Search NASA⌕ Search

SEARCH · Search NASA

Results for “Chlorophyll A”

Search indexed NASA NTRS and DOE OSTI research on propulsion, heat transfer, battery materials and energy systems. Follow report and document links to the original sources.

Quote a phrase for an exact phrase match. Source license links do not imply unrestricted reuse.

At least 37 records · Page 2

Optical properties of intact leaves for estimating chlorophyll concentration

Changes in leaf chlorophyll content can serve as relative indicators of plant vigor and environmental quality. This study identified reflectance, transmittance, and absorptance wavebands and band ratios within the 400- to 850-nm range for intact leaves that could be used to estimate extracted leaf chlorophyll per unit leaf area (areal concentration) with minimal error. Leaf optical properties along with chlorophyll a, b, and a + b concentrations were measured for the planar-leaved sweetgum (Liquidambar styraciflua L.), red maple (Acer rubrum L.), wild grape (Vitis rotundifolia Michx.), and switchcane [Arundinaria gigantea (Walter) Muhl.], and for needles of longleaf pine (Pinus palustris Miller). Generally, reflectance, transmittance, and absorptance corresponded most precisely with chlorophyll concentrations at wavelengths near 700 nm, although regressions were also strong in the 550- to 625-nm range. A power function was superior to a simple linear function in yielding low standard deviations of the estimate (s). When data were combined among the planar-leaved species, s values were low at approximately 50 mumol/m2 out of a 940 mumol/m2 range in chlorophyll a + b at best-fit wavelengths of 707 to 709 nm. Minimal s values for chlorophyll a + b ranged from 32 to 62 mumol/m2 across species when band ratios having numerator wavelengths of 693 to 720 nm were used with the application of a power function. Optimal denominator wavelengths for the band ratios were 850 nm for reflectance and transmittance and 400 nm for absorptance. This information can be applied in designing field portable chlorophyll meters and in the landscape-scale remote sensing of plant responses to the environment.

Chlorophyll/analysis↗

Completing the Feedback Loop: The Impact of Chlorophyll Data Assimilation on the Ocean State

In anticipation of the integration of a full biochemical model into the next generation GMAO coupled system, an intermediate solution has been implemented to estimate the penetration depth (1Kd_PAR) of ocean radiation based on the chlorophyll concentration. The chlorophyll is modeled as a tracer with sources-sinks coming from the assimilation of MODIS chlorophyll data. Two experiments were conducted with the coupled ocean-atmosphere model. In the first, climatological values of Kpar were used. In the second, retrieved daily chlorophyll concentrations were assimilated and Kd_PAR was derived according to Morel et al (2007). No other data was assimilated to isolate the effects of the time-evolving chlorophyll field. The daily MODIS Kd_PAR product was used to validate the skill of the penetration depth estimation and the MERRA-OCEAN re-analysis was used as a benchmark to study the sensitivity of the upper ocean heat content and vertical temperature distribution to the chlorophyll input. In the experiment with daily chlorophyll data assimilation, the penetration depth was estimated more accurately, especially in the tropics. As a result, the temperature bias of the model was reduced. A notably robust albeit small (2-5 percent) improvement was found across the equatorial Pacific ocean, which is a critical region for seasonal to inter-annual prediction.

MERRA-OCEAN↗

Forecasting Ocean Chlorophyll in the Equatorial Pacific

Using a global ocean biogeochemical model combined with a forecast of physical oceanic and atmospheric variables from the NASA Global Modeling and Assimilation Office, we assess the skill of a chlorophyll concentrations forecast in the Equatorial Pacific for the period 2012-2015 with a focus on the forecast of the onset of the 2015 El Niño event. Using a series of retrospective 9-month hindcasts, we assess the uncertainties of the forecasted chlorophyll by comparing the monthly total chlorophyll concentration from the forecast with the corresponding monthly ocean chlorophyll data from the Suomi-National Polar-orbiting Partnership Visible Infrared Imaging Radiometer Suite (S-NPP VIIRS) satellite. The forecast was able to reproduce the phasing of the variability in chlorophyll concentration in the Equatorial Pacific, including the beginning of the 2015-2016 El Niño. The anomaly correlation coefficient (ACC) was significant (p less than 0.05) for forecast at 1-month (R=0.33), 8-month (R=0.42) and 9-month (R=0.41) lead times. The root mean square error (RMSE) increased from 0.0399 microgram ch1 L(exp -1) for the 1-month lead forecast to a maximum of 0.0472 microgram ch1 L(exp -1) for the 9-month lead forecast indicating that the forecast of the amplitude of chlorophyll concentration variability was getting worse. Forecasts with a 3-month lead time were on average the closest to the S-NPP VIIRS data (23% or 0.033 microgram ch1 L(exp -1)) while the forecast with a 9-month lead time were the furthest (31% or 0.042 microgram ch1 L(exp -1)). These results indicate the potential for forecasting chlorophyll concentration in this region but also highlights various deficiencies and suggestions for improvements to the current biogeochemical forecasting system. This system provides an initial basis for future applications including the effects of El Niño events on fisheries and other ocean resources given improvements identified in the analysis of these results.

