term changes in temperature were compared. The effect of growth temperature on root and leaf soluble sugar and nitrogen concentrations was examined. Light‐saturated photosynthesis (A_sat) was also measured in the LT acclimation experiment. Our results show that Q₁₀ values and the degree of acclimation are highly variable amongst species and that roots exhibit lower Q₁₀ values than leaves over the 15–25 °C measurement temperature range. Differences in RGR or concentrations of soluble sugars/nitrogen could not account for the inter‐specific differences in the Q₁₀ or degree of acclimation. There were no systematic differences in the ability of roots and leaves to acclimate when plants developed under contrasting temperatures (LT acclimation). However, acclimation was greater in both leaves and roots that developed at the growth temperature (LT acclimation) than in pre‐existing leaves and roots shifted from one temperature to another (ST acclimation). The balance between leaf R and A_sat was maintained in plants grown at different temperatures, regardless of their inherent relative growth rate. We conclude that there is tight coupling between the respiratory acclimation and the temperature under which leaves and roots developed and that acclimation plays an important role in determining the relationship between respiration and photosynthesis.     

Leaf respiration is differentially affected by leaf vs. stand-level night-time warming

Plant respiration is an important physiological process in the global carbon cycle serving as a major carbon flux from the biosphere to the atmosphere. Respiration is sensitive to temperature providing a link between environmental variability, climate change and the global carbon cycle. We measured leaf respiration in Populus deltoides after manipulating the air temperature surrounding part of a single leaf, and compared this to the temperature response of the same leaves after manipulating the temperature of the stand. The short‐term temperature response of respiration (Q₁₀– change in the respiration rate with a 10 °C increase in leaf temperature) was 1.7 when the leaf temperature was manipulated, but 2.1 when the stand‐level temperature was changed. As a result, total night‐time carbon release during the five‐day experiment was 21% lower when using the Q₁₀ estimates from the tradition leaf manipulation compared to the stand‐level manipulation. We conclude that the temperature response of leaf respiration is related to whole plant carbon and energy demands, and that appropriate experimental procedures are required in examining respiratory CO₂ release under variable temperature conditions.     

Evergreen leaf respiration acclimates to long-term nocturnal warming under field conditions

Acclimation of plant respiration rates (R) to climate warming is highly variable and many results appear contradictory. We tested the recently suggested hypotheses that pre‐existing, long‐lived leaves should exhibit a relatively limited ability for R to acclimate to climate warming, and that acclimation would occur via changes in the short‐term temperature sensitivity of respiration. Seedlings of a subalpine, evergreen tree species (Eucalyptus pauciflora) were grown under naturally fluctuating conditions within its natural distribution. We used a free air temperature increase (FATI) system of infra‐red ceramic lamps to raise night‐time leaf temperatures by 0.3±0.1, 1.3±0.1, and 2.2±0.1 °C above ambient for 1 year. Light‐saturated assimilation rates and plant growth did not change with nocturnal FATI treatments. Leaf R measured at prevailing temperatures did not differ between FATI treatments. Within each FATI treatment, nocturnal leaf R was highly sensitive to artificial temperature changes within minutes, and also correlated strongly with natural nocturnal and seasonal temperature variation. The corresponding values of Q₁₀ of R varied according to time scale of measurements, but did not vary between FATI treatments. Instead, acclimation of R to nocturnal FATI occurred through changes in the base rate of respiration.     

土壤呼吸对温度升高的适应

土壤呼吸是陆地生态系统碳循环的重要环节之一 ,其对温度升高的敏感程度在相当大的程度上决定着全球气候变化与碳循环之间的反馈关系。土壤呼吸对温度升高的适应是个比较普遍的现象 ,其表现形式主要为随着温度的持续升高和升温时间的延长 ,土壤呼吸对温度升高反应的敏感程度下降。产生这一现象的机制包括影响因子主导地位的转移和温度以外其他因子的协同变化。土壤呼吸对温度升高的适应可以视为碳循环对全球变暖的负反馈效应 ,它可能会在一定程度上缓和陆地生态系统对全球气候系统之间的耦合作用 ,并且导致土壤呼吸对全球温度升高响应的时空差异。由于目前生态系统模型多数没有考虑土壤呼吸的对温度升高的适应性 ,而采用统一的 Q1 0 值 ,其对未来土壤呼吸和未来气候变化幅度的预测可能存在偏差