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W Photoprotection in Tropical Marine Organisms

Increasing levels of ultraviolet (UV) radiation reaching the earth's surface which results from stratospheric ozone depletions could have serious implications for terrestrial plants and for aquatic organisms within the euphotic zone. A documented 9% decline in ozone at mid-latitudes is considered to produce a 12% increase in harmful UV radiation. The biologically damaging effects of higher UV levels, particularly W-B (280-320 rim), could manifest earlier in the tropics because of the relative thinness of the earth's equatorial ozone layer. Tropical marine organisms are also living close to their upper tolerance levels of water temperature, However, despite the large potential effects on plants and animals, little is known about UV effects on tropical ecosystems. Long-term ecological studies are needed to quantify the effects of increased UV radiation on terrestrial and marine ecosystems and to produce reliable data for prediction. Plants have developed several mechanisms to protect themselves from harmful UV radiation, one of which is the production of secondary leaf pigments that absorb W-B radiation (screening pigments). A higher concentration of screening pigments (e.g. flavonoids) in leaves may be interpreted as a natural response to increased W radiation. If higher concentrations of flavonoids filter out the excessive W radiation, no damage will occur, as suggested by Caldwell et al. (1989) and Tevini (1993). Failure to screen all W-B may result in deleterious effects on photosynthesis, plant genetic material, and plant and leaf morphology and growth. Eventually this will have an impact on ecosystem processes, structure, species composition, and productivity. This paper describes an ongoing project that is assessing the responses of mangroves, seagrasses and corals to W radiation by studying pigment concentrations, biophysical parameters, and variations in spectral reflectance in the field and in W-reduction experiments. Preliminary results on the distribution of W-absorbing flavonoid compounds in red mangroves (Rhizophora mangle) and the seagrass Thalassia testudinum, are presented. This research also provides, for the first time, a permanent record of daily W irradiance measurements at a tropical -location.

Armstrong, Roy A.↗

Lipophilic pigments from cyanobacterial (blue-green algal) and diatom mats in Hamelin Pool, Shark Bay, Western Australia

Lipophilic pigments were examined in microbial mat communities dominated by cyanobacteria in the intertidal zone and by diatoms in the subtidal and sublittoral zones of Hamelin Pool, Shark Bay, Western Australia. These microbial mats have evolutionary significance because of their similarity to lithfied stromatolites from the Proterozoic and Early Paleozoic eras. Fucoxanthin, diatoxanthin, diadinoxanthin, beta-carotene, and chlorophylls a and c characterized the diatom mats, whereas cyanobacterial mats contained myxoxanthophyll, zeaxanthin, echinenone, beta-carotene, chlorophyll a and, in some cases, sheath pigment. The presence of bacteriochlorophyll a within the mats suggest a close association of photosynthetic bacteria with diatoms and cyanobacteria. The high carotenoids : chlorophyll a ratios (0.84-2.44 wt/wt) in the diatom mats suggest that carotenoids served a photoprotective function in this high light environment. By contrast, cyanobacterial sheath pigment may have largely supplanted the photoprotective role of carotenoids in the intertidal mats.

NASA Discipline Exobiology↗

Sources and geochemical evolution of cyanide and formaldehyde

The major source of cyanide has, in current paleoatmospheric models, been assumed to be the reaction of photodissociated thermospheric nitrogen with a limiting supply of stratospheric methane. Formaldehyde may be produced with more ease from an atmosphere of carbon dioxide as the dominant carbon species, and from carbonate in solution or sorbed in double layer hydroxide minerals. Potentially more important sources for cyanide and other carbon containing molecules are the partially photoprotected northern and southern auroral ovals where continuous currents reaching several mega-amperes induce ion-molecule reactions, extending into the lower stratosphere. In simulated environments of this kind, the cyanide ion is known to be produced from oxidized carbon species potentially more abundant than methane. Rainout of cyanide and formaldehyde place them in two different geochemical reaction reservoirs. In the anoxic Archean hydrosphere, about 1mM in Fe2(+), the cyanide ion would have been efficiently converted to the stable ferrocyanide complex Fe(CN) sub 6(4-), protecting it from the commonly considered fate of decomposition by hydrolysis, and eventually incorporating it in pyroaurite type minerals, most efficiently in green rust where it converts to insoluble ferriferrocyanide, prussian blue.

Arrhenius, G.↗

Remote sensing of the biological dynamics of large-scale salt evaporation ponds

Optical properties of salt evaporation ponds associated with Exportadora de Sal, a salt production company in Baja California Sur, Mexico, were analyzed using a combination of spectroradiometer and extracted pigment data, and Landsat-5 Thematic Mapper imagery. The optical characteristics of each pond are determined by the biota, which consists of dense populations of algae and photosynthetic bacteria containing a wide variety of photosynthetic and photoprotective pigments. Analysis has shown that spectral and image data can differentiate between taxonomic groups of the microbiota, detect changes in population distributions, and reveal large-scale seasonal dynamics.

Richardson, Laurie L.↗

SeaWiFS Technical Report Series. Volume 29: SeaWiFS CZCS-type pigment algorithm

The Sea-viewing Wide Field-of-view Sensor (SeaWiFS) mission will provide operational ocean color that will be superior to the previous Coastal Zone Color Sensor (CZCS) proof-of-concept mission. an algorithm is needed that exploits the full functionality of SeaWiFS whilst remaining compatible in concept with algorithms used for the CZCS. This document describes the theoretical rationale of radiance band-radio methods for determining chlorophyll alpha and other important biogeochemical parameters, and their implementation for the SeaWiFS mission. Pigment interrelationships are examined to explain the success of the CZCS algorithms. In the context where chlorophyll alpha absorbs only weakly at 520 nm, the success of the 520 nm to 550 nm CZCS band ratio needs to be explained. This is explained by showing that in pigment data from a range of oceanic provinces chlorophyll alpha (absorbing at less than 490 nm), carotenoids (absorbing at greater than 460 nm), and total pigment are highly correlated. Correlations within pigment groups particularly photoprotectant and photosynthetic carotenoids are less robust. The sources of variability in optical data re examined using the NIMBUS Experiment Team (NET) bio-optical data set and bio-optical model. In both the model and NET data, the majority of the variance in the optical data is attributed to variability in pigment (chlorophyll alpha, and total particulates, with less than 5% of the variability resulting from pigment assemblage. The relationships between band ratios and chlorophyll is examined analytically, and a new formulation based on a dual hyperbolic model is suggested which gives a better calibration curve than the conventional log-log linear regression fit. The new calibration curve shows that 490:555 ratio is the best single-band ratio and is the recommended CZCS-type pigment algorithm. Using both the model and NET data, a number of multiband algorithms are developed; the best of which is an algorithm based on the 443:555 and 490:555 ratios. From model data, the form of potential algorithms for other products, such as total particulates and dissolved organic matter (DOM), are suggested.

Hooker, Stanford B.↗

Evaluation of the photochemical reflectance index in AVIRIS imagery

In this paper, we evaluate the potential for extracting the 'photochemical reflectance index' (PRI; previously called the 'physiological reflectance index') from AVIRIS data. This index, which is derived from narrow-band reflectance at 531 and 570 nm, has proven to be a useful indicator of photosynthetic function at the leaf and canopy scales. At the leaf level, PRI varies with photosynthetic capacity, radiation-use efficiency, and vegetation type (unpublished data). This finding is consistent with the hypothesis that vegetation types exhibiting chronically reduced photosynthesis during periods of stress (e.g. drought-tolerant evergreens) invest proportionally more in photoprotective processes than vegetation with high photosynthetic capacity (e.g. crops or deciduous perennials). Vertical transects in tropical and boreal forest canopies have indicated declines in PRI associated with downregulation of photosynthesis at the canopy tops under sunny, dry midday conditions (unpublished data). This reduced PRI in upper canopy levels provides a further basis for examining this signal with the 'view from above' afforded by aircraft overflights. Although many factors could confound interpretation of a subtle physiological signal at the landscape scale, we conducted a preliminary examination of PRI extracted from existing, AVIRIS imagery of Stanford University's Jasper Ridge Biological Preserve obtained on the June 2nd, 1992, overflight. The goal was to use the hyperspectral capabilities of AVIRIS to evaluate the potential of this index for obtaining useful physiological data at the landscape scale. The expectation based on leaf- and canopy-level studies was that regions containing vegetation of reduced photosynthetic capacity (e.g. chaparral or evergreen woodland) would exhibit lower PRI values than regions of high capacity (e.g. deciduous woodland).

Gamon, John A.↗

SeaWiFS Technical Report Series: The SeaWiFS CZCS-Type Pigment Algorithm - Volume 29

The Sea-viewing Wide Field-of-view Sensor (SeaWiFS) mission will provide operational ocean color that will be superior to the previous Coastal Zone Color Sensor (CZCS) proof-of-concept mission. An algorithm is needed that exploits the full functionality of SeaWiFS whilst remaining compatible in concept with algorithms used for the CZCS. This document describes the theoretical rationale of radiance band-ratio methods for determining chlorophyll-a and other important biogeochemical parameters, and their implementation for the SeaWIFS mission. Pigment interrelationships are examined to explain the success of the CZCS algorithms. In the context where chlorophyll-a absorbs only weakly at 520 nm, the success of the 520 nm to 550 nm CZCS band ratio needs to be explained. This is explained by showing that in pigment data from a range of oceanic provinces chlorophyll-a (absorbing at less than 490 nm), carotenoids (absorbing at greater than 460 nm), and total pigment are highly correlated. Correlations within pigment groups particularly photoprotectant and photosynthetic carotenoids are less robust. The sources of variability in optical data are examined using the NIMBUS Experiment Team (NET) bio-optical data set and bio-optical model. In both the model and NET data, the majority of the variance in the optical data is attributed to variability in pigment (chlorophyll-a), and total particulates, with less than 5% of the variability resulting from pigment assemblage. The relationships between band ratios and chlorophyll is examined analytically, and a new formulation based on a dual hyperbolic model is suggested which gives a better calibration curve than the conventional log-log linear regression fit. The new calibration curve shows the 490:555 ratio is the best single-band ratio and is the recommended CZCS-type pigment algorithm. Using both the model and NET data, a number of multiband algorithms are developed; the best of which is an algorithm based on the 443:555 and 490:555 ratios. From model data, the form of potential algorithms for other products, such as total particulates and dissolved organic matter (DOM), are suggested.

Hooker, Stanford B.↗

UV-Visible Spectroscopic Method and Models for Assessment and Monitoring of Harmful Algal Blooms

The development of an enhanced predictive and early warning capability for the occurrence and impact of harmful algal blooms (HABs) would be of great benefit to coastal communities. A critical issue for early detection and monitoring of HABs is the need to detect harmful algal species within a mixed-species phytoplankton assemblage. Possession of UV-absorbing compounds called mycosporine-like amino acids (MAAs) may be one factor that allows HAB species to out-compete their phytoplankton neighbors. Possession of MAAs, which we believe can be inferred from strong UV-absorption signals in phytoplankton absorption coefficients, can be used as a flag for potential HAB outbreak. The goal of this project was to develop a solar simulating UV-visible incubator to grow HAB dinoflagellates, to begin MAA analysis of samples collected on global cruises, and to carry out initial experiments on HAB dinoflagellate species in pure culture. Our scientific objectives are to quantify MAA production and spectral induction mechanisms in HAB species, to characterize spectral absorption of MAAs, and to define the ecological benefit of MAAs (i.e. photoprotection). Data collected on cruises to the global oceans will be used to parameterize phytoplankton absorption in the UV region, and this parameterization could be incorporated into existing models of seawater optical properties in the UV spectral region. Data collected in this project were used for graduate fellowship applications by Elizabeth Frame. She has been awarded an EPA STAR fellowship to continue the work initiated by this project.

Mitchell, B. Greg↗

Problems in Assessment of the UV Penetration into Natural Waters from Space-based Measurements

Satellite instruments currently provide global maps of surface UV (ultraviolet) irradiance by combining backscattered radiance data with radiative transfer models. The models are often limited by uncertainties in physical input parameters of the atmosphere and surface. Global mapping of the underwater UV irradiance creates further challenges for the models. The uncertainties in physical input parameters become more serious because of the presence of absorbing and scattering quantities caused by biological processes within the oceans. In this paper we summarize the problems encountered in the assessment of the underwater UV irradiance from space-based measurements, and propose approaches to resolve the problems. We have developed a radiative transfer scheme for computation of the UV irradiance in the atmosphere-ocean system. The scheme makes use of input parameters derived from satellite instruments such as TOMS (Total Ozone Mapping Spectrometer) and SeaWiFS (Sea-viewing Wide Field-of-view Sensor). The major problem in assessment of the surface UV irradiance is to accurately quantify the effects of clouds. Unlike the standard TOMS UV algorithm, we use the cloud fraction products available from SeaWiFS and MODIS (Moderate Resolution Imaging Spectrometer) to calculate instantaneous surface flux at the ocean surface. Daily UV doses can be calculated by assuming a model of constant cloudiness throughout the day. Both SeaWiFS and MODIS provide some estimates of seawater optical properties in the visible. To calculate the underwater UV flux the seawater optical properties must be extrapolated down to shorter wavelengths. Currently, the problem of accurate extrapolation of visible data down to the UV spectral range is not solved completely, and there are few available measurements. The major difficulty is insufficient correlation between photosynthetic and photoprotective pigments of phytoplankton absorbing in the visible and UV respectively. We propose to empirically parameterize seawater absorption in the UV on a basis of available data sets of bio-optical measurements from a variety of ocean waters. Another problem is the lack of reliable data on pure seawater absorption in the UV. Laboratory measurements of the UV absorption of both pure water and pure seawater are required.

Vasilkov, Alexander P.↗

High Spectral Resolution MODIS Algorithms for Ocean Chlorophyll in Case II Waters

The Case 2 chlorophyll a algorithm is based on a semi-analytical, bio-optical model of remote sensing reflectance, R(sub rs)(lambda), where R(sub rs)(lambda) is defined as the water-leaving radiance, L(sub w)(lambda), divided by the downwelling irradiance just above the sea surface, E(sub d)(lambda,0(+)). The R(sub rs)(lambda) model (Section 3) has two free variables, the absorption coefficient due to phytoplankton at 675 nm, a(sub phi)(675), and the absorption coefficient due to colored dissolved organic matter (CDOM) or gelbstoff at 400 nm, a(sub g)(400). The R(rs) model has several parameters that are fixed or can be specified based on the region and season of the MODIS scene. These control the spectral shapes of the optical constituents of the model. R(sub rs)(lambda(sub i)) values from the MODIS data processing system are placed into the model, the model is inverted, and a(sub phi)(675), a(sub g)(400) (MOD24), and chlorophyll a (MOD21, Chlor_a_3) are computed. Algorithm development is initially focused on tropical, subtropical, and summer temperate environments, and the model is parameterized in Section 4 for three different bio-optical domains: (1) high ratios of photoprotective pigments to chlorophyll and low self-shading, which for brevity, we designate as 'unpackaged'; (2) low ratios and high self-shading, which we designate as 'packaged'; and (3) a transitional or global-average type. These domains can be identified from space by comparing sea-surface temperature to nitrogen-depletion temperatures for each domain (Section 5). Algorithm errors of more than 45% are reduced to errors of less than 30% with this approach, with the greatest effect occurring at the eastern and polar boundaries of the basins. Section 6 provides an expansion of bio-optical domains into high-latitude waters. The 'fully packaged' pigment domain is introduced in this section along with a revised strategy for implementing these variable packaging domains. Chlor_a_3 values derived semi-analytically and Chlor_a_2 values derived empirically using the O Reilly et al. OC3M algorithm from MODIS Terra radiances are compared to field chlorophyll-a concentrations in Sections 7 and 8.

Carder, Kendall L.↗

A genomic timescale of prokaryote evolution: insights into the origin of methanogenesis, phototrophy, and the colonization of land

BACKGROUND: The timescale of prokaryote evolution has been difficult to reconstruct because of a limited fossil record and complexities associated with molecular clocks and deep divergences. However, the relatively large number of genome sequences currently available has provided a better opportunity to control for potential biases such as horizontal gene transfer and rate differences among lineages. We assembled a data set of sequences from 32 proteins (approximately 7600 amino acids) common to 72 species and estimated phylogenetic relationships and divergence times with a local clock method. RESULTS: Our phylogenetic results support most of the currently recognized higher-level groupings of prokaryotes. Of particular interest is a well-supported group of three major lineages of eubacteria (Actinobacteria, Deinococcus, and Cyanobacteria) that we call Terrabacteria and associate with an early colonization of land. Divergence time estimates for the major groups of eubacteria are between 2.5-3.2 billion years ago (Ga) while those for archaebacteria are mostly between 3.1-4.1 Ga. The time estimates suggest a Hadean origin of life (prior to 4.1 Ga), an early origin of methanogenesis (3.8-4.1 Ga), an origin of anaerobic methanotrophy after 3.1 Ga, an origin of phototrophy prior to 3.2 Ga, an early colonization of land 2.8-3.1 Ga, and an origin of aerobic methanotrophy 2.5-2.8 Ga. CONCLUSIONS: Our early time estimates for methanogenesis support the consideration of methane, in addition to carbon dioxide, as a greenhouse gas responsible for the early warming of the Earths' surface. Our divergence times for the origin of anaerobic methanotrophy are compatible with highly depleted carbon isotopic values found in rocks dated 2.8-2.6 Ga. An early origin of phototrophy is consistent with the earliest bacterial mats and structures identified as stromatolites, but a 2.6 Ga origin of cyanobacteria suggests that those Archean structures, if biologically produced, were made by anoxygenic photosynthesizers. The resistance to desiccation of Terrabacteria and their elaboration of photoprotective compounds suggests that the common ancestor of this group inhabited land. If true, then oxygenic photosynthesis may owe its origin to terrestrial adaptations.

Methane/metabolism↗

Seasonal And Regional Differentiation Of Bio-Optical Properties Within The North Polar Atlantic

Using data collected during spring and summer seasons in the north polar Atlantic we examined the variability of the spectral absorption, a(lambda), and backscattering, b(sub b)(lambda), coefficients of surface waters and its relation to phytoplankton pigment concentration and composition. For a given chlorophyll a concentration (TChla), the concentrations of photosynthetic carotenoids (PSC), photoprotective carotenoids (PPC), and total accessory pigments (AP) were consistently lower in spring than in summer. The chlorophyll-specific absorption coefficients of phytoplankton and total particulate matter were also lower in spring, which can be partly attributed to lower proportions of PPC, PSC, and AP in spring. The spring values of the green-to-blue band ratio of the absorption coefficient were higher than the summer ratios. The blue-to-green ratios of backscattering coefficient were also higher in spring. The higher b(sub b) values and lower blue-to-green b(sub b) ratios in summer were likely associated with higher concentrations of detrital particles in summer compared to spring. Because the product of the green-to-blue absorption ratio and the blue-to-green backscattering ratio is a proxy for the blue-to-green ratio of remote-sensing reflectance, we conclude that the performance of ocean color band-ratio algorithms for estimating pigments in the north polar Atlantic is significantly affected by seasonal shifts in the relationships between absorption and TChla as well as between backscattering and TChla. Intriguingly, however, fairly good estimate of the particulate beam attenuation coefficient at 660 nm (potential measure of total particulate matter or particulate organic carbon concentration) can be obtained by applying a single blue-to-green band-ratio algorithm for both spring and summer seasons.

Stramska, Malgorzata↗

Diurnal and Directional Responses of Chlorophyll Fluorescence and the PRI in a Cornfield

Determining the health and vigor of vegetation using high spectral resolution remote sensing is an important goal which has application to monitoring agriculture and ecosystem productivity and carbon exchange. Two spectral indices used to assess whether vegetation is performing near-optimally or exhibiting symptoms of environmental stress (e.g., drought or nutrient deficiency, non-optimal temperatures, etc.) are the Photochemical Reflectance Index (PRI) and solar-induced red and far-red Chlorophyll Fluorescence (Fs). Both the PRI and Fs capture the dynamics of photoprotection mechanisms within green foliage: the PRI is based on the association of the reflected radiation in the green spectrum with the xanthophyll cycle, whereas Fs measures the emitted radiation in the red and far-red spectrum. Fs was determined from retrievals in the atmospheric oxygen absorption features centered at 688 and 760 nm using a modified Fraunhofer Line Depth (FLD) method. We previously demonstrated diurnal and seasonal PRI differences for sunlit vs. shaded foliage in a conifer forest canopy, as expressed in the hotspot and darkspot of the Bidirectional Reflectance Function (BRF). In a USDA-ARS experimental field site located in Beltsville, MD, USA, measurements were acquired over a corn crop from a nadir view in 2008 with an ASD FieldSpec Pro (Analytical Spectral Devices, Inc., Boulder, CO, USA) to study the behavior of the PRI for sunlit and shaded foliage as captured in reflectance variations associated with the BRF, in a I m tall canopy in the vegetative growth stage. Those observations were compared to simulations obtained from two radiative transfer models. Measurements were then acquired to examine whether the PRI and Fs were influenced by view zenith and azimuth geometries at different times of day. Those measurements were made in 2010 with the Ocean Optics USB4000 Miniature Fiber Optic Spectrometer (Ocean Optics Inc., Dunedin, Florida, USA) at several times during the day on multiple days throughout the growing season. We found that the PRI consistently had higher values, indicating lower stress, in the BRF darkspot associated with shaded foliage than in the hotspot associated with sunlit foliage. We also found that Fs exhibited differences associated with sunlit and shaded canopy sectors, which were most pronounced for the red/far-red Fs ratio. Values indicated greater physiological stress in afternoons compared to mornings, and in the early senescent canopy as compared to the vegetative growth stage, BRFs for both the PRI and the red/far-red Fs ratio were bowl-shaped for the full azimuth sweep of the canopy. These two spectral indices (PRI, Fs ratio) provided complementary information on the photosynthetic function of the corn canopy.

Middleton, Elizabeth↗