Anomaly Correlation Coefficient (ACC)↗

Assessing the Skill of Chlorophyll Forecasts: Latest Development and Challenges Ahead Using the Case of the Equatorial Pacific

Using a global ocean biogeochemical model combined with a forecast of physical oceanic and atmospheric variables from the NASA Global Modeling and Assimilation Office, we assess the skill of a chlorophyll concentrations forecast in the Equatorial Pacific for the period 2012-2015 with a focus on the forecast of the onset of the 2015 El Nino event. Using a series of retrospective 9-month hindcasts, we assess the uncertainties of the forecasted chlorophyll by comparing the monthly total chlorophyll concentration from the forecast with the corresponding monthly ocean chlorophyll data from the Suomi-National Polar-orbiting Partnership Visible Infrared Imaging Radiometer Suite (S-NPP VIIRS) satellite. The forecast was able to reproduce the phasing of the variability in chlorophyll concentration in the Equatorial Pacific, including the beginning of the 2015-2016 El Nino. The anomaly correlation coefficient (ACC) was significant (p less than 0.05) for forecast at 1-month (R=0.33), 8-month (R=0.42) and 9-month (R=0.41) lead times. The root mean square error (RMSE) increased from 0.0399 microgram chl L(exp -1) for the 1-month lead forecast to a maximum of 0.0472 microgram chl L(exp -1) for the 9-month lead forecast indicating that the forecast of the amplitude of chlorophyll concentration variability was getting worse. Forecasts with a 3-month lead time were on average the closest to the S-NPP VIIRS data (23% or 0.033 microgram chl L(exp -1)) while the forecast with a 9-month lead time were the furthest (31% or 0.042 microgram chl L(exp -1)). These results indicate the potential for forecasting chlorophyll concentration in this region but also highlights various deficiencies and suggestions for improvements to the current biogeochemical forecasting system. This system provides an initial basis for future applications including the effects of El Nino events on fisheries and other ocean resources given improvements identified in the analysis of these results.

El Nin↗

The Effects of Chlorophyll Assimilation on Carbon Fluxes in a Global Biogeochemical Model

In this paper, we investigated whether the assimilation of remotely-sensed chlorophyll data can improve the estimates of air-sea carbon dioxide fluxes (FCO2). Using a global, established biogeochemical model (NASA Ocean Biogeochemical Model, NOBM) for the period 2003-2010, we found that the global FCO2 values produced in the free-run and after assimilation were within -0.6 mol C m(sup -2) y(sup -1) of the observations. The effect of satellite chlorophyll assimilation was assessed in 12 major oceanographic regions. The region with the highest bias was the North Atlantic. Here the model underestimated the fluxes by 1.4 mol C m(sup -2) y(sup -1) whereas all the other regions were within 1 mol C m(sup -2) y(sup -1) of the data. The FCO2 values were not strongly impacted by the assimilation, and the uncertainty in FCO2 was not decreased, despite the decrease in the uncertainty in chlorophyll concentration. Chlorophyll concentrations were within approximately 25% of the database in 7 out of the 12 regions, and the assimilation improved the chlorophyll concentration in the regions with the highest bias by 10-20%. These results suggest that the assimilation of chlorophyll data does not considerably improve FCO2 estimates and that other components of the carbon cycle play a role that could further improve our FCO2 estimates.

