北京大气中SO2、NOx、CO和O3体积分数变化分析

为了探讨北京市大气污染物的污染水平和变化特征,2004年8月—2005年7月对北京大气中SO2、NOx、O3和CO体积分数进行了连续观测,并对比分析不同季节的变化特征。结果表明,SO2体积分数呈双峰曲线日变化,在08:00和23:00出现峰值。SO2采暖季日振幅明显高于非采暖季日振幅,采暖季SO2体积分数要比非采暖季高出3倍以上。NOx、CO体积分数在早晨07:00和傍晚20:00左右出现峰值,NOx体积分数最大值可达130×10-9,而CO体积分数最大值可达3300×10-9。NO有明显的日变化和季节变化,而NO2白天夜晚都维持在同一水平,且季节变化也不大。O3体积分数夏季远远高于冬季,日变化均呈单峰型分布,午后14:00—15:00出现峰值。     

油菜地CO2、N2O排放及其影响因素

2005年11月至2006年5月采用静态箱法对成都平原典型水稻 - 油菜轮作区油菜地CO2、N2O排放通量进行原位测定.结果表明,CO2排放通量为121.4～1 585.8 mg·m-2·h-1,平均656.8 mg·m-2·h-1;N2O排放通量为18.0～521.0 μg·m-2·h-1,平均168.0 μg·m-2·h-1.在整个油菜生长期内,地下5 cm土壤温度与CO2、N2O排放通量之间呈指数函数关系.3种不同处理油菜地CO2、N2O排放通量均为常规处理＞无氮处理＞裸地处理.土壤温度、施氮和植物生长是影响油菜地CO2、N2O排放的主要因素.     

三江平原湿地岛状林CH4和N2O排放通量的特征

利用静态箱-气相色谱法对三江平原湿地岛状林生长季N2O、CH4的排放通量进行了为期4个月的原位测定,分析了三江平原湿地岛状林CH4、N2O排放通量的季节特征,并初步探讨了CH4、N2O排放通量与温度和土壤水分的关系.湿地岛状林CH4、N2O通量均呈现出明显的正负交替的变化特征, 通量范围分别为-560.45～706.35 μg/(m2·h)和-28.87～43.59 μg/(m2·h), 季节均值分别为41.88 μg/(m2·h)和11.56 μg/(m2·h).结果表明, 湿地三江平原湿地岛状林土壤是CH4和N2O的汇.相关分析表明,CH4和N2O的排放与箱内温度具有弱相关性.土壤水分是影响湿地岛状林土壤CH4吸收、排放关系的重要因素, 而土壤的干湿交替能促进岛状林土壤N2O的排放.     

免耕对华北平原潮土N2O和CO2排放的影响

采用静态箱-气相色谱法研究了免耕和常规耕作下玉米生长季华北平原潮土N2O和CO2的排放特征.结果表明,免耕土壤N2O累积排放量(以N2O-N计,下同)为0.31 kg· hm-2,略高于常规耕作土壤的0.27 kg·hm-2,两者没有显著差异.灌水、强烈降水或连续阴天会诱发土壤大量排放N2O,免耕处理N2O排放峰值(28.1 ～38.4μg·m-2·h-1)高于常规耕作处理(18.6 ～25.7 μg·m-2·h-1).免耕处理CO2累积排放量(以CO2-C计,下同)为1 880 kg·hm-2,显著高于常规耕作土壤的1 333 kg·hm-2.土壤N2O和CO2排放通量与土壤温度呈显著指数相关,常规耕作处理下的相关程度更高.     

CO2-enrichment and nutrient availability alter ectomycorrhizal fungal communities

Ectomycorrhizal fungi (EMF), a phylogenetically and physiologically diverse guild, form symbiotic associations with many trees and greatly enhance their uptake of nutrients and water. Elevated CO2, which increases plant carbon supply and demand for mineral nutrients, may change the composition of the EMF community, possibly altering nutrient uptake and ultimately forest productivity. To assess CO2 effects on EMF communities, we sampled mycorrhizae from the FACTS-I (Forest-Atmosphere Carbon Transfer and Storage) research site in Duke Forest, Orange County, North Carolina, USA, where Pinus taeda forest plots are maintained at either ambient or elevated CO2 (200 ppm above ambient) concentrations. Mycorrhizae were identified by DNA sequence similarity of the internal transcribed spacer ribosomal RNA gene region. EMF richness was very high; 72 distinct phylotypes were detected from 411 mycorrhizal samples. Overall EMF richness and diversity were not affected by elevated CO2, but increased CO2 concentrations altered the relative abundances of particular EMF taxa colonizing fine roots, increased prevalence of unique EMF species, and led to greater EMF community dissimilarity among individual study plots. Natural variation among plots in mean potential net nitrogen (N) mineralization rates was a key determinant of EMF community structure; increasing net N mineralization rate was negatively correlated with EMF richness and had differential effects on the abundance of particular EMF taxa. Our results predict that, at CO2 concentrations comparable to that predicted for the year 2050, EMF community composition and structure will change, but diversity will be maintained. In contrast, high soil N concentrations can negatively affect EMF diversity; this underscores the importance of considering CO2 effects on forest ecosystems in the context of background soil chemical parameters and other environmental perturbations such as acid deposition or fertilizer runoff.     

近地层O3污染对陆地生态系统的影响

随着全球气候变化对生态环境的影响日益增加,近地层臭氧(O3)污染的环境生态效应备受人们关注.现有研究表明,陆地生态系统的温室气体NOx和CH4释放、矿质能源消耗和机动车辆尾气排放量的增加将加剧近地层O3污染.O3污染通过降低植物叶片气孔导度、光合速率和净同化作用,改变同化物的分配,进而抑制植物生长和加速植物老化,导致作物和林木减产.O3污染导致植物-土壤系统碳积累和周定降低势必影响未来全球碳动力学和碳预算,而植物和根系生长受到抑制则不利于土壤养分、水分的吸收进而影响植物-土壤系统养分循环,但目前报导极少,尚无法准确判断对全球碳和养分循环的影响,亟待深入研究.由于环境因素间具有互作效应,目前模拟研究过多集中O3与CO2增加对陆地生态系统的复合效应方面,而与其它环境因子(如O3与NO、SO2、水分、温度等)的复合效应研究偏少,不利于在全球气候变化背景下深入了解与预测O3污染对陆地生态系统的影响程度与趋势.基于研究现状,未来应加强:(1)地表O3监测网络建设和监测,结合田间试验和建模加强草地、森林和农田生态系统对O3污染的响应研究;(2)长期定位研究,侧重陆地生态系统对O3污染连合其它温室气体、温度增加等模拟未来气候情景下的环境响应研究;(3)O3污染下土壤.植物系统碳循环和固定研究;(4)O3污染条件下优势植物和农作物在不同时空条件下的土壤.植物系统养分利用研究;以期为判断和预测全球气候变化背景下陆地生态系统对近地层O3污染加剧的响应程度与趋势提供数据资料和科学依据.     

Litterfall 15N abundance indicates declining soil nitrogen availability in a free-air CO2 enrichment experiment

Forest productivity increases in response to carbon dioxide (CO2) enrichment of the atmosphere. However, in nitrogen-limited ecosystems, increased productivity may cause a decline in soil nitrogen (N) availability and induce a negative feedback on further enhancement of forest production. In a free-air CO2 enrichment (FACE) experiment, the response of sweetgum (Liquidambar styraciflua L.) productivity to elevated CO2 concentrations [CO2] has declined over time, but documenting an associated change in soil N availability has been difficult. Here we assess the time history of soil N availability through analysis of natural 15N abundance in archived samples of freshly fallen leaf litterfall. Litterfall delta15N declined from 1998 to 2005, and the rate of decline was significantly faster in elevated [CO2]. Declining leaf litterfall delta15N is indicative of a tighter ecosystem N cycle and more limited soil N availability. By integrating N availability over time and throughout the soil profile, temporal dynamics in leaf litterfall delta15N provide a powerful tool for documenting changes in N availability and the critical feedbacks between C and N cycles that will control forest response to elevated atmospheric CO2 concentrations.     

大气CO2与植物氮素营养的关系

大气CO2浓度升高对植物吸收氮素,以及对植物和土壤中的氮浓度、C/N比和氮循环都存在着影响。大气CO2浓度与植物氮素营养之间存在着交互作用。大气CO2浓度升高对植物氮素营养影响的结果与氮浓度、氮形态等因素有关。     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

施肥与不施肥措施下小麦田的CO2、CH4、N2O排放日变化特征

二氧化碳（CO2）、甲烷（CH4）、氧化亚氮（N2O）是对全球气候变化影响最大的温室气体。由于土壤与大气之间的水热交换需要一定的传导平衡时间,因此土壤温室气体与温湿度之间的关系存在不同的表现形式。目前,有关温室气体研究多集中于季节性排放特征,而关于CO2、CH4、N2O的日变化研究却少见报道。以北京小麦（Triticum aestivuml）农田土壤为研究对象,对施肥和不施肥条件下CO2、CH4、N2O交换通量和气温、土壤温度进行连续观测,来探讨3种温室气体的日变化特征。采用人工静态暗箱法对小麦田土壤进行连续48 h原位观测,每2 h测定1次,每次盖箱时间为30 min。气体样品中的CO2、CH4、N2O用气相色谱仪（Agilent 6890A,FID/ECD）测定。结果表明：施肥与不施肥条件下小麦生育后期麦田土壤CO2、CH4、N2O交换通量具有明显的日变化特征。土壤表现为CH4的吸收汇、CO2和N2O的排放源。CH4的吸收通量、CO2和N2O的排放通量均表现为施肥区＞对照区。CO2、CH4的交换通量的70%以上出现在白天,而施肥区和对照区的N2O白天排放通量分别达到全天的81.8%、91.1%。另外,相关性分析表明,CO2、N2O交换通量的日变化与气温和5 cm地温呈极显著（P＜0.01）或显著（P＜0.05）的正相关关系,且N2O交换通量日变化与10 cm地温呈现极显著的正相关关系,说明温度是影响CO2、N2O交换通量日变化的重要因素;而气温、5 cm地温、10 cm地温对CH4交换通量日变化不存在显著性影响。     

Efficient and sustainable management of complex forest ecosystems

A large range of models has been developed for the analysis of optimal forest management strategies, with the well-known Faustmann models dating back to the mid-19th century. To date, however, there has been relatively little attention for the implications of complex ecosystem dynamics for optimal forest management. This paper examines the implications of irreversible ecosystem responses for efficient and sustainable forest management. The paper is built around two forest models that comprise two ecosystem components, forest cover and topsoil, the interactions between these components, and the supply of the ecosystem services ‘wood’ and ‘erosion control’. The first model represents a forest that responds in a reversible way to overharvesting. In the second model, an additional ecological process has been included and the ecosystem irreversibly collapses below certain thresholds in forest cover and topsoil depth. The paper presents a general model, and demonstrates the implications of pursuing efficient as well as sustainable forest management for the two forest ecosystems. Both fixed and variable harvesting cycles are examined. Efficient and sustainable harvesting cycles are compared, and it is shown that irreversible ecosystem behaviour reduces the possibilities to reconcile efficient and sustainable forest management through a variable harvesting cycle.     

Nitrous oxide nitrification and denitrification 15N enrichment factors from Amazon forest soils.

The isotopic signatures of 15N and 18O in N2O emitted from tropical soils vary both spatially and temporally, leading to large uncertainty in the overall tropical source signature and thereby limiting the utility of isotopes in constraining the global N2O budget. Determining the reasons for spatial and temporal variations in isotope signatures requires that we know the isotope enrichment factors for nitrification and denitrification, the two processes that produce N2O in soils. We have devised a method for measuring these enrichment factors using soil incubation experiments and report results from this method for three rain forest soils collected in the Brazilian Amazon: soil with differing sand and clay content from the Tapajos National Forest (TNF) near Santarém, Pará, and Nova Vida Farm, Rond?nia. The 15N enrichment factors for nitrification and denitrification differ with soil texture and site: -111 per thousand +/- 12 per thousand and -31 per thousand +/- 11 per thousand for a clay-rich Oxisol (TNF), -102 per thousand +/- 5 per thousand and -45 per thousand +/- 5 per thousand for a sandier Ultisol (TNF), and -10.4 per thousand +/- 3.5 per thousand (enrichment factor for denitrification) for another Ultisol (Nova Vida) soil, respectively. We also show that the isotopomer site preference (delta15Nalpha - delta15Nbeta, where alpha indicates the central nitrogen atom and beta the terminal nitrogen atom in N2O) may allow differentiation between processes of production and consumption of N2O and can potentially be used to determine the contributions of nitrification and denitrification. The site preferences for nitrification and denitrification from the TNF-Ultisol incubated soils are: 4.2 per thousand +/- 8.4 per thousand and 31.6 per thousand +/- 8.1 per thousand, respectively. Thus, nitrifying and denitrifying bacteria populations under the conditions of our study exhibit significantly different 15N site preference fingerprints. Our data set strongly suggests that N2O isotopomers can be used in concert with traditional N2O stable isotope measurements as constraints to differentiate microbial N2O processes in soil and will contribute to interpretations of the isotopic site preference N2O values found in the free troposphere.     

长期不同施肥条件下红壤小麦和玉米季CO2、N2O排放特征

基于中国农业科学院红壤实验站红壤旱地小麦-玉米轮作长期定位试验,采用静态箱/气相色谱法,研究红壤旱地连续施肥16 a后,不同施肥条件下小麦季和玉米季土壤CO2和N2O的排放特征。结果表明,CO2和N2O排放具有明显的季节性,施肥对土壤CO2和N2O排放有明显影响,且有机肥的施用显著促进了土壤CO2和N2O排放。不施肥对照（CK）、氮磷配施（NP）、氮钾配施（NK）、氮磷钾配施（NPK）和有机无机肥配施（NPKM）处理小麦季和玉米季土壤CO2累积排放量分别为5 904、8 062、4 298、9 235、14 098和4 708、7 530、5 435、7 089、15 472 kg.hm-2,土壤N2O累积排放量分别为0.34、0.63、0.44、0.62、1.00和0.25、0.39、0.35、0.52、1.73 kg.hm-2,小麦休闲期土壤CO2和N2 O累积排放量平均占小麦生长季的63.52%和28.43%,玉米休闲季平均占玉米生长季的49.98%和32.72%,说明休闲期土壤CO2和N2O累积排放量不容忽视。除玉米季NP、NK、NPK处理外,其他各处理小麦季和玉米季土壤CO2排放通量与5 cm深处土壤温度显著相关;而土壤N2O排放通量与土壤温度间均未表现出显著相关性;除NPKM处理外,其他各处理土壤CO2或N2O排放通量与土壤水分间相关性均未达显著水平。     

Lignin and its effects on litter decomposition in forest ecosystems

Lignin is a major component of plant litter. In this review, we found lignin comprises a complex class of organic compounds whose concentration differs greatly both between and within plant species. There are many analytical methods for detecting the composition and structure of lignins. As lignins are enormously complex compounds, chemical assay is difficult and different methods vary with the results. Lignin plays a significant role in the carbon cycle, sequestering atmospheric carbon into the living tissues of woody perennial vegetation. It has also great effects on nitrogen dynamics of forest ecosystems as well as other ecological processes. Lignin is one of the most slowly decomposing components of dead vegetation, contributing a major fraction of the material that becomes humus as it decomposes. Lignin is highly correlated with decomposition of litter. Thus, there is evidence that the lignin concentration is a more influential factor than the other chemical concentrations, in determining the rate of leaf litter decomposition of different forest ecosystems. Although a great number of researchers have addressed lignin's role in litter decomposition, still there are many aspects of lignin biogeochemistry that are not known. This lack of information hinders complete amalgam of lignin effects on litter decomposition processes and dynamics of nutrient cycling.     

湖滨带温室气体氧化亚氮(N2O)排放研究

利用不锈钢气体采集箱,在太湖梅梁湾湖区,沿水体至陆地方向对植被型和裸露型湖滨带进行温室气体N2O的原位排放观测。结果发现两种类型湖滨带N2O排放均显著高于临近的开阔水体,可以达到10~100倍,水位变幅区是湖滨带温室气体N2O排放的峰值区,是陆向和水向辐射区的5~10倍。在观测期间,N2O排放通量的范围为-159.2～1565.6μg·m-2·h-1,具有明显的时空梯度变化,时间上,9月份最高,随着气温的下降和芦苇的衰亡而逐渐减少,在1月份最低;空间上,自水向辐射区至陆向辐射区,先逐渐升高,在水位变幅区达到峰值,然后再降低。该结果初步说明了湖滨带是温室气体N2O的一个极其重要的排放源,而目前IPCC对水体N2O排放的估算可能存在很大的疏漏;同时也从一定程度上反映了湖滨带是湖泊脱氮的重要区域,其对缓解湖泊氮污染起到了举足轻重的作用。     

施氮水平对小麦-玉米轮作体系氨挥发与氧化亚氮排放的影响

试验采用密闭室间歇通气法研究华北平原冬小麦（Triticum aestivum Linneaus）-夏玉米（Zea mays Linneaus）轮作体系在不同施氮水平下农田氨挥发和N2O排放。结果表明：冬小麦季和夏玉米季的氨挥发速率与N2O排放通量呈现出季节性动态变化,且随着施氮量的增加而增加,均在施肥后第2~3天出现峰值。玉米季氨挥发量和N2O排放量均高于小麦季,分别占整个轮作周期气态损失的53.5%~68.9%和54.7%~82.3%。轮作体系中氨挥发净损失量（以N计）为4.28~31.31 kg.hm-2,占施氮量的3.17%~5.80%;N2O净排放损失量（以N计）为50.95~1051.03 g.hm-2,占施氮量的比例为0.04%~0.23%。因此,施氮量是影响冬小麦-夏玉米轮作体系气态损失重要因素,且夏玉米季是控制控制气态损失的关键时期。     

退耕土壤的碳、氮固存及其对CO2、N2O通量的影响

采样分析陇中黄土高原地区农田退耕种植苜蓿3 a、5 a、8 a后0～5、5～10、10～20 cm土层土壤有机碳(SOC)、全氮(TN)、活性有机碳(SAOC)及矿质氮(NO3-N、NH4-N)含/储量的变化,并用静态箱-气质联用法对样地的COO2、NO2O排放通量进行了测定,研究碳氮变化对土壤CO2、N2O排放通量的影响.结果表明:(1)SOC、TN基础含量很低的贫瘠土壤退耕后表现出明显的碳、氮固存效应,有很强碳、氮固存潜力.与未退耕休闲农田相比,退耕3 a、5 a、8 a后0～20 cm SOC储量分别提高了9.12%、20.18%、34.39%,SOC平均固存率分别为0.17、0.23、0.25mg/(hm2·a).TN储量在5～10、10～20 cm增加不明显,在0～5 cm退耕3 a、5 a、8 a后储量分别提高14.29%,35.71%和64.29%,各退耕年限0～20 cm TN平均固存率均为0.2 mg/(hm2.a);(2)退耕后各年限草地土壤活性有机碳(SAOC)含量有所增加,但各层含量变化不明显,其增加量远小于SOC的增加,说明退耕初期阶段积累了较多的土壤惰性碳;NO3-N含量增加明显,0～5、5～10 cm土壤各退耕年限含量达5%的显著性差异,但退耕前后NH4-N含量无明显变化.(3)土壤CO2通量与SOC含量、SAOC含量、TN含量及N2O通量显著正相关;N2O通量与SOC含量、矿质氮含量及CO2通量显著正相关.说明在环境因素稳定的条件下,退耕后土壤碳、氮含量的增加会导致CO2、N2O排放的加剧,表现出大气CO2、N2O的"源"效应.     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Elevated CO2 stimulates net accumulations of carbon and nitrogen in land ecosystems: a meta-analysis

The capability of terrestrial ecosystems to sequester carbon (C) plays a critical role in regulating future climatic change yet depends on nitrogen (N) availability. To predict long-term ecosystem C storage, it is essential to examine whether soil N becomes progressively limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. A critical parameter to indicate the long-term progressive N limitation (PNL) is net change in ecosystem N content in association with C accumulation in plant and soil pools under elevated CO2. We compiled data from 104 published papers that study C and N dynamics at ambient and elevated CO2. The compiled database contains C contents, N contents, and C:N ratio in various plant and soil pools, and root:shoot ratio. Averaged C and N pool sizes in plant and soil all significantly increase at elevated CO2 in comparison to those at ambient CO2, ranging from a 5% increase in shoot N content to a 32% increase in root C content. The C and N contents in litter pools are consistently higher in elevated than ambient CO2 among all the surveyed studies whereas C and N contents in the other pools increase in some studies and decrease in other studies. The high variability in CO2-induced changes in C and N pool sizes results from diverse responses of various C and N processes to elevated CO2. Averaged C:N ratios are higher by 3% in litter and soil pools and 11% in root and shoot pools at elevated relative to ambient CO2. Elevated CO2 slightly increases root:shoot ratio. The net N accumulation in plant and soil pools at least helps prevent complete down-regulation of, and likely supports, long-term CO2 stimulation of C sequestration. The concomitant C and N accumulations in response to rising atmospheric CO2 may reflect intrinsic nature of ecosystem development as revealed before by studies of succession over hundreds to millions of years.