基于Biome-BGC模型和集合卡尔曼滤波方法的阔叶红松林生态系统水碳通量模拟

数据同化为模型与遥感观测结合提供了一条有效的途径,通过在模型运行过程中融入遥感观测数据,调整模型运行轨迹从而降低模型误差,提高模拟精度。本文利用集合卡尔曼滤波(En KF)算法同化生长季中分辨率成像光谱仪(MODIS)叶面积指数(LAI)与Biome-BGC模型模拟的LAI模拟长白山阔叶红松林的水碳通量。同时,通过改进模拟的雪面升华与土壤温度计算方法的参数,旨在降低冬季生态呼吸的模拟误差。结果表明,相对于原始模型,数据同化与模型改进后使得生态系统总初级生产力(GPP)的模拟值与观测值之间的相关系数提高0.06,中心化均方根误差(RMSE)降低0.48 g C·m~(-2)·d~(-1);生态系统呼吸(RE)的相关系数提高0.02,中心化均方根误差降低0.20 g C·m~(-2)·d~(-1);净生态系统碳交换量(NEE)相关系数提高0.35,中心化均方根误差降低0.50 g C·m~(-2)·d~(-1)。同时,数据同化对蒸散发(ET)的模拟精度没有显著影响,改进的模型提高了其相关系数。基于En KF算法的数据同化提高了长白山阔叶红松林碳通量模拟精度,对于精确估算区域碳通量有着重要的意义。     

中亚热带人工针叶林生态系统碳通量拆分差异分析

涡度通量观测可直接获取陆地生态系统与大气之间CO2净交换量(NEE),但深入认识碳循环过程和校验生态系统模型需要不同时间尺度总初级生产力(GPP)和生态系统呼吸(Re)等碳通量数据。利用中国陆地生态系统通量观测与研究网络(ChinaFLUX)中亚热带人工针叶林生态系统2003—2009年的涡度通量和气象观测数据,分析了两种NEE拆分方法对不同时间尺度GPP和Re评估的影响,结果表明:(1)两种拆分方法得到的生态系统碳通量组分(GPP和Re)的季节动态变化一致,都在生长季7、8月份达到峰值;(2)非线性回归模型拆分得到的全年Re和GPP相较于光响应曲线模型分别高出2%—28.6%和1.6%—23%,最大高出317.6 gC·m-2·a-1(2006年),逐月最大差值主要发生在8、9月份;(3)不同时间尺度上,两种方法拆分得到的GPP和Re之间差值的环境响应因子不同。在广泛采用非线性回归模型进行拆分时,如果当月光合有效辐射接近到905mol·m-2·月-1,月平均空气饱和水汽压差接近1.18 kPa时,需要考虑使用光响应曲线模型拆分该月通量,结合两种拆分方法以减小全年的误差。     

涡度相关观测的太湖CH4通量数据插补方法评价

涡度相关法是在湖泊开展CH₄通量长期连续观测的重要方法。受到多种因素的影响,CH₄通量观测数据存在大量缺失。为重构完整的CH₄通量时间序列,就需要适宜的数据插补方法。本研究利用太湖涡度通量观测网络东部的避风港站点2014—2017年的常规气象数据及涡度相关观测的CH₄通量数据,在分析半小时尺度以及日尺度CH₄通量影响要素的基础上,测试了非线性回归法以及随机森林算法和误差反向传播算法在半小时尺度及日尺度上插补CH₄通量缺失数据的可行性。结果表明: 在半小时尺度上,避风港站生长季CH₄通量主要受到底泥温度、摩擦风速、气温、相对湿度、潜热通量和20 cm处水温的影响,非生长季主要受到相对湿度、潜热通量、风速、感热通量和底泥温度的影响,而在日尺度上CH₄通量主要受潜热通量和相对湿度的影响。在对CH₄通量缺失数据的插补中,随机森林模型在所有时间尺度上都表现为最佳的插补性能,其中,将日序、太阳高度角、底泥温度、摩擦风速、气温、20 cm处水温、相对湿度、气压和风速作为输入变量的随机森林模型更适用于半小时尺度缺失数据的插补;将日序、底泥温度、摩擦风速、气温、20 cm处水温、相对湿度、气压、风速和向下短波辐射作为输入变量的随机森林模型更适用于日尺度缺失数据的插补;整体上,插补模型对日尺度缺失数据的插补优于半小时尺度。     