Biogeochemical Model↗

Charge transfer as a mechanism for chlorophyll fluorescence concentration quenching

Highly concentrated solutions of chlorophyll display rapid fluorescence quenching. The same devastating energy loss is not seen in photosynthetic light-harvesting antenna complexes, despite the need for chromophores to be in close proximity to facilitate energy transfer. A promising, though unconfirmed mechanism for the observed quenching is energy transfer from an excited chlorophyll monomer to a closely associated chlorophyll pair that subsequently undergoes rapid nonradiative decay to the ground state via a short-lived intermediate charge-transfer state. In this work, we make use of newly emerging fast methods in quantum chemistry to assess the feasibility of this proposed mechanism. We calculate rate constants for the initial charge separation, based on Marcus free-energy surfaces extracted from molecular dynamics simulations of solvated chlorophyll pairs, demonstrating that this pathway will compete with fluorescence (i.e., drive quenching) at experimentally measured quenching concentrations. We show that the rate of charge separation is highly sensitive to interchlorophyll distance and the relative orientations of chromophores within a quenching pair. We discuss possible solvent effects on the rate of charge separation (and consequently the degree of quenching), using the light-harvesting complex II (LH2) protein from rps. acidophila as a specific example of how this process might be controlled in a protein environment. Crucially, we reveal that the LH2 antenna protein prevents quenching, even at the high chlorophyll concentrations required for efficient energy transfer, by restricting the range of orientations that neighboring chlorophyll pairs can adopt.

59 BASIC BIOLOGICAL SCIENCES↗

Wind effects on coastal zone color scanner chlorophyll patterns in the U.S. Mid-Atlantic Bight during spring 1979

Coastal zone color scanner (CZCS) chlorophyll concentration increases in the Mid-Atlantic Bight were associated with high wind speeds in continental shelf waters during March and May 1979. Maximum spring CZCS chlorophyll concentrations occurred during April when the water column was not thermally stratified and were spatially and temporally associated with reductions in wind speed both in onshelf and in offshelf regions. Increased chlorophyll concentrations in offshelf waters were associated with high wind speeds during May when a deep chlorophyll maximum was present. Chlorophyll patchiness was observed on length scales typical of those controlled by biological processes during the April low-wind period but not during March or May when wind speeds were greater. The spring CZCS chlorophyll maximum in the southern portion of the Mid-Atlantic Bight occurred in response to a reduction in mixed layer depth caused by decreased wind speeds and not by increased water column stratification.

Eslinger, David L.↗

Aircraft laser derived chlorophyll distribution across the Iceland-Faeroe front

The ocean surface color (water-leaving radiances) and thermal structure across the Iceland-Faeroe Front under both clear and cloudy conditions were measured from a NASA research aircraft on 25 May 1989. The measurements were made along four north-south lines that were 125 km in length and spaced 35 km apart. The color measurements were made with a 14 channel, non-polarized Multispectral Airborne Radiometer System (MARS) while the thermal data were collected by a thermal radiometer and aircraft bathythermographs. The satellite imagery (NOAA AVHRR) sequence show the development of meanders through the frontal region. These aircraft ocean color and thermal data characterize the biological distribution and are closely coupled to the physical processes occurring in the frontal systems. The ratio of several channels of the ocean color data are used to determine the surface chlorophyll. The retrieved data correlate well with laser-induced chlorophyll fluorescence obtained at the same time and historical chlorophyll data. The observed chlorophyll patchiness appearing across the Iceland-Faeroe Front is believed to be directly and indirectly related to primary and secondary circulation processes in ocean frontal systems. High chlorophyll concentrations were observed on the north side of the front and are inferred to be the result from the advection of Icelandic Coastal Water into the region. The sharp chlorophyll decline at and south of the thermal frontal boundary is clearly related to the subsurface thermal structure.

Arnone, Robert A.↗

Global Seasonal Climatologies of Ocean Chlorophyll: Blending In situ and Satellite Data for the CZCS Era

The historical archives of in situ (National Oceanographic Data Center) and satellite (Coastal Zone Color Scanner) chlorophyll data were combined using the blended analysis method of Reynolds [1988] in an attempt to construct an improved climatological seasonal representation of global chlorophyll distributions. The results of the blended analysis differed dramatically from the CZCS representation: global chlorophyll estimates increased 8-35% in the blended analysis depending upon season. Regional differences were even larger, up to 140% in the equatorial Indian Ocean in summer (during the southwest monsoon). Tropical Pacific chlorophyll values increased 25-41%. The results suggested that the CZCS generally underestimates chlorophyll. Regional and seasonal differences in the blended analysis were sufficiently large as to produce a different representation of global chlorophyll distributions than otherwise inferred from CZCS data alone. Analyses of primary production and biogeochemical cycles may be substantially impacted by these results.

Gregg, Watson W.↗

Multispectral In-situ Measurements of Organic Matter and Chlorophyll Fluorescence in Seawater: Documenting the Intrusion of the Mississippi River Plume in the West Florida Shelf

We performed multispectral in-situ fluorescence measurement of colored dissolved organic matter and chlorophyll in surface water of the West Florida Shelf using West Labs Spectral absorption and Fluorescence Instrument (SAFIre). Continuous measurements underway allowed us to simultaneously map the dispersion of riverine organic material and chlorophyll on the shelf. By using two fluorescence emission ratios we were able to differentiate between riverine and marine CDOM. Our data also showed unusually high concentrations of CDOM offshore. These were attributed to an intrusion of the Mississippi River Plume. We performed limited comparisons between in-situ chlorophyll concentrations measured with SAFIre and chlorophyll values obtained from SeaWiFS satellite data using OC4 and MODIS algorithm. Our results show that, although both algorithms overestimated chlorophyll, MODIS performed better than OC4, particularly in areas with high CDOM concentrations. Analysis of the relationship between chlorophyll and CDOM concentrations within the study area showed regional variability causes by differences in river source.

DelCastillo, Carlos E.↗

Sampling Biases in MODIS and SeaWiFS Ocean Chlorophyll Data

Although modem ocean color sensors, such as MODIS and SeaWiFS are often considered global missions, in reality it takes many days, even months, to sample the ocean surface enough to provide complete global coverage. The irregular temporal sampling of ocean color sensors can produce biases in monthly and annual mean chlorophyll estimates. We quantified the biases due to sampling using data assimilation to create a "truth field", which we then sub-sampled using the observational patterns of MODIS and SeaWiFS. Monthly and annual mean chlorophyll estimates from these sub-sampled, incomplete daily fields were constructed and compared to monthly and annual means from the complete daily fields of the assimilation model, at a spatial resolution of 1.25deg longitude by 0.67deg latitude. The results showed that global annual mean biases were positive, reaching nearly 8% (MODIS) and >5% (SeaWiFS). For perspective the maximum interannual variability in the SeaWiFS chlorophyll record was about 3%. Annual mean sampling biases were low (<3%) in the midlatitudes (between -40deg and 40deg). Low interannual variability in the global annual mean sampling biases suggested that global scale trend analyses were valid. High latitude biases were much higher than the global annual means, up to 20% as a basin annual mean, and over 80% in some months. This was the result of the high solar zenith angle exclusion in the processing algorithms. Only data where the solar angle is <75deg are permitted, in contrast to the assimilation which samples regularly over the entire area and month. High solar zenith angles do not facilitate phytoplankton photosynthesis and consequently low chlorophyll concentrations occurring here are missed by the data sets. Ocean color sensors selectively sample in locations and times of favorable phytoplankton growth, producing overestimates of chlorophyll. The biases derived from lack of sampling in the high latitudes varied monthly, leading to artifacts in the apparent seasonal cycle from ocean color sensors. A false secondary peak in chlorophyll occurred in May-August, which resulted from lack of sampling in the Antarctic.

Gregg, Watson W.↗

Evaluation of Vertical Patterns in Chlorophyll-A Derived From A Data Assimilating Model of Satellite-Based Ocean Color

Satellite-based sensors of ocean color have become the primary tool to infer changes in surface chlorophyll, while BGC-Argo floats are now filling the information gap at depth. Here we use BGC-Argo data to assess depth-resolved information on chlorophyll-a derived from an ocean biogeochemical model constrained by the assimilation of surface ocean color remote sensing. The data-assimilating model replicates well the general seasonality and meridional gradients in surface and depth-resolved chlorophyll-a inferred from the float array in the Southern Ocean. On average, the model tends to overestimate float-based chlorophyll, particularly at times and locations of high productivity such as the beginning of the spring bloom, subtropical deep chlorophyll maxima, and non-iron limited regions of the Southern Ocean. The highest model RMSE in the upper 50 m with respect to the float array is of 0.6 mg Chl m −3 , which should allow the detection of seasonal changes in float-based biomass (varying between 0.01 and >1 mg Chl m −3 ) but might hinder the identification of subtle changes in chlorophyll at narrow local scales. Both model and float profiling data show good agreement with in situ data from station ALOHA, with model estimates showing a slight accuracy edge in inferring depth-resolved observations. Uncertainties in float bio-optical estimates impede their use as a reliable benchmark for validation, but the general qualitative agreement between model and float data provides confidence in the ability of model to replicate biogeochemical features below the surface, where data is not directly constrained by the assimilation of satellite ocean color.

Lionel A Quintero↗

Unique contributions of chlorophyll and nitrogen to predict crop photosynthetic capacity from leaf spectroscopy

The photosynthetