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通量观测是定量描述土壤-植被-大气间物质循环和能量交换过程的基础。涡度相关技术作为直接测量植被冠层与大气间能量与物质交换通量的技术手段, 已经逐步发展成为国际通用的通量观测标准方法。随着涡度相关技术在全球碳水循环研究中的广泛应用, 长期连续的通量观测正在为准确评价生态系统碳固持能力、水分和能量平衡状况、生态系统对全球气候变化的反馈作用、区域和全球尺度模型的优化与验证、极端事件对生态系统结构与功能影响等方面的研究提供重要数据支撑和机制理解途径。通过站点尺度通量长期动态观测, 明确了不同气候区和植被类型生态系统碳水通量强度基线及其季节与年际变异特征。通过多站点联网观测, 在区域和全球尺度研究生态系统碳通量空间变异特征, 揭示了区域尺度上温度和降水对生态系统碳通量空间格局的生物地理学控制机制。该文概括地介绍了涡度相关技术的基本原理、假设与系统构成, 总结了涡度通量长期联网观测在陆地生态系统碳水通量研究中的主要应用, 并对通量研究发展前景进行了展望。 相似文献
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CHRISTIAN WILLE LARS KUTZBACH TORSTEN SACHS† DIRK WAGNER† EVA-MARIA PFEIFFER‡ 《Global Change Biology》2008,14(6):1395-1408
Eddy covariance measurements of methane flux were carried out in an arctic tundra landscape in the central Lena River Delta at 72°N. The measurements covered the seasonal course of mid‐summer to early winter in 2003 and early spring to mid‐summer in 2004, including the periods of spring thaw and autumnal freeze back. The study site is characterized by very cold and deep permafrost and a continental climate with a mean annual air temperature of ?14.7 °C. The surface is characterized by wet polygonal tundra, with a micro‐relief consisting of raised moderately dry sites, depressed wet sites, polygonal ponds, and lakes. We found relatively low fluxes of typically 30 mg CH4 m?2 day?1 during mid‐summer and identified soil temperature and near‐surface atmospheric turbulence as the factors controlling methane emission. The influence of atmospheric turbulence was attributed to the high coverage of open water surfaces in the tundra. The soil thaw depth and water table position were found to have no clear effect on methane fluxes. The excess emission during spring thaw was estimated to be about 3% of the total flux measured during June–October. Winter emissions were modeled based on the functional relationships found in the measured data. The annual methane emission was estimated to be 3.15 g m?2. This is low compared with values reported for similar ecosystems. Reason for this were thought to be the very low permafrost temperature in the study region, the sandy soil texture and low bio‐availability of nutrients in the soils, and the high surface coverage of moist to dry micro‐sites. The methane emission accounted for about 14% of the annual ecosystem carbon balance. Considering the global warming potential of methane, the methane emission turned the tundra into an effective greenhouse gas source. 相似文献
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Assessing the eddy covariance technique for evaluating carbon dioxide exchange rates of ecosystems: past, present and future 总被引:38,自引:0,他引:38
DENNIS D. BALDOCCHI 《Global Change Biology》2003,9(4):479-492
The eddy covariance technique ascertains the exchange rate of CO2 across the interface between the atmosphere and a plant canopy by measuring the covariance between fluctuations in vertical wind velocity and CO2 mixing ratio. Two decades ago, the method was employed to study CO2 exchange of agricultural crops under ideal conditions during short field campaigns. During the past decade the eddy covariance method has emerged as an important tool for evaluating fluxes of carbon dioxide between terrestrial ecosystems and the atmosphere over the course of a year, and more. At present, the method is being applied in a nearly continuous mode to study carbon dioxide and water vapor exchange at over a hundred and eighty field sites, worldwide. The objective of this review is to assess the eddy covariance method as it is being applied by the global change community on increasingly longer time scales and over less than ideal surfaces. The eddy covariance method is most accurate when the atmospheric conditions (wind, temperature, humidity, CO2) are steady, the underlying vegetation is homogeneous and it is situated on flat terrain for an extended distance upwind. When the eddy covariance method is applied over natural and complex landscapes or during atmospheric conditions that vary with time, the quantification of CO2 exchange between the biosphere and atmosphere must include measurements of atmospheric storage, flux divergence and advection. Averaging CO2 flux measurements over long periods (days to year) reduces random sampling error to relatively small values. Unfortunately, data gaps are inevitable when constructing long data records. Data gaps are generally filled with values produced from statistical and empirical models to produce daily and annual sums of CO2 exchange. Filling data gaps with empirical estimates do not introduce significant bias errors because the empirical algorithms are derived from large statistical populations. On the other hand, flux measurement errors can be biased at night when winds are light and intermittent. Nighttime bias errors tend to produce an underestimate in the measurement of ecosystem respiration. Despite the sources of errors associated with long‐term eddy flux measurements, many investigators are producing defensible estimates of annual carbon exchange. When measurements come from nearly ideal sites the error bound on the net annual exchange of CO2 is less than ±50 g C m?2 yr?1. Additional confidence in long‐term measurements is growing because investigators are producing values of net ecosystem productivity that are converging with independent values produced by measuring changes in biomass and soil carbon, as long as the biomass inventory studies are conducted over multiple years. 相似文献
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Impacts of tropospheric ozone and climate change on net primary productivity and net carbon exchange of China's forest ecosystems 总被引:1,自引:0,他引:1
Wei Ren Hanqin Tian Bo Tao Art Chappelka Ge Sun Chaoqun Lu Mingliang Liu Guangsheng Chen Xiaofeng Xu 《Global Ecology and Biogeography》2011,20(3):391-406
Aim We investigated how ozone pollution and climate change/variability have interactively affected net primary productivity (NPP) and net carbon exchange (NCE) across China's forest ecosystem in the past half century. Location Continental China. Methods Using the dynamic land ecosystem model (DLEM) in conjunction with 10‐km‐resolution gridded historical data sets (tropospheric O3 concentrations, climate variability/change, and other environmental factors such as land‐cover/land‐use change (LCLUC), increasing CO2 and nitrogen deposition), we conducted nine simulation experiments to: (1) investigate the temporo‐spatial patterns of NPP and NCE in China's forest ecosystems from 1961–2005; and (2) quantify the effects of tropospheric O3 pollution alone or in combination with climate variability and other environmental stresses on forests' NPP and NCE. Results China's forests acted as a carbon sink during 1961–2005 as a result of the combined effects of O3, climate, CO2, nitrogen deposition and LCLUC. However, simulated results indicated that elevated O3 caused a 7.7% decrease in national carbon storage, with O3‐induced reductions in NCE (Pg C year?1) ranging from 0.4–43.1% among different forest types. Sensitivity experiments showed that climate change was the dominant factor in controlling changes in temporo‐spatial patterns of annual NPP. The combined negative effects of O3 pollution and climate change on NPP and NCE could be largely offset by the positive fertilization effects of nitrogen deposition and CO2. Main conclusions In the future, tropospheric O3 should be taken into account in order to fully understand the variations of carbon sequestration capacity of forests and assess the vulnerability of forest ecosystems to climate change and air pollution. Reducing air pollution in China is likely to increase the resilience of forests to climate change. This paper offers the first estimate of how prevention of air pollution can help to increase forest productivity and carbon sequestration in China's forested ecosystems. 相似文献
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Yu. S. Ravkin V. P. Sedel’nikov M. G. Sergeev A. A. Titlyanova V. A. Khmelev I. N. Bogomolova S. M. Tsybulin 《Contemporary Problems of Ecology》2011,4(6):568-581
Results of classification of terrestrial ecosystems using an average similarity matrix are reported for the West Siberian Plain. Initial indices are first calculated separately for four components of an ecosystem. These components (blocks) include the underground block (soil humus, mortmass, and underground phytomass), above-ground vegetation, and invertebrates and vertebrates. Mismatch of boundaries in separate blocks of ecosystems and in comparison with the inhomogeneity of ecosystems in general was demonstrated. These differences are observed in both the typological and typological-chorological analysis. The indicated features of spatial succession within the blocks generate continuity of ecosystems and the conventional character of all the classifications and drawn boundaries. 相似文献
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BACKGROUND AND AIMS: Distinguishing between, and quantifying, the different components of ecosystem C fluxes is critical in predicting the responses of ecosystem C cycling to climate change. The aims of this study were to quantify the photosynthetic and respiratory fluxes of a 50-year-old Scots pine (Pinus sylvestris) ecosystem, and to distinguish respiration of branches with needles from that of stems, and that of soil. METHODS: The CO2 flux of the ecosystem was continuously measured using the eddy covariance (EC) method, and its components (respiration and photosynthesis of a branch with needles, stem and soil surface) were measured with an automated chamber system, from 2001 to 2004. KEY RESULTS: All values below are chamber based. The average temperature coefficient (Q10) of respiration was 2.7, 2.2 and 4.0, respectively, for branch (Rbran), stem (Rstem) and the soil surface (Rsoil). Respiration at a reference temperature of 15 degrees C (R15) was 1.27, 0.49 and 4.02 micromol CO2 m(-2) ground s(-1) for the three components, respectively. Over 4 years, the annual Rbran, Rstem and Rsoil ranged from 196 to 256, 56 to 83 and 439 to 598 g C m(-2) ground year(-1), respectively, with a 4-year average of 227, 72 and 507 g C m(-2) ground year(-1). Annual ecosystem respiration (Reco) was 731, 783, 909 and 751 g C m(-2) ground year(-1) in years 2001-2004, respectively, gross primary production (GPP) was 922, 1030, 1138 and 1001 g C m(-2) ground year(-1), and net ecosystem production (NEP) was 191, 247, 229 and 251 g C m(-2) ground year(-1). The average contribution of Rbran, Rstem and Rsoil to Reco was 29, 9 and 62 %, respectively. Overstorey photosynthesis accounted for 96 % of GPP. The average Reco/GPP ratio was 0.78. Net primary production (NPP) in the 4 years was 469, 581, 600 and 551 g C m(-2) year(-1), respectively, with the NPP/GPP ratio 0.54 averaged over the years. CONCLUSIONS: Respiration from the soil is the dominant component of ecosystem respiration. Differences between years in Reco were due to differences in temperature during the growing season. Rsoil was more sensitive to temperature than Rbran and Rstem, and differences in Rsoil were responsible for the differences in Reco between years. 相似文献
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湖北省主要森林类型生态系统生物量与碳密度比较 总被引:2,自引:0,他引:2
利用野外调查数据对湖北省封山育林下的次生林、次生林、人工林森林生态系统碳密度进行了分析,结果表明:封山育林下的次生林、次生林和人工林生态系统乔木层平均碳密度分别为133.87、73.42和111.62t·hm-2,灌木层平均碳密度分别为1.65、1.40和1.52t·hm-2,草本层平均碳密度分别为0.13、0.09和0.13t·hm-2,枯落物层平均碳密度分别为0.47、1.34和0.93t·hm-2,乔木层碳密度作为生态系统碳储量的主要贡献者占总生物碳密度的98.35%、96.29%和97.74%,林下植被(灌木层和草本层)碳密度分别占1.31%、1.95%和1.44%,凋落物层碳密度分别占0.34%、1.76%和0.82%。土壤(0~100cm)碳密度平均值分别为57.04、66.92和54.12t·hm-2,土壤碳密度的60%储存在0~40cm土壤中,并随土层深度增加,各层次土壤碳密度逐渐减少。森林生态系统的乔木层、灌木层、草本层、凋落物层生物量和土壤层碳密度均表现出:封山育林下的次生林、次生林大于人工林。封山育林下的次生林、次生林和人工林碳密度分布序列为土壤(0~100cm)>乔木层>灌木层>草本层>枯落物层。可见,封山育林下的次生林更有助于提高森林碳汇,实施近自然林经营是提升该区域森林碳汇能力的重要途径。 相似文献
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Ecosystems with high rates of nitrogen fixation often have high loss rates through leaching or possibly denitrification. However, there is no formal theoretical context to examine why this should be the case nor of how nitrogen accumulates in such open systems. Here, we propose a simple model coupling nitrogen inputs and losses to carbon inputs and losses. The nitrogen balance of this model system depends on plant (nitrogen fixer) growth rate, its carrying capacity, N fixed/C fixed, residence time of nitrogen and carbon in biomass, litter decay rate, litter N/C, and fractional loss rate of mineralized nitrogen. The model predicts the requirements for equilibrium in a nitrogen-fixing system, and the conditions on nitrogen fixation and losses in order for the system to accumulate nitrogen and carbon. In particular, the accumulation of nitrogen and carbon in a nitrogen-fixing system depend on an interaction between residence time in vegetation and litter decay rate in soil. To reflect a possible increased uptake of soil nitrogen and decreased respiratory cost of symbiotic nitrogen fixers, the model was then modified so that fixation rate decreased and growth rate increased as nitrogen capital accumulated. These modifications had only small effects on carbon and nitrogen accumulation. This suggests that switching from uptake of atmospheric nitrogen to mineral soil nitrogen as nitrogen capital accumulates simply results in a trade-off between energetic limitations and soil nitrogen limitations to carbon and nitrogen accumulation. Experimental tests of the model are suggested. 相似文献
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Biodiversity and the productivity and stability of ecosystems 总被引:2,自引:0,他引:2
Attempts to unveil the relationships between the taxonomic diversity, productivity and stability of ecosystems continue to generate inconclusive, contradictory and controversial conclusions. New insights from recent studies support the hypothesis that species diversity enhances productivity and stability in some ecosystems, but not in others. Appreciation is growing for the ways that particular ecosystem features, such as environmental variability and nutrient stress, can influence biotic interactions. Alternatives to the diversity-stability hypothesis have been proposed, and experimental approaches are starting to evolve to test these hypotheses and to elucidate the mechanisms underlying the functional role of species diversity. 相似文献
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David Schimel Ryan Pavlick Joshua B. Fisher Gregory P. Asner Sassan Saatchi Philip Townsend Charles Miller Christian Frankenberg Kathy Hibbard Peter Cox 《Global Change Biology》2015,21(5):1762-1776
Terrestrial ecosystem and carbon cycle feedbacks will significantly impact future climate, but their responses are highly uncertain. Models and tipping point analyses suggest the tropics and arctic/boreal zone carbon–climate feedbacks could be disproportionately large. In situ observations in those regions are sparse, resulting in high uncertainties in carbon fluxes and fluxes. Key parameters controlling ecosystem carbon responses, such as plant traits, are also sparsely observed in the tropics, with the most diverse biome on the planet treated as a single type in models. We analyzed the spatial distribution of in situ data for carbon fluxes, stocks and plant traits globally and also evaluated the potential of remote sensing to observe these quantities. New satellite data products go beyond indices of greenness and can address spatial sampling gaps for specific ecosystem properties and parameters. Because environmental conditions and access limit in situ observations in tropical and arctic/boreal environments, use of space‐based techniques can reduce sampling bias and uncertainty about tipping point feedbacks to climate. To reliably detect change and develop the understanding of ecosystems needed for prediction, significantly, more data are required in critical regions. This need can best be met with a strategic combination of remote and in situ data, with satellite observations providing the dense sampling in space and time required to characterize the heterogeneity of ecosystem structure and function. 相似文献
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C. S. Reynolds 《Journal of Aquatic Ecosystem Stress and Recovery (Formerly Journal of Aquatic Ecosystem Health)》1993,2(1):3-8
This paper summarizes the conceptual basis for ecosystemic water management, principally as it is discernible through a series of discussion papers contributed to the UN/ECE Seminar held in Oslo in May, 1991. The ecosystems approach seeks the objective management of water quality in lakes and river catchments, the sustainable exploitation of water resources (sensu lato) and the maintenance of biodiversity within aquatic catchments. It also seeks an attitude founded upon the sharing of habitat with other ecosystem components and the minimization of human impact. Moreover, there is no final condition: rather, there is an ethos always to improve performance. Emphasis is nevertheless placed on the unevolved, subclimactic state in which many ecosystems find themselves and the elasticity of structure which this imparts and which may, within limits, be exploited. Methods for assessing environmental quality and for measuring the performance of corrective management are briefly discussed. Future progress is suggested to be less than easy but proper ecosystemic attitudes and approaches are seen to be essential ingredients if past mistakes are to be overcome. 相似文献
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盐生荒漠净生态系统碳交换的涡度相关法和箱式法对比 总被引:1,自引:0,他引:1
将叶面积指数的季节动态,与箱式法同步观测得到的同化枝净光合(呼吸)速率和土壤呼吸速率相结合,对群落碳交换进行估算,并以此验证盐生荒漠涡度相关数据的可靠性。结果表明:盐生荒漠生态系统年叶片生物量为51.30±5.56 g·m-2,其中90.45%以上来源于多枝柽柳的贡献;而整个生长季,群落叶面积指数(LAI)呈单峰形式变化,从5月30日—9月30日,LAI介于0.180.30,并在第197天达到最大值。涡度相关法和箱式法对群落碳交换的测定结果表明,群落碳交换存在显著的季节变化,并于7月中旬达到碳同化峰值,与LAI有显著的相关性(P<0.001)。对比发现,两种测量方法对群落碳交换日过程的测定结果有很好的一致性,但对夜间生态系统呼吸的测定,涡度相关法较箱式法存在略微的低估,引起这种低估的原因可能是夜间湍流较弱。 相似文献
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Christopher S. Martens Thomas J. Shay† Howard P. Mendlovitz Daniel M. Matross‡ Scott R. Saleska‡ Steven C. Wofsy‡ W. Stephen Woodward Mary C. Menton §¶ José M. S. De Moura¶ Patrick M. Crill Osvaldo L. L. De Moraes†† Risonaldo L. Lima¶ 《Global Change Biology》2004,10(5):618-629
Radon‐222 (Rn‐222) is used as a transport tracer of forest canopy–atmosphere CO2 exchange in an old‐growth, tropical rain forest site near km 67 of the Tapajós National Forest, Pará, Brazil. Initial results, from month‐long periods at the end of the wet season (June–July) and the end of the dry season (November–December) in 2001, demonstrate the potential of new Rn measurement instruments and methods to quantify mass transport processes between forest canopies and the atmosphere. Gas exchange rates yield mean canopy air residence times ranging from minutes during turbulent daytime hours to greater than 12 h during calm nights. Rn is an effective tracer for net ecosystem exchange of CO2 (CO2 NEE) during calm, night‐time hours when eddy covariance‐based NEE measurements are less certain because of low atmospheric turbulence. Rn‐derived night‐time CO2 NEE (9.00±0.99 μmol m?2 s?1 in the wet season, 6.39±0.59 in the dry season) was significantly higher than raw uncorrected, eddy covariance‐derived CO2 NEE (5.96±0.51 wet season, 5.57±0.53 dry season), but agrees with corrected eddy covariance results (8.65±1.07 wet season, 6.56±0.73 dry season) derived by filtering out lower NEE values obtained during calm periods using independent meteorological criteria. The Rn CO2 results suggest that uncorrected eddy covariance values underestimate night‐time CO2 loss at this site. If generalizable to other sites, these observations indicate that previous reports of strong net CO2 uptake in Amazonian terra firme forest may be overestimated. 相似文献
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We used a combination of eddy flux, chamber and environmental measurements with an integrated suite of models to analyse the seasonality of net ecosystem carbon uptake (FCO2) in an 8-year-old, closed canopy Pinus radiata D.Don plantation in New Zealand (42°52′ S, 172°45′ E). The analyses utilized a biochemically based, big-leaf model of tree canopy photosynthesis (Ac), coupled to multiplicative environmental-constraint functions of canopy stomatal conductance (Gc) via environmental measurements, a temperature-dependent model of ecosystem respiration (Reco), and a soil water balance model. Available root zone water storage capacity at the measurement site is limited to about 50 mm for the very stony soil, and annual precipitation is only 660 mm, distributed evenly throughout the year. Accordingly the site is prone to soil moisture deficit throughout the summer. G c and Ac obtained maximum rates early in the growing season when plentiful soil water supply was associated with sufficient quantum irradiance (Qabs), and moderate air saturation deficit (D) and temperature (T). From late spring onwards, soil water deficit and D confined Gc and Ac congruously, which together with the solely temperature dependency of Reco resulted in the pronounced seasonality in FCO2. Reflecting a light-limitation of Ac in the closed canopy, modelled annual carbon (C) uptake was most sensitive to changes in Qabs. However, Qabs did not vary significantly between years, and changes in annual FCO2 were mostly due to variability in summer rainfall and D. Annual C-uptake of the forest was 717 g C m–2 in a near-average rainfall year, exceeding by one third the net uptake in a year with 20% less than average rainfall (515 g C m–2). 相似文献
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Karyotypes and morphology of three species with close relations toC. riihimakiensis Wülker 1971 are described from material of Tuva and Altai in Siberia. Only one of them could be reared to the male adult and was namedC. tuvanicus n.sp. 相似文献
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涡度相关观测的能量闭合状况及其对农田蒸散测定的影响 总被引:3,自引:0,他引:3
涡度相关法被认为是测定农田蒸散量的标准方法。然而,能量不闭合现象在涡度相关测量中普遍存在。分析能量不闭合对涡度相关观测的影响,对于提高涡度相关观测精度具有重要意义。以蒸渗仪法为参照,探讨涡度相关观测的能量闭合状况对农田蒸散测定的影响,在导致涡度相关观测能量不闭合的诸多因素中,寻找对蒸散测定有影响的因素。结果表明:涡度相关观测的白天能量平衡比率(EBR)呈秋冬高、春夏低的变化特征,麦季日均EBR范围在0.26—2.84之间,平均1.15;玉米季日均EBR范围在0.19—2.59之间,平均0.78。无论麦季或玉米季,涡度相关法测定的平均蒸散量(ETec)均明显低于蒸渗仪法观测值(ETL),但两者显著相关(P<0.01),并有相似的季节变化。平均蒸散比(ETec/ETL)麦季约为0.61,玉米季约为0.50。在冬小麦田和夏玉米田,ETec/ETL均与EBR显著相关(P<0.01)。麦田种植密度大,下垫面较均匀,蒸散比与EBR成正比(P<0.01),且不受叶面积指数(LAI)大小影响;反之,玉米田种植密度小,只有当LAI>1,下垫面变得较均匀后,蒸散比与EBR的关系才变得显著(P<0.01)。风速小时ETec/ETL与EBR显著相关,风速增加时二者相关性减弱。尤其在玉米田,当摩擦风速(u*)大于0.3 m/s时,ETec/ETL与EBR的相关性不再显著。风速小时,大气湍流微弱,湍流的涡旋较大。在有限的观测时段(0.5h)内,涡度相关仪的传感器难以捕捉足够的湍涡能量,所测湍流能量偏低,导致能量不闭合。以上结果为应用能量平衡比率校正农田蒸散提供了可能途径。 相似文献
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V. P. Sedel’nikov Yu. S. Ravkin A. A. Titlyanova I. N. Bogomolova O. N. Nikolaeva 《Contemporary Problems of Ecology》2011,4(3):229-240
Similarity coefficients were quantified for the ecosystems of the West Siberian Plain on the basis of the air-dry masses of dominant and subdominant plant species in five life forms (mosses, lichens, grasses, dwarf shrubs, shrubs, and trees) indicated in the formation ranges of the phytogeographic map of the West Siberian Plain. The resultant matrix was studied with cluster analysis and two classifications were made: one averaged over the formation ranges and the other over groups of the formation ranges. Expected informativity of the proposed classifications expressed in explained variance was 11–16% higher than that of the vegetation-based division used in the original phytogeographic map. 相似文献
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A new automated procedure for nanomolar nitrate analysis was applied to the study of uptake of low nitrate concentrations (< 100 ngat l–1) by phytoplankton. The precision of this analytical method (± 3 ngat l–1) made it possible to monitor the absorption of very low quantities of nitrate over short term periods by a low cell-density culture of the marine diatom Phaeodactylum tricornutum, where the levels of particulate nitrogen and chlorophyll were equivalent to those found in oligotrophic areas (0.5 µgat N l–1 and 0.4 µg l–1 respectively). By continuous monitoring of nitrate disappearance from the culture medium, we are able to describe accurately the transient uptake responses of the diatom after a spike addition of trace quantities of nitrate and thus to provide new information on the still largely unknown small-scale phenomenon of pulsed nitrate supply in the upper layer of stratified oceans and rapid uptake of these nitrate patches by phytoplankton.The results show that a N-limited culture of Phaeodactylum tricornutum is immediately capable of taking up trace quantities of nitrate (< 100 ngat l–1) at high rates (0.10–0.14 h–1) . These initial rates are one order of magnitude higher than the theoretical rates calculated from the Michaelis-Menten equation and are close to the level of V
max (0.15 h–1) obtained when cells are exposed to saturating nitrate concentrations. This rapid initial uptake would be a considerable advantage in oligotrophic areas where nanomolar nitrate supply is thought to be episodic. The present results suggest that phytoplankton evolve adaptations to utilize the available nitrate at the spatial and temporal scales at which it occurs. On the other hand, we can consider this physiological adaptation as evidence of nitrate pulses in the field which would invalidate the steady-state approach to the oligotrophic ecosystems. 相似文献