双集合卡尔曼滤波LAI同化结合BEPS模型的竹林生态系统碳通量模拟

叶面积指数(LAI)是森林生态系统碳循环研究的重要观测数据,也是驱动森林生态系统模型模拟碳循环的重要参数.本文以毛竹林和雷竹林为研究对象,首先利用双集合卡尔曼滤波,同化两种竹林生态系统观测站点2014—2015年MODIS LAI时间序列数据,然后将同化的高质量毛竹LAI和雷竹LAI作为输入数据驱动BEPS模型,模拟两种竹林生态系统总初级生产力(GPP)、净生态系统碳交换量(NEE)和总生态系统呼吸(TER)等碳循环数据,并用通量站实际观测值评价模拟结果;另外,还对比不同质量LAI对碳循环模拟的影响.结果表明: 双集合卡尔曼滤波同化得到的毛竹林和雷竹林LAI与实测LAI之间的相关关系极为显著,R²分别为0.81和0.91,且均方根误差和绝对偏差均较小,极大地提高了MODIS LAI的产品精度;在同化得到的LAI驱动下,BEPS模型模拟的毛竹林GPP、NEE和TER与实际观测值之间的R²分别为0.66、0.47和0.64,雷竹林分别为0.66、0.45和0.73,模拟结果均好于三次样条帽盖算法平滑LAI模拟得到的GPP、NEE和TER,其中,毛竹林、雷竹林NEE的模拟精度提高幅度最大,分别为11.2%和11.8%.     

基于FLUXNET的CLM模型总初级生产力模拟评价与误差分析

总初级生产力(gross primary productivity,GPP)的准确估计是陆地生态系统碳循环研究以及未来气候变化预测的基础。本文利用全球通量网的通量观测数据评估了CLM4.5的GPP模拟效果,结合叶面积指数(leaf area index,LAI)观测数据分析了CLM4.5模型GPP模拟误差的主要原因,并对CLM4.5模型主要的光合参数进行敏感性分析,探讨改善光合作用模拟精度的可能途径。结果表明:CLM4.5对GPP的模拟效果优于CLM4,月尺度和年尺度GPP模拟值的平均绝对偏差分别降低15%和29%。但与观测值相比,CLM4.5模拟的GPP年总量仍然具有较大偏差,平均偏差为366.06 g C·m-2·a-1。不同植被功能型GPP模拟误差具有不同的季节变化特征,误差主要发生在春季和夏季。冠层顶部比叶面积、叶片碳氮比和叶片氮含量中Rubisco氮含量所占比例是GPP模拟的3个敏感参数。未来应主要从物候期及LAI模拟的改进、磷循环过程的影响、生态系统水平光合参数集的构建等方面实现对GPP模拟精度的提高。     

利用大气二氧化碳和羰基硫浓度评估陆地生态系统模型碳通量模拟的不确定性

生态系统模型是模拟全球陆地生态系统碳循环的重要工具,但是其在全球不同区域的模拟存在很大的不确定性。如何评估陆地生态系统模型的不确定性是一项重要的研究。以北美地区为例,利用8个高塔观测站点同步获取的大气CO_2和羰基硫(OCS)浓度数据,结合WRF-STILT大气粒子扩散模型,评估了CASA-GFED3、SiB3和SiBCASA三种陆地生态系统模型模拟总初级生产力(GPP)和净生态系统CO_2交换(NEE)通量的不确定性。结果表明,SiB3模型能很好地模拟北美陆地生态系统GPP和NEE的季节变化时相和幅度,在3种模型中具有最佳的模拟能力;CASA-GFED3模型模拟的NEE季节变化较为理想、但对生长季GPP的模拟存在较大的误差,SiBCASA模型在模拟冬季晚期和春季早期的NEE和GPP时表现较不理想。研究证明了大气CO_2和OCS在评估陆地生态系统模型碳通量模拟的不确定性中的作用,为利用大气CO_2和OCS观测数据优化计算陆地生态系统光合和呼吸碳通量提供了理论支撑。     

雨养玉米农田生态系统的蒸散特征及其作物系数

基于雨养玉米农田生态系统2007年整个生长季的涡度相关通量资料,对蒸散的日、季动态进行分析.结果表明：玉米农田生态系统蒸散的日、季动态均呈单峰型变化,最大值分别出现在12:00左右和7月.结合修正的Penman-Monteith公式与相应的生态、气象观测要素,对作物系数(K指数)影响因子的分析结果表明,K指数主要受叶面积指数(LAI)、气温(T_a)、净辐射(R_n)以及表层土壤含水量的影响.在此基础上,初步建立了半小时尺度的作物系数(K指数)模型.     

温带阔叶红松林生态系统潜热通量模拟——气孔导度组合模型在Shuttleworth-Wallace双源模型中的应用

叶片水平的气孔导度组合模型已被成功扩展到冠层水平,并被应用于冬小麦生态系统潜热通量的模拟研究,但该研究仅基于1a的数据,有必要研究模型在更长时间尺度和其它生态系统类型的适用性.以长白山阔叶红松林(CBS)为研究对象,将组合模型进一步应用于Shuttleworth-Wallace双源模型,模拟了CBS 3a生长季内的潜热通量,利用涡度相关系统观测的潜热通量数据对模型进行验证,并对比了双源模型与单源模型的模拟结果.结果显示,双源模型较单源模型能取得更高的模拟精度,生长季不同时期的潜热通量模拟值和实测值的日变化较一致.对双源模型模拟值和实测潜热通量的相关分析显示,二者直线回归斜率和R2分别为0.96和0.72.对长白山阔叶红松林生态系统的蒸散和植被蒸腾的季节和年际变异分析发现,影响冠层蒸散和植被蒸腾季节动态的主要因素是饱和差和辐射,而影响它们年际动态的主要因素则是饱和差和温度.     

基于模型数据融合的千烟洲亚热带人工林碳水通量模拟

人工林生态系统是我国森林生态系统的重要组成部分,在全球碳平衡中的作用越来越受到重视.利用千烟洲亚热带人工针叶林通量观测站的碳水通量和气象观测数据,通过模型数据融合方法对碳水循环过程模型——SIPNET模型关键参数进行反演,模拟了2004-2009年千烟洲人工林生态系统的碳水通量.结果表明:仅用碳通量观测数据优化模型参数时,净生态系统碳交换量(NEE)模拟效果较好(R2=0.934),而生态系统蒸散(ET)模拟效果较差(R2=0.188);同时用碳水通量观测数据优化时,NEE模拟效果稍差(R2=0.929),但ET模拟效果显著提升(R2=0.824),说明利用碳水通量观测数据同时优化,SIPNET模型才能较好地模拟试验站点碳水通量.在此基础上,开展了人工林生态系统碳通量对降水变化响应的敏感性分析,发现降水量减少对光合作用的影响比对呼吸作用的影响更为强烈,且碳水通量同时参与优化时模型才能较好地模拟碳通量随降水减少而快速降低的趋势,表明如果不能同时利用碳水通量进行参数优化,模型无法正确揭示生态系统碳循环对降水变异的响应.     

3PG碳生产模型在长白山阔叶红松林总初级生产力估算中的应用

利用ChinaFLUX长白山站阔叶红松林的通量观测数据以及同期卫星遥感数据,对3PG模型中的植被光合模型(VPM)、光能利用率模型(EC-LUE)、陆地生态系统模型(TEM)、卡内基-埃姆斯-斯坦福方法模型(CASA)4种模型进行参数重组,通过对比通量观测值与估算值的均方根误差、决定系数及平均误差确定模型的最适合参数;并利用实测的通量观测数据对优化后的模型进行拟合度验证,以提高其估算长白山阔叶红松林总初级生产力(GPP)的准确性.结果表明: 采用温度、增强植被指数、地表水分指数分别表征原模型中的温度限制因子、光合有效辐射吸收比例、水分限制因子估算长白山阔叶红松林GPP时,结果最优,优化后模型的精度(R²=0.948,RMSE=0.035 mol·m^-2·month^-1)明显优于原模型(R²=0.854,RMSE=0.177 mol·m^-2·month^-1),且能够有效改善原模型生长季明显高估的现象;通过敏感性分析可知,温度是对GPP估算不确定性影响最大的参数,其次为增强型植被指数和光合有效辐射,地表水分指数最小,且变量间的交互作用对GPP估算不确定性也存在影响.     

Tuning of functional heme reduction potentials in Shewanella fumarate reductases

The fumarate reductases from S. frigidimarina NCIMB400 and S. oneidensis MR-1 are soluble and monomeric enzymes located in the periplasm of these bacteria. These proteins display two redox active domains, one containing four c-type hemes and another containing FAD at the catalytic site. This arrangement of single-electron redox co-factors leading to multiple-electron active sites is widespread in respiratory enzymes. To investigate the properties that allow a chain of single-electron co-factors to sustain the activity of a multi-electron catalytic site, redox titrations followed by NMR and visible spectroscopies were applied to determine the microscopic thermodynamic parameters of the hemes. The results show that the redox behaviour of these fumarate reductases is similar and dominated by a strong interaction between hemes II and III. This interaction facilitates a sequential transfer of two electrons from the heme domain to FAD via heme IV.     

Spatio-temporal variability in the photosynthetic characteristics of Zostera tasmanica measured by PAM

Rapid light curves (RLCs), based on pulse amplitude modulated (PAM) fluorometry, were used to investigate the spatio-temporal variability in photosynthesis versus irradiance parameters (α, I_k and P_max) and the F_v/F_m ratio of the seagrass Zostera tasmanica (formerly Heterozostera tasmanica). Spatial variation was examined across scales ranging from within a leaf (cms) to across the bed (ms), using a nested analysis of covariance sampling design. Overall, significant variation was identified at all scales examined, excluding the largest scale (area). Patterns of variability differed among individual parameters; however a high percentage of the variation was consistently assigned to the covariates, age (within and between leaves) for all parameters, except P_max.     

Water deficit in field-grown Gossypium hirsutum primarily limits net photosynthesis by decreasing stomatal conductance,increasing photorespiration,and increasing the ratio of dark respiration to gross photosynthesis

Much effort has been expended to improve irrigation efficiency and drought tolerance of agronomic crops; however, a clear understanding of the physiological mechanisms that interact to decrease source strength and drive yield loss has not been attained. To elucidate the underlying mechanisms contributing to inhibition of net carbon assimilation under drought stress, three cultivars of Gossypium hirsutum were grown in the field under contrasting irrigation regimes during the 2012 and 2013 growing season near Camilla, Georgia, USA. Physiological measurements were conducted on three sample dates during each growing season (providing a broad range of plant water status) and included, predawn and midday leaf water potential (Ψ_PD and Ψ_MD), gross and net photosynthesis, dark respiration, photorespiration, and chlorophyll a fluorescence. End-of-season lint yield was also determined. Ψ_PD ranged from −0.31 to −0.95 MPa, and Ψ_MD ranged from −1.02 to −2.67 MPa, depending upon irrigation regime and sample date. G. hirsutum responded to water deficit by decreasing stomatal conductance, increasing photorespiration, and increasing the ratio of dark respiration to gross photosynthesis, thereby limiting P_N and decreasing lint yield (lint yield declines observed during the 2012 growing season only). Conversely, even extreme water deficit, causing a 54% decline in P_N, did not negatively affect actual quantum yield, maximum quantum yield, or photosynthetic electron transport. It is concluded that P_N is primarily limited in drought-stressed G. hirsutum by decreased stomatal conductance, along with increases in respiratory and photorespiratory carbon losses, not inhibition or down-regulation of electron transport through photosystem II. It is further concluded that Ψ_PD is a reliable indicator of drought stress and the need for irrigation in field-grown cotton.     

Thermoregulation in male temperate bats depends on habitat characteristics

Several energy-saving strategies have evolved in animals, one example being the short-term reduction of metabolism and body temperature (torpor) in endotherms. For bats, pronounced torpor behaviour has been described. The aim of this study was to assess individual variation in torpor expression of male Myotis daubentonii, and to analyse whether this variation is related to habitat characteristics. For that we measured skin temperatures of bats from different habitats using radio transmitters and also recorded ambient temperature. Skin temperature was corrected for ambient temperature and individual body mass. Cluster analysis of residuals revealed two different thermoregulatory strategies. Males in cluster 1 were more often encountered torpid and reached lower minimum skin temperatures than males in cluster 2. The differences in behaviour were related to environmental variables (water surface area near the roost, roost altitude, precipitation, ambient temperature in the warmest quarter of the year). Males from cluster 1 occupied less favourable habitats (less water surface, higher altitudes, wetter and colder climate) than males from cluster 2. Our data suggest a linkage between torpor behaviour and habitat characteristics. These characteristics could be used to identify favourable and marginal habitats for M. daubentonii.     

Electron transfer kinetics between soluble modules of Paracoccus denitrificans cytochrome c1 and its physiological redox partners

The transient electron transfer (ET) interactions between cytochrome c₁ of the bc₁-complex from Paracoccus denitrificans and its physiological redox partners cytochrome c₅₅₂ and cytochrome c₅₅₀ have been characterized functionally by stopped-flow spectroscopy. Two different soluble fragments of cytochrome c₁ were generated and used together with a soluble cytochrome c₅₅₂ module as a model system for interprotein ET reactions. Both c₁ fragments lack the membrane anchor; the c₁ core fragment (c_1CF) consists of only the hydrophilic heme-carrying domain, whereas the c₁ acidic fragment (c_1AF) additionally contains the acidic domain unique to P. denitrificans. In order to determine the ionic strength dependencies of the ET rate constants, an optimized stopped-flow protocol was developed to overcome problems of spectral overlap, heme autoxidation and the prevalent non-pseudo first order conditions. Cytochrome c₁ reveals fast bimolecular rate constants (10⁷ to 10⁸ M^− 1 s^− 1) for the ET reaction with its physiological substrates c₅₅₂ and c₅₅₀, thus approaching the limit of a diffusion-controlled process, with 2 to 3 effective charges of opposite sign contributing to these interactions. No direct involvement of the N-terminal acidic c₁-domain in electrostatically attracting its substrates could be detected. However, a slight preference for cytochrome c₅₅₀ over c₅₅₂ reacting with cyochrome c₁ was found and attributed to the different functions of both cytochromes in the respiratory chain of P. denitrificans.     

Thermodynamic and kinetic characterisation of individual haems in multicentre cytochromes c3

The characterisation of individual centres in multihaem proteins is difficult due to the similarities in the redox and spectroscopic properties of the centres. NMR has been used successfully to distinguish redox centres and allow the determination of the microscopic thermodynamic parameters in several multihaem cytochromes c₃ isolated from different sulphate-reducing bacteria. In this article we show that it is also possible to discriminate the kinetic properties of individual centres in multihaem proteins, if the complete microscopic thermodynamic characterisation is available and the system displays fast intramolecular equilibration in the time scale of the kinetic experiment. The deconvolution of the kinetic traces using a model of thermodynamic control provides a reference rate constant for each haem that does not depend on driving force and can be related to structural factors. The thermodynamic characterisation of three tetrahaem cytochromes and their kinetics of reduction by sodium dithionite are reported in this paper. Thermodynamic and kinetic data were fitted simultaneously to a model to obtain microscopic reduction potentials, haem-haem and haem-proton interacting potentials, and reference rate constants for the haems. The kinetic information obtained for these cytochromes and recently published data for other multihaem cytochromes is discussed with respect to the structural factors that determine the reference rates. The accessibility for the reducing agent seems to play an important role in controlling the kinetic rates, although is clearly not the only factor.     

The use of small subcutaneous transponders for quantifying thermal biology and torpor in small mammals

Remote measurements of body temperature (T_b) in animals require implantation of relatively large temperature-sensitive radio-transmitters or data loggers, whereas rectal temperature (T_rec) measurements require handling and therefore may bias the results. We investigated whether ∼0.1 g temperature-sensitive subcutaneously implanted transponders can be reliably used to quantify thermal biology and torpor use in small mammals. We examined (i) the precision of transponder readings as a function of temperature and (ii) whether subcutaneous transponders can be used to remotely record subcutaneous temperature (T_sub). Five adult male dunnarts (Sminthopsis macroura, body mass 24 g) were implanted with subcutaneous transponders to determine T_sub as a function of time and ambient temperature (T_a), and in comparison to thermocouple readings of T_rec. Transponder temperature was highly correlated with water bath temperature (r²=0.96–0.99) over a range of approximately 10.0–40.0 °C. Transponders provided reliable data (±0.6 °C) over the T_sub of 21.4–36.9 °C and could be read from a distance of up to 5 cm. Below 21.4 °C, accuracy was reduced to ±2.8 °C, but individual transponder accuracy varied. Consequently, small subcutaneous transponders are useful to remotely quantify thermal physiology and torpor patterns without having to disturb the animal and disrupt torpor. Even at T_sub<21.4 °C where the accuracy of the temperature readings was reduced, transponders do provide reliable data on whether and when torpor is used.     

Determination of catalase activity using chromogenic probe involving iso-nicotinicacidhydrazide and pyrocatechol

A biocatalatic pathway involving chromogenic probe has been proposed for the determination of catalase activity by means of iso-nicotinicacidhydrazide (INH) and pyrocatechol (PC). The assay is based on the enzymatic consumption of hydrogen peroxide using INH-PC system. The response of the catalase activity was ascertained by the rate of the reaction involving 14.10 mM H₂O₂. On addition of H₂O₂, INH-PC indicator system formed a chromogenic product with absorbance maxima at 490 nm. Hence the activity of catalase was directly measured by the chromogenic response in the formation of the coupled product. The catalase assay was elaborated by the kinetic response of the INH-PC system. The linearity of the catalase activity and H₂O₂ was in the range 0.2-7.0 units and 1.76-7.0 mM, respectively in 3 ml solution. The catalytic efficiency and catalytic power were calculated. The Michaelis-Menten constant of INH, PC and H₂O₂ were found to be 0.344, 0.176 and 8.82 mM, respectively. The indicator reaction was applied in the determination of catalase activity in mycelia mats and culture media.     

Crystal structure of the 16 heme cytochrome from Desulfovibrio gigas: a glycosylated protein in a sulphate-reducing bacterium

Sulphate-reducing bacteria have a wide variety of periplasmic cytochromes involved in electron transfer from the periplasm to the cytoplasm. HmcA is a high molecular mass cytochrome of 550 amino acid residues that harbours 16 c-type heme groups. We report the crystal structure of HmcA isolated from the periplasm of Desulfovibrio gigas. Crystals were grown using polyethylene glycol 8K and zinc acetate, and diffracted beyond 2.1 A resolution. A multiple-wavelength anomalous dispersion experiment at the iron absorption edge enabled us to obtain good-quality phases for structure solution and model building. DgHmcA has a V-shape architecture, already observed in HmcA isolated from Desulfovibrio vulgaris Hildenborough. The presence of an oligosaccharide molecule covalently bound to an Asn residue was observed in the electron density maps of DgHmcA and confirmed by mass spectrometry. Three modified monosaccharides appear at the highly hydrophobic vertex, possibly acting as an anchor of the protein to the cytoplasmic membrane.