Integrating in-situ measurements, land surface models and satellite remote sensing to understand impacts of environmental changes on terrestrial ecosystem processes at multiple scales

How terrestrial ecosystems respond to environmental changes affects the well-being of human society. Thus, extreme climate events, increasing the atmospheric concentration of CO₂, and drastic changes in temperature are sources of major concern. However, our current capacity to understand and predict these responses is still limited because a myriad of physical, chemical, and biological processes are involved. While many mechanistic-based land surface models have been developed, their performances remain relatively poor and require continuous improvement. While ground-based and space-based observational datasets of the surface of the Earth have been available for a long time, their linkages to the functional aspects of the processes in terrestrial ecosystems often are weak. In this study, I used the approach of integrating in-situ measurements, land surface models, and remote sensing by satellites. I hypothesized that, through such integration, the impacts of environmental changes on terrestrial processes at multiple scales could be better understood and even predicted, especially the impacts related to the functions of important ecosystems. I tested this hypothesis at the local, regional, and global scales. At the local scale, i.e., at a Midwest forest site known as the isoprene volcano of the world, I examined the effects of droughts on the emissions of isoprene, which is the most abundant, non-methane, biogenic volatile organic compound. I compared flux tower observations with simulations performed by a state-of-the-art land model (CLM) coupled with the model of emissions of gases and aerosols from Nature version 2.1 (MEGAN2.1), and I used these observations to develop an understanding of how the amount of moisture in the soil and the ambient temperature affect the prediction of isoprene emissions during droughts. I found that temperature had a delaying effect on isoprene emissions, which are sensitive to variations in the moisture content of the soil. Thus, during drought conditions, both the delaying effect and the sensitivity to moisture are overlooked by the model. A better model that does not have these two shortcomings is required for realistic predictions of isoprene emissions. At the regional scale, I investigated the potential of using sun-induced chlorophyll fluorescence (SIF) retrieved from a satellite to monitor vegetation activities in an arid region and a semi-arid region in Australia. I chose these two types of regions for this investigation because the ecosystems in such regions have important effects on the global carbon cycle, while their contributions are poorly constrained in global carbon budgets. I found that SIF was synchronized better with the activity of vegetation than other indices that are commonly used for this purpose. I quantified the relationships between the various activities of plants and the amount and frequency of precipitation, and I was able to demonstrate that, over the region being studied, SIF represented the activity of vegetation in response to the availability of water better than other, remotely-sensed variables. At the global scale, I used multiple model ensembles to determine the climatic and anthropogenic controls on the terrestrial evapotranspiration trends from 1982 to 2010. After climatic influences, increases in CO₂ were found to be the second-most dominant factor that affected the trend of ET. CO₂ causes a decreasing trend in the canopy’s transpiration and ET, and this is especially of concern for tropical forests and high-latitude shrub lands. The increased deposition of nitrogen amplifies the global ET slightly due to enhanced growth of plants. On a global scale, land-use-induced ET responses are minor, but they can be significant locally, particularly over regions with intensive changes in the land-cover. The results of my studies demonstrated that integrating in-situ measurements, models of the surface on the land, and remote sensing using satellites can provide insights regarding the impacts of environmental changes on terrestrial processes at multiple scales. This approach is particularly important when models are imperfect and observations are lacking. My findings indicated ways that future models can be improved and identified key observational needs for the functions of terrestrial ecosystems.Geological Science

New Methods for Measurements of Photosynthesis from Space

Our ability to close the Earth's carbon budget and predict feedbacks in a warming climate depends critically on knowing where, when, and how carbon dioxide (CO2) is exchanged between the land and atmosphere. In particular, determining the rate of carbon fixation by the Earth's biosphere (commonly referred to as gross primary productivity, or GPP) and the dependence of this productivity on climate is a central goal. Historically, GPP has been inferred from spectral imagery of the land and ocean. Assessment of GPP from the color of the land and ocean requires, however, additional knowledge of the types of plants in the scene, their regulatory mechanisms, and climate variables such as soil moisture—just the independent variables of interest! Sunlight absorbed by chlorophyll in photosynthetic organisms is mostly used to drive photosynthesis, but some can also be dissipated as heat or re‐radiated at longer wavelengths (660–800 nm). This near‐infrared light re‐emitted from illuminated plants is termed solarinduced fluorescence (SIF), and it has been found to strongly correlate with GPP. To advance our understanding of SIF and its relation to GPP and environmental stress at the planetary scale, the Keck Institute for Space Studies (KISS) convened a workshop—held in Pasadena, California, in August 2012—to focus on a newly developed capacity to monitor chlorophyll fluorescence from terrestrial vegetation by satellite. This revolutionary approach for retrieving global observations of SIF promises to provide direct and spatially resolved information on GPP, an ideal bottom‐up complement to the atmospheric net CO2 exchange inversions. Workshop participants leveraged our efforts on previous studies and workshops related to the European Space Agency’s FLuorescence EXplorer (FLEX) mission concept, which had already targeted SIF for a possible satellite mission and had developed a vibrant research community with many important publications. These studies, mostly focused on landscape, canopy, and leaf‐level interpretation, provided the ground‐work for the workshop, which focused on the global carbon cycle and synergies with atmospheric net flux inversions. Workshop participants included key members of several communities: plant physiologists with experience using active fluorescence methods to quantify photosynthesis; ecologists and radiative transfer experts who are studying the challenge of scaling from the leaf to regional scales; atmospheric scientists with experience retrieving photometric information from space‐borne spectrometers; and carbon cycle experts who are integrating new observations into models that describe the exchange of carbon between the atmosphere, land and ocean. Together, the participants examined the link between “passive” fluorescence observed from orbiting spacecraft and the underlying photochemistry, plant physiology and biogeochemistry of the land and ocean. This report details the opportunity for forging a deep connection between scientists doing basic research in photosynthetic mechanisms and the more applied community doing research on the Earth System. Too often these connections have gotten lost in empiricism associated with the coarse scale of global models. Chlorophyll fluorescence has been a major tool for basic research in photosynthesis for nearly a century. SIF observations from space, although sensing a large footprint, probe molecular events occurring in the leaves below. This offers an opportunity for direct mechanistic insight that is unparalleled for studies of biology in the Earth System. A major focus of the workshop was to review the basic mechanisms that underlie this phenomenon, and to explore modeling tools that have been developed to link the biophysical and biochemical knowledge of photosynthesis with the observable—in this case, the radiance of SIF—seen by the satellite. Discussions led to the identification of areas where knowledge is still lacking. For example, the inability to do controlled illumination observations from space limits the ability to fully constrain the variables that link fluorescence and photosynthesis. Another focus of the workshop explored a “top‐down” view of the SIF signal from space. Early studies clearly identified a strong correlation between the strength of this signal and our best estimate of the rate of photosynthesis (GPP) over the globe. New studies show that this observation provides improvements over conventional reflectance‐based remote sensing in detecting seasonal and environmental (particularly drought related) modulation of photosynthesis. Apparently SIF responds much more quickly and with greater dynamic range than typical greenness indices when GPP is perturbed. However, discussions at the workshop also identified areas where top‐down analysis seemed to be “out in front” of mechanistic studies. For example, changes in SIF based on changes in canopy light interception and the light use efficiency of the canopy, both of which occur in response to drought, are assumed equivalent in the top‐down analysis, but the mechanistic justification for this is still lacking from the bottom‐up side. Workshop participants considered implications of these mechanistic and empirical insights for large‐scale models of the carbon cycle and biogeochemistry, and also made progress toward incorporating SIF as a simulated output in land surface models used in global and regional‐scale analysis of the carbon cycle. Comparison of remotely sensed SIF with modelsimulated SIF may open new possibilities for model evaluation and data assimilation, perhaps leading to better modeling tools for analysis of the other retrieval from GOSAT satellite, atmospheric CO2 concentration. Participants also identified another application for SIF: a linkage to the physical climate system arising from the ability to better identify regional development of plant water stress. Decreases in transpiration over large areas of a continent are implicated in the development and “locking‐in” of drought conditions. These discussions also identified areas where current land surface models need to be improved in order to enable this research. Specifically, the radiation transport treatments need dramatic overhauls to correctly simulate SIF. Finally, workshop participants explored approaches for retrieval of SIF from satellite and ground‐based sensors. The difficulty of resolving SIF from the overwhelming flux of reflected sunlight in the spectral region where fluorescence occurs was once a major impediment to making this measurement. Placement of very high spectral resolution spectrometers on GOSAT (and other greenhouse gas–sensing satellites) has enabled retrievals based on infilling of solar Fraunhofer lines, enabling accurate fluorescence measurements even in the presence of moderately thick clouds. Perhaps the most interesting challenge here is that there is no readily portable ground‐based instrumentation that even approaches the capability of GOSAT and other planned greenhouse gas satellites. This strongly limits scientists’ ability to conduct ground‐based studies to characterize the footprint of the GOSAT measurement and to conduct studies of radiation transport needed to interpret SIF measurement. The workshop results represent a snapshot of the state of knowledge in this area. New research activities have sprung from the deliberations during the workshop, with publications to follow. The introduction of this new measurement technology to a wide slice of the community of Earth System Scientists will help them understand how this new technology could help solve problems in their research, address concerns about the interpretation, identify future research needs, and elicit support of the wider community for research needed to support this observation. Somewhat analogous to the original discovery that vegetation indices could be derived from satellite measurements originally intended to detect clouds, the GOSAT observations are a rare case in which a (fortuitous) global satellite dataset becomes available before the research community had a consolidated understanding on how (beyond an empirical correlation) it could be applied to understanding the underlying processes. Vegetation indices have since changed the way we see the global biosphere, and the workshop participants envision that fluorescence can perform the next indispensable step by complementing these measurements with independent estimates that are more indicative of actual (as opposed to potential) photosynthesis. Apart from the potential FLEX mission, no dedicated satellite missions are currently planned. OCO‐2 and ‐3 will provide much more data than GOSAT, but will still not allow for regional studies due to the lack of mapping capabilities. Geostationary observations may even prove most useful, as they could track fluorescence over the course of the day and clearly identify stress‐related down‐regulation of photosynthesis. Retrieval of fluorescence on the global scale should be recognized as a valuable tool; it can bring the same quantum leap in our understanding of the global carbon cycle as vegetation indices once did

Satellite-detected fluorescence reveals global physiology of ocean phytoplankton

© 2009 The Authors. This is an open-access article distributed under the terms of the Creative Commons Attribution 3.0 License. The definitive version was published in Biogeosciences 6 (2009): 779-794, doi: 10.5194/bg-6-779-2009Phytoplankton photosynthesis links global ocean biology and climate-driven fluctuations in the physical environment. These interactions are largely expressed through changes in phytoplankton physiology, but physiological status has proven extremely challenging to characterize globally. Phytoplankton fluorescence does provide a rich source of physiological information long exploited in laboratory and field studies, and is now observed from space. Here we evaluate the physiological underpinnings of global variations in satellite-based phytoplankton chlorophyll fluorescence. The three dominant factors influencing fluorescence distributions are chlorophyll concentration, pigment packaging effects on light absorption, and light-dependent energy-quenching processes. After accounting for these three factors, resultant global distributions of quenching-corrected fluorescence quantum yields reveal a striking consistency with anticipated patterns of iron availability. High fluorescence quantum yields are typically found in low iron waters, while low quantum yields dominate regions where other environmental factors are most limiting to phytoplankton growth. Specific properties of photosynthetic membranes are discussed that provide a mechanistic view linking iron stress to satellite-detected fluorescence. Our results present satellite-based fluorescence as a valuable tool for evaluating nutrient stress predictions in ocean ecosystem models and give the first synoptic observational evidence that iron plays an important role in seasonal phytoplankton dynamics of the Indian Ocean. Satellite fluorescence may also provide a path for monitoring climate-phytoplankton physiology interactions and improving descriptions of phytoplankton light use efficiencies in ocean productivity models.This work was supported by grants from the NASA Ocean Biology and Biogeochemistry Program and the NSF Biological Oceanography Program

Satellite-Detected Fluorescence Reveals Global Physiology of Ocean Phytoplankton

Phytoplankton photosynthesis links global ocean biology and climate-driven fluctuations in the physical environment. These interactions are largely expressed through changes in phytoplankton physiology, but physiological status has proven extremely challenging to characterize globally. Phytoplankton fluorescence does provide a rich source of physiological information long exploited in laboratory and field studies, and is now observed from space. Here we evaluate the physiological underpinnings of global variations in satellite-based phytoplankton chlorophyll fluorescence. The three dominant factors influencing fluorescence distributions are chlorophyll concentration, pigment packaging effects on light absorption, and light-dependent energy-quenching processes. After accounting for these three factors, resultant global distributions of quenching-corrected fluorescence quantum yields reveal a striking consistency with anticipated patterns of iron availability. High fluorescence quantum yields are typically found in low iron waters, while low quantum yields dominate regions where other environmental factors are most limiting to phytoplankton growth. Specific properties of photosynthetic membranes are discussed that provide a mechanistic view linking iron stress to satellite-detected fluorescence. Our results present satellite-based fluorescence as a valuable tool for evaluating nutrient stress predictions in ocean ecosystem models and give the first synoptic observational evidence that iron plays an important role in seasonal phytoplankton dynamics of the Indian Ocean. Satellite fluorescence may also provide a path for monitoring climate-phytoplankton physiology interactions and improving descriptions of phytoplankton light use efficiencies in ocean productivity models

The effect of climate change on the carbon balance between photosynthesis and respiration in Antarctic microalgae

The biological process of the carbon cycle in the Antarctic Ocean is controlled by the photosynthetic activity of the primary producers. The amount of fixed carbon does not only depend on the photosynthetic activity but also on the carbon losses due to respiration. Thus, the ratio photosynthesis to respiration (rP/R) is an important parameter to predict the effect of climate change on the Antarctic ecosystem. Indeed, the ongoing changes in climate change are influencing the dynamics of environmental conditions, which has tremendous effects on the phytoplankton community. Therefore, two ecologically relevant species from the Southern Ocean were here investigated: the diatom Chaetoceros sp. and the prymnesiophyte Phaeocystis antarctica, studying the changes in the rP/R under global climate change conditions. Three main parameters were examined i.e temperature, salinity and iron limitation. The P/R ratio was significantly affected by temperature, while salinity had only a secondary importance, although with species-specific differences. More specifically, the values were ranging from 12.3 to 7.5 for Chaetoceros sp. and from 12.4 to 2.5 for P. antarctica. The changes in this ratio were principally due to variations in respiration, rather than in photosynthesis. Chaetoceros sp. appears to be less flexible in the regulation of the extent of photoprotective mechanisms (non-photochemical quenching and alternative electrons), but its photoprotective level was generally higher than in P. antarctica. Regarding iron limitation, data were successfully collected only for Chaetoceros sp.. The P/R ratio, equal to 2.8, did not change under iron limitation, with iron limited cells showing a very efficient acclimation to the lowered assimilatory metabolism by decreasing their respiratory losses

First assessment of the plant phenology index (PPI) for estimating gross primary productivity in African semi-arid ecosystems

The importance of semi-arid ecosystems in the global carbon cycle as sinks for CO2 emissions has recently been highlighted. Africa is a carbon sink and nearly half its area comprises arid and semi-arid ecosystems. However, there are uncertainties regarding CO2 fluxes for semi-arid ecosystems in Africa, particularly savannas and dry tropical woodlands. In order to improve on existing remote-sensing based methods for estimating carbon uptake across semi-arid Africa we applied and tested the recently developed plant phenology index (PPI). We developed a PPI-based model estimating gross primary productivity (GPP) that accounts for canopy water stress, and compared it against three other Earth observation-based GPP models: the temperature and greenness model, the greenness and radiation model and a light use efficiency model. The models were evaluated against in situ data from four semi-arid sites in Africa with varying tree canopy cover (3 to 65 percent). Evaluation results from the four GPP models showed reasonable agreement with in situ GPP measured from eddy covariance flux towers (EC GPP) based on coefficient of variation, root-mean-square error, and Bayesian information criterion. The PPI-based GPP model was able to capture the magnitude of EC GPP better than the other tested models. The results of this study show that a PPI-based GPP model is a promising tool for the estimation of GPP in the semi-arid ecosystems of Africa.Comment: Accepted manuscript; 12 pages, 4 tables, 9 figure

Developing screening tools for abiotic stresses using cowpea [Vigna unguiculata (L.) Walp.] as a model crop

Abiotic stresses cause extensive loss to agriculture production worldwide. Cowpea is an important legume crop grown widely in tropical and subtropical regions where high temperature, ultraviolet-B (UVB) radiation and drought are the common stress factors limiting production. Various vegetative, physiological, biochemical and reproductive plant attributes were assessed under a range of UVB radiation levels in Experiment I and in a combination with two doses of each carbon dioxide concentration [CO2], temperature, and UVB radiation and their interactions in Experiment II by using six cowpea genotypes and sunlit plant growth chambers. The dynamics of photosynthesis and fluorescence processes were assessed in 15 cowpea genotypes under drought condition in Experiment III in pot-grown plants under sunlit conditions. A distinct response pattern was not observed in cowpea in response to UVB radiation form 0 to 15 kJ; however, plants grown under elevated UVB showed reduced photosynthesis resulting in shorter plants and produced smaller flowers and lower seed yield. Increased phenolic compounds appeared to be a defense response to UVB radiation. The growth enhancements observed by doubling of [CO2] were not observed when plants were grown in combination with elevated UVB or temperature which also showed the most detrimental effects on plant growth and seed yield. Results form Experiment I and II revealed that cowpea reproductive traits were highly sensitive to abiotic stresses compared to the vegetative growth and development. A total stress response index (TSRI) technique, derived from all vegetative and reproductive parameters, was used to screen genotypes for their stress tolerance to UVB or combination of stresses. An increase in water use efficiency while maintaining higher rate of photosynthesis was an important drought tolerance mechanism in tolerant cowpea genotypes. Using principal component analysis technique, four groups of the genotypes were identified for their drought tolerance. Evaluating same genotypes across stress conditions revealed that no single genotype has the absolute tolerance characters to all stress conditions. The identified diversity for abiotic stress tolerance among cowpea genotypes and associated traits can be used to develop tolerant genotypes suitable for an agro-ecological niche though traditional breeding or genetic engineering methods

New Methods for Measurements of Photosynthesis from Space

Our ability to close the Earth's carbon budget and predict feedbacks in a warming climate depends critically on knowing where, when, and how carbon dioxide (CO2) is exchanged between the land and atmosphere. In particular, determining the rate of carbon fixation by the Earth's biosphere (commonly referred to as gross primary productivity, or GPP) and the dependence of this productivity on climate is a central goal. Historically, GPP has been inferred from spectral imagery of the land and ocean. Assessment of GPP from the color of the land and ocean requires, however, additional knowledge of the types of plants in the scene, their regulatory mechanisms, and climate variables such as soil moisture—just the independent variables of interest! Sunlight absorbed by chlorophyll in photosynthetic organisms is mostly used to drive photosynthesis, but some can also be dissipated as heat or re‐radiated at longer wavelengths (660–800 nm). This near‐infrared light re‐emitted from illuminated plants is termed solarinduced fluorescence (SIF), and it has been found to strongly correlate with GPP. To advance our understanding of SIF and its relation to GPP and environmental stress at the planetary scale, the Keck Institute for Space Studies (KISS) convened a workshop—held in Pasadena, California, in August 2012—to focus on a newly developed capacity to monitor chlorophyll fluorescence from terrestrial vegetation by satellite. This revolutionary approach for retrieving global observations of SIF promises to provide direct and spatially resolved information on GPP, an ideal bottom‐up complement to the atmospheric net CO2 exchange inversions. Workshop participants leveraged our efforts on previous studies and workshops related to the European Space Agency’s FLuorescence EXplorer (FLEX) mission concept, which had already targeted SIF for a possible satellite mission and had developed a vibrant research community with many important publications. These studies, mostly focused on landscape, canopy, and leaf‐level interpretation, provided the ground‐work for the workshop, which focused on the global carbon cycle and synergies with atmospheric net flux inversions. Workshop participants included key members of several communities: plant physiologists with experience using active fluorescence methods to quantify photosynthesis; ecologists and radiative transfer experts who are studying the challenge of scaling from the leaf to regional scales; atmospheric scientists with experience retrieving photometric information from space‐borne spectrometers; and carbon cycle experts who are integrating new observations into models that describe the exchange of carbon between the atmosphere, land and ocean. Together, the participants examined the link between “passive” fluorescence observed from orbiting spacecraft and the underlying photochemistry, plant physiology and biogeochemistry of the land and ocean. This report details the opportunity for forging a deep connection between scientists doing basic research in photosynthetic mechanisms and the more applied community doing research on the Earth System. Too often these connections have gotten lost in empiricism associated with the coarse scale of global models. Chlorophyll fluorescence has been a major tool for basic research in photosynthesis for nearly a century. SIF observations from space, although sensing a large footprint, probe molecular events occurring in the leaves below. This offers an opportunity for direct mechanistic insight that is unparalleled for studies of biology in the Earth System. A major focus of the workshop was to review the basic mechanisms that underlie this phenomenon, and to explore modeling tools that have been developed to link the biophysical and biochemical knowledge of photosynthesis with the observable—in this case, the radiance of SIF—seen by the satellite. Discussions led to the identification of areas where knowledge is still lacking. For example, the inability to do controlled illumination observations from space limits the ability to fully constrain the variables that link fluorescence and photosynthesis. Another focus of the workshop explored a “top‐down” view of the SIF signal from space. Early studies clearly identified a strong correlation between the strength of this signal and our best estimate of the rate of photosynthesis (GPP) over the globe. New studies show that this observation provides improvements over conventional reflectance‐based remote sensing in detecting seasonal and environmental (particularly drought related) modulation of photosynthesis. Apparently SIF responds much more quickly and with greater dynamic range than typical greenness indices when GPP is perturbed. However, discussions at the workshop also identified areas where top‐down analysis seemed to be “out in front” of mechanistic studies. For example, changes in SIF based on changes in canopy light interception and the light use efficiency of the canopy, both of which occur in response to drought, are assumed equivalent in the top‐down analysis, but the mechanistic justification for this is still lacking from the bottom‐up side. Workshop participants considered implications of these mechanistic and empirical insights for large‐scale models of the carbon cycle and biogeochemistry, and also made progress toward incorporating SIF as a simulated output in land surface models used in global and regional‐scale analysis of the carbon cycle. Comparison of remotely sensed SIF with modelsimulated SIF may open new possibilities for model evaluation and data assimilation, perhaps leading to better modeling tools for analysis of the other retrieval from GOSAT satellite, atmospheric CO2 concentration. Participants also identified another application for SIF: a linkage to the physical climate system arising from the ability to better identify regional development of plant water stress. Decreases in transpiration over large areas of a continent are implicated in the development and “locking‐in” of drought conditions. These discussions also identified areas where current land surface models need to be improved in order to enable this research. Specifically, the radiation transport treatments need dramatic overhauls to correctly simulate SIF. Finally, workshop participants explored approaches for retrieval of SIF from satellite and ground‐based sensors. The difficulty of resolving SIF from the overwhelming flux of reflected sunlight in the spectral region where fluorescence occurs was once a major impediment to making this measurement. Placement of very high spectral resolution spectrometers on GOSAT (and other greenhouse gas–sensing satellites) has enabled retrievals based on infilling of solar Fraunhofer lines, enabling accurate fluorescence measurements even in the presence of moderately thick clouds. Perhaps the most interesting challenge here is that there is no readily portable ground‐based instrumentation that even approaches the capability of GOSAT and other planned greenhouse gas satellites. This strongly limits scientists’ ability to conduct ground‐based studies to characterize the footprint of the GOSAT measurement and to conduct studies of radiation transport needed to interpret SIF measurement. The workshop results represent a snapshot of the state of knowledge in this area. New research activities have sprung from the deliberations during the workshop, with publications to follow. The introduction of this new measurement technology to a wide slice of the community of Earth System Scientists will help them understand how this new technology could help solve problems in their research, address concerns about the interpretation, identify future research needs, and elicit support of the wider community for research needed to support this observation. Somewhat analogous to the original discovery that vegetation indices could be derived from satellite measurements originally intended to detect clouds, the GOSAT observations are a rare case in which a (fortuitous) global satellite dataset becomes available before the research community had a consolidated understanding on how (beyond an empirical correlation) it could be applied to understanding the underlying processes. Vegetation indices have since changed the way we see the global biosphere, and the workshop participants envision that fluorescence can perform the next indispensable step by complementing these measurements with independent estimates that are more indicative of actual (as opposed to potential) photosynthesis. Apart from the potential FLEX mission, no dedicated satellite missions are currently planned. OCO‐2 and ‐3 will provide much more data than GOSAT, but will still not allow for regional studies due to the lack of mapping capabilities. Geostationary observations may even prove most useful, as they could track fluorescence over the course of the day and clearly identify stress‐related down‐regulation of photosynthesis. Retrieval of fluorescence on the global scale should be recognized as a valuable tool; it can bring the same quantum leap in our understanding of the global carbon cycle as vegetation indices once did

Application of FTIR spectroscopy for monitoring water quality in a hypertrophic aquatic ecosystem (Lake Auensee, Leipzig)

FTIR spectroscopy as molecular fingerprint has been used to assess macromolecular and ele-mental stoichiometry as well as growth rates of phytoplankton cells. Chemometric models have been developed to extract quantitative information from FTIR spectra to reveal macro-molecular composition (of proteins, carbohydrates and lipids), C:N ratio, and growth potential. In this study, we tested these chemometric models based on lab-cultured algal species in mon-itoring changes of phytoplankton community structure in a hypertrophic lake (Lake Auensee, Leipzig, Germany), where a seasonal succession of spring green algal bloom followed by cya-nobacterial dominance in summer can be commonly observed. Our results demonstrated that green algae reacted to environmental changes such as nitrogen limitation (due to imbalanced nitrogen and phosphorus supply) with restricted growth by changing carbon allocation from protein synthesis to storage carbohydrates and/or lipids, and increased C:N ratio. By contrast, cyanobacteria proliferated under nitrogen limiting conditions. Furthermore, the FTIR-based growth potential of green alga matched well with the population biomass determined by the Chl-a concentration. However, the predicted growth potential based on FTIR spectroscopy cannot describe the realistic growth development of cyanobacteria in this lake. These results revealed that green algae and cyanobacteria have different strategies of C-allocation stoichi-ometry and growth patterns in response to environmental changes. These taxon-specific re-sponses may explain at a molecular level why green algae bloomed in the spring under condi-tions with sufficient nutrient, lower pH and lower water temperature; while cyanobacteria overgrew green algae and dominated in the summer under conditions with limited nutrient availability, higher pH and higher water temperature. In addition, the applicability of these chemometric models for predicting field cyanobacterial growth is of limited value. This may be attributed to other special adaptation properties of cyanobacterial species under stress growth conditions. We used flow cytometry to isolate functional algal groups from the water samples. Despite some drawbacks of the flow cytometry combined FTIR spectroscopy tech-nique, this method provides prospects of monitoring water quality and early warning of harmful algal blooms

근접 표면 원격 센싱 시스템들을 이용한 지속적 식물 계절 및 태양 유도 엽록소 형광물질 관측

학위논문(박사) -- 서울대학교대학원 : 환경대학원 협동과정 조경학, 2022.2. 류영렬.Monitoring phenology, physiological and structural changes in vegetation is essential to understand feedbacks of vegetation between terrestrial ecosystems and the atmosphere by influencing the albedo, carbon flux, water flux and energy. To this end, normalized difference vegetation index (NDVI) and solar-induced chlorophyll fluorescence (SIF) from satellite remote sensing have been widely used. However, there are still limitations in satellite remote sensing as 1) satellite imagery could not capture fine-scale spatial resolution of SIF signals, 2) satellite products are strongly influenced by condition of the atmosphere (e.g. clouds), thus it is challenging to know physiological and structural changes in vegetation on cloudy days and 3) satellite imagery captured a mixed signal from over- and understory, thus it is difficult to study the difference between overstory and understory phenology separately. Therefore, in order to more accurately understand the signals observed from the satellite, further studies using near-surface remote sensing system to collect ground-based observed data are needed. The main purpose of this dissertation is continuous observation of vegetation phenology and SIF using near-surface remote sensing system. To achieve the main goal, I set three chapters as 1) developing low-cost filter-based near-surface remote sensing system to monitor SIF continuously, 2) monitoring SIF in a temperate evergreen needleleaf forest continuously, and 3) understanding the relationships between phenology from in-situ multi-layer canopies and satellite products. In Chapter 2, I developed the filter-based smart surface sensing system (4S-SIF) to overcome the technical challenges of monitoring SIF in the field as well as to decrease sensor cost for more comprehensive spatial sampling. I verified the satisfactory spectral performance of the bandpass filters and confirmed that digital numbers (DN) from 4S-SIF exhibited linear relationships with the DN from the hyperspectral spectroradiometer in each band (R2 > 0.99). In addition, we confirmed that 4S-SIF shows relatively low variation of dark current value at various temperatures. Furthermore, the SIF signal from 4S-SIF represents a strong linear relationship with QEpro-SIF either changing the physiological mechanisms of the plant using DCMU (3-(3, 4-dichlorophenyl)-1, 1-dimethyurea) treatment. I believe that 4S-SIF will be a useful tool for collecting in-situ data across multiple spatial and temporal scales. Satellite-based SIF provides us with new opportunities to understand the physiological and structural dynamics of vegetation from canopy to global scales. However, the relationships between SIF and gross primary productivity (GPP) are not fully understood, which is mainly due to the challenges of decoupling structural and physiological factors that control the relationships. In Chapter 3, I reported the results of continuous observations of canopy-level SIF, GPP, absorbed photosynthetically active radiation (APAR), and chlorophyll: carotenoid index (CCI) in a temperate evergreen needleleaf forest. To understand the mechanisms underlying the relationship between GPP and SIF, I investigated the relationships of light use efficiency (LUE_p), chlorophyll fluorescence yield (Φ_F), and the fraction of emitted SIF photons escaping from the canopy (f_esc) separately. I found a strongly non-linear relationship between GPP and SIF at diurnal and seasonal time scales (R2 = 0.91 with a hyperbolic regression function, daily). GPP saturated with APAR, while SIF did not. In addition, there were differential responses of LUE_p and Φ_F to air temperature. While LUE_p reached saturation at high air temperatures, Φ_F did not saturate. I also found that the canopy-level chlorophyll: carotenoid index was strongly correlated to canopy-level Φ_F (R2 = 0.84) implying that Φ_F could be more closely related to pigment pool changes rather than LUE_p. In addition, I found that the f_esc contributed to a stronger SIF-GPP relationship by partially capturing the response of LUE_p to diffuse light. These findings can help refine physiological and structural links between canopy-level SIF and GPP in evergreen needleleaf forests. We do not fully understand what satellite NDVI derived leaf-out and full leaf dates actually observe because deciduous broadleaf forest consists of multi-layer canopies typically and mixed-signal from multi-layer canopies could affect satellite observation. Ultimately, we have the following question: What phenology do we actually see from space compared to ground observations on multi-layer canopy phenology? In Chapter 4, I reported the results of 8 years of continuous observations of multi-layer phenology and climate variables in a deciduous broadleaf forest, South Korea. Multi-channel spectrometers installed above and below overstory canopy allowed us to monitor over- and understory canopy phenology separately, continuously. I evaluated the widely used phenology detection methods, curvature change rate and threshold with NDVI observed above top of the canopy and compared leaf-out and full leaf dates from both methods to in-situ observed multi-layer phenology. First, I found that NDVI from the above canopy had a strong linear relationship with satellites NDVI (R2=0.95 for MODIS products and R2= 0.85 for Landsat8). Second, leaf-out dates extracted by the curvature change rate method and 10% threshold were well matched with understory leaf-out dates. Third, the full-leaf dates extracted by the curvature change rate method and 90% threshold were similar to overstory full-leaf dates. Furthermore, I found that overstory leaf-out dates were closely correlated to accumulated growing degree days (AGDD) while understory leaf-out dates were related to AGDD and also sensitive to the number of chill days (NCD). These results suggest that satellite-based leaf-out and full leaf dates represent understory and overstory signals in the deciduous forest site, which requires caution when using satellite-based phenology data into future prediction as overstory and understory canopy show different sensitivities to AGDD and NCD.식물 계절 및 식생의 생리학적, 구조적인 변화를 지속적으로 모니터링 하는 것은 육상생태계와 대기권 사이의 에너지, 탄소 순환 등의 피드백을 이해하는데 필수적이다. 이를 관측하기 위하여 위성에서 관측된 정규화 식생 지수 (NDVI) 태양 유도 엽록소 형광물질 (SIF)는 대중적으로 사용되고 있다. 그러나, 우주 위성 기반의 자료는 다음과 같은 한계점들이 존재한다. 1) 아직까지 고해상도의 위성 기반 SIF 자료는 없고, 2) 위성 자료들은 대기의 질 (예, 구름)에 영향을 받아, 흐린 날의 식생의 생리학적, 구조적 변화를 탐지하기 힘들다. 또한, 3) 위성 이미지는 상부 식생과 하부 식생이 혼합되어 섞인 신호를 탐지하기 때문에, 각 층의 식물 계절을 각각 연구하기에 어려움이 있다. 그러므로, 위성에서 탐지한 신호를 평가하고, 식생의 생리학적, 구조적 변화를 보다 정확히 이해하기 위해서는 근접 표면 원격 센싱 시스템을 이용한 실측 자료 기반의 연구들이 요구된다. 본 학위논문의 주 목적은 근접 표면 원격 센싱 시스템을 이용하여 식물 계절 및 SIF를 현장에서 지속적으로 실측하고, 위성 영상 기반의 연구가 갖고 있는 한계점 및 궁금증들을 해결 및 보완하는 것이다. 이 목적을 달성하기 위하여, 아래와 같은 세가지 Chapter: 1) SIF를 관측하기 위한 필터 기반의 저렴한 근접 표면 센싱 시스템 개발, 2)온대 침엽수림에서의 연속적인 SIF 관측, 3)위성 기반의 식물 계절과 실측한 다층 식생의 식물 계절 비교로 구성하고, 이를 진행하였다. SIF는 식생의 구조적, 생리학적 변화를 이해하고, 추정하는 인자로 사용될 수 있어, SIF를 현장에서 관측하기 위한 다양한 근접 표면 원격 센싱 시스템들이 최근 제시되어 오고 있다. 그러나, 아직까지 SIF를 관측하기 위한 상업적으로 유통되는 관측 시스템은 현저히 부족하며, 분광계의 구조적 특성상 현장에서 관측 시스템을 보정 및 관리하기가 어려워 높은 질의 SIF를 취득하는 것은 매우 도전 적인 분야이다. 본 학위 논문의 Chapter 2에서는 SIF를 현장에서 보다 손쉽게 관측하기 위한 필터 기반의 근접 표면 센싱 시스템(Smart Surface Sensing System, 4S-SIF)을 개발하였다. 센서는 대역 필터들과 포토다이오드가 결합되어 있으며, 서보 모터를 사용하여 대역 필터 및 관측 방향을 자동적으로 변경함으로써, 한 개의 포토다이오드가 3개의 파장 범위(757, 760, 770 nm)의 빛 및 태양으로부터 입사되는 광량과 식생으로 반사/방출된 광량을 관측할 수 있도록 고안되었다. 포토다이오드로부터 인식된 디지털 수치 값은 상업적으로 판매되는 초고해상도 분광계(QE Pro, Ocean Insight)와 뚜렷한 선형 관계를 보이는 것을 확인하였다 (R2 > 0.99). 추가적으로, 4S-SIF에서 관측된 SIF와 초고해상도 분광계를 이용하여 추출한 SIF가 선형적인 관계를 이루는 것을 확인하였다. 식생의 생리학적 변화를 일으키는 화학 물질인 DCMU(3-(3, 4-dichlorophenyl)-1, 1-dimethyurea)을 처리했음에도 불구하고, 산출된 SIF들은 선형 관계를 보였다. 본 센서는 기존 시스템들에 비해 사용하기 쉽고 간단하며, 저렴하기 때문에 다양한 시공간적 스케일의 SIF를 관측할 수 있다는 장점이 있다. 위성 기반의 SIF를 이용하여 총일차생산성(gross primary productivity, GPP)을 추정하는 연구는 최근 탄소 순환 연구 분야에서 각광받고 있는 연구 주제이다. 그러나, SIF와 GPP의 관계는 여전히 많은 불확실성을 지니고 있는데, 이는 SIF-GPP 관계를 조절하는 식생의 구조적 및 생리학적 요인을 따로 분리하여 고찰한 연구들이 부족하기 때문이다. 본 학위 논문의 Chapter 3에서는 지속적으로 SIF, GPP, 흡수된 광합성유효복사량 (absorbed photosynthetically active radiation, APAR), 그리고 클로로필과 카로티노이드의 비율 인자 (chlorophyll: carotenoid index, CCI)를 온대침엽수림에서 연속적으로 관측하였다. SIF-GPP 관계의 구체적인 메커니즘 관계를 밝히기 위하여, 광 이용효율 (light use efficiency, LUE_p), 엽록소 형광 수득률 (chlorophyll fluorescence yield, Φ_F) 그리고 SIF 광자가 군락으로부터 방출되는 비율 (escape fraction, f_esc)을 각각 도출하고 탐구하였다. SIF와 GPP의 관계는 뚜렷한 비 선형적인 관계를 보이는 것을 확인했으며(R2 = 0.91 with a hyperbolic regression function, daily), 일주기 단위에서 SIF는 APAR에 대해 선형적이었지만 GPP는 APAR에 대해 뚜렷한 포화 관계를 보이는 것을 확인하였다. 추가적으로 LUE_p 와 Φ_F 가 대기 온도에 따라 반응하는 정도가 다른 것을 확인하였다. LUE_p는 높은 온도에서 포화 되었지만, Φ_F는 포화 패턴을 확인할 수 없었다. 또한, 군락 수준에서의 CCI와 Φ_F가 뚜렷한 상관 관계를 보였다(R2 = 0.84). 이는 Φ_F가 엽록소 색소에 영향을 LUE_p에 비해 더 강한 관계가 있을 수 있음을 시사한다. 마지막으로, f_esc가 태양광의 산란된 정도에 따라 반응을 하여, Φ_F와 LUE_p의 강한 상관 관계를 형성하는데 기여하는 것을 확인하였다. 이러한 발견은 온대 침엽수림에서 군락 수준의 SIF-GPP관계를 생리학적 및 구조적 측면에서 이해하고 규명하는데 큰 도움이 될 것이다. 식물 계절은 식생이 철을 따라 주기적으로 나타내는 변화를 관측하는 반응이다. 식물 계절은 육상생태계와 대기권 사이의 물질 순환을 이해하는데 매우 중요하다. 위성 기반의 NDVI는 식물 계절을 탐지하고 연구하는데 가장 대중적으로 사용된다. 그러나, 활엽수림에서의 위성 NDVI 기반의 개엽 시기 및 성숙 시기가 실제 어느 시점을 탐지하는지는 불분명하다. 실제 활엽수림은 다층 식생 구조의 삼차원으로 이루어져 있는 반면, 위성 영상은 다층 식생의 신호가 섞여 있는 이차원의 결과물이기 때문이다. 따라서, 위성 NDVI 기반의 식물 계절이 다층 식생 구조를 이루고 있는 활엽수림에서 실제 현장 관측과 비교하였을 때 어느 시점을 탐지하는지에 대한 궁금증이 남는다. 본 학위 논문의 Chapter 4에서는 지속적으로 8년 동안 활엽수림내의 다층 식생의 식물 계절을 근접 표면 원격 센싱 시스템을 이용하여 관측하고, 위성 NDVI 기반의 식물 계절과 비교하였다. 다채널 분광계를 상부 식생의 위와 아래에 설치함으로써, 상부 식생과 하부 식생의 식물 계절을 각각 연속적으로 관측하였다. 식물 계절을 탐지하기 위하여 가장 많이 사용되는 방법인 1) 역치를 이용하는 방법과 2) 이계도함수를 이용하는 방법을 사용하여 개엽 시기 및 성숙 시기를 계산하고 이를 다층 식생의 식물 계절과 비교하였다. 본 연구 결과, 첫번째로, 군락의 상층부에서 실측한 NDVI와 위성 기반의 NDVI가 강한 선형 관계를 보이는 것을 확인했다 (R2=0.95 는 MODIS 영상들 및 R2= 0.85 는 Landsat8). 두번째로, 이계도함수 방법과 10%의 역치 값을 이용한 방법이 비슷한 개엽 시기를 추정하는 것을 확인하였으며, 하부 식생의 개엽 시기와 비슷한 시기임을 확인하였다. 세번째로, 이계도함수 방법과 90%의 역치 값을 이용한 방법이 비슷한 성숙 시기를 산출하였으며, 이는 상부 식생의 성숙 시기와 비슷하였다. 추가적으로 상부 식생의 개엽 시기와 하부 식생의 개엽 시기가 온도와 반응하는 정도가 뚜렷하게 차이가 나는 것을 확인할 수 있었다. 상부 식생의 개엽 시기는 적산 생장 온도 일수 (AGDD)와 강한 상관성을 보였고, 하부 식생의 개엽 시기는 AGDD와 연관성을 갖고 있을 뿐만 아니라 추위 일수(NCD)에도 민감하게 반응하는 것을 확인하였다. 이러한 결과는 위성 NDVI 기반의 개엽 시기는 하부 식생의 개엽 시기와 연관성이 높고, 성숙 시기는 상부 식생의 성숙 시기와 비슷하다는 것을 의미한다. 또한, 상부 식생과 하부 식생이 온도에 다른 민감성을 갖고 있어, 위성에서 산출된 식물 계절을 이용하여 기후변화를 이해하고자 할 때, 어떤 층의 식생이 위성 영상에 주된 영향을 미치는지 고려해야 한다는 것을 시사한다. 위성은 넓은 지역의 변화를 손쉽게 모니터링할 수 있어 많은 가능성을 갖고 있는 도구이지만, 보다 정확한 위성 관측 값을 이해하기 위해서는 현장에서 관측된 자료를 기반으로 한 검증이 요구된다. 본 학위 논문에서는 1) 근접 표면 센싱 시스템을 개발, 2) 근접 표면 센싱 시스템을 활용한 식생의 생리학적 구조적 변화의 지속적인 관측, 3) 다층 식생 구조에서 관측되는 식물 계절 및 위성에서 추정된 식물 계절의 연관성 평가를 수행하였다. 개발한 근접 표면 센서는 상업 센서들과 비교했을 때, 가격적으로 저렴하고 손 쉽게 사용할 수 있었으며, 성능적으로도 부족함이 없었다. 근접 표면 센싱 시스템을 이용하여 SIF를 온대 침엽수림에서 지속적으로 관측한 결과, 총일차생산성과 SIF는 비선형 관계를 갖는 것을 확인하였다. 이는 많은 선행 연구들에서 발표한 위성 기반의 SIF와 GPP가 선형적인 관계를 보인다는 것과는 다소 상반된 결과이다. 다측 식생의 봄철 식물 계절을 연속적으로 관측하고, 위성 기반의 식물 계절과 비교평가한 연구에서는 위성 기반의 개엽 시기는 하부 식생에 영향을 주로 받고, 성숙 시기는 상부 식생의 시기와 비슷한 것을 확인하였다. 즉, 근접 표면 센싱 시스템을 이용하여 현장에서 실측한 결과는 위성 영상을 활용한 연구들과는 다른 결과를 보일 수도 있으며, 위성 영상을 평가 및 이해하는데 사용될 수 있다. 따라서, 보다 정확한 식생의 구조적, 생리학적 메커니즘을 이해하기 위해서는 근접 표면 센싱을 활용한 현장에서 구축한 자료 기반의 더 많은 연구들이 필요하다는 것을 시사한다.Abstract i Chapter 1. Introduction 2 1. Background 2 2. Purpose 5 Chapter 2. Monitoring SIF using a filter-based near surface remote sensing system 9 1. Introduction 9 2. Instrument desing and technical spefications of the filter-based smart surface sensing system (4S-SIF) 12 2.1. Ultra-narrow band pass filter 14 2.2. Calibration of 4S-SIF 15 2.3. Temperature and humidity response 16 2.4. Evaluate SIF quality from 4S-SIF in the field 17 3. Results 20 4. Discussion 23 Chapter 3. SIF is non-linearly related to canopy photosynthesis in a temperate evergreen needleleaf forest during fall transition 27 1. Introduction 27 2. Methods and Materials 31 2.1. Study site 31 2.2. Leaf-level fluorescence measurement 32 2.3. Canopy-level SIF and spectral reflectance measurement 34 2.4. SIF retrieval 37 2.5. Canopy-level photosynthesis estimates 38 2.6. Meteorological variables and APAR 39 2.7. Statistical analysis 40 3. Results 41 4. Discussion 48 4.1. Non-linear relationships between SIF and GPP 49 4.2. Role of f_esc in SIF-GPP relationship 53 4.3. Implications of non-linear SIF-GPP relationship in temperate ENF 54 5. Conclusion 57 6. Appendix 59 Chapter 4. Monitoring spring phenology of multi-layer canopy in a deciduous broadleaf forest: What signal do satellites actually see in space 65 1. Introduction 65 2. Materials and Methods 69 2.1. Study site 69 2.2. Multi-layer spectral reflectance and transmittance measurement 70 2.3. Phenometrics detection 72 2.4. In-situ multi-layer phenology 74 2.5. Satellite remote sensing data 75 2.6. Meteorological variables 75 3. Results 76 3.1. Seasonal to interannual variations of NDVI, 1-transmittance, and air temperature 76 3.2. Inter-annual variation of leaf-out and full-leaf dates 78 3.3. The relationships between dates calculated according tothreshold and in-situ multi-layer phenology 80 3.4. The relationship between multi-layer phenology, AGDD and NCD 81 4. Discussion 82 4.1. How do satellite-based leaf-out and full-leaf dates differ from in-situ multi-layer phenology 83 4.2. Are the 10 % and 90 % thresholds from satellite-basedNDVI always well matched with the leaf-out and full-leaf dates calculated by the curvature change rate 86 4.3. What are the implications of the difference between satellite-based and multi-layer phenology 87 4.4. Limitations and implications for future studies 89 5. Conclusion 91 6. Appendix 92 Chapter 5. Conclusion 114 Abstract in Korean 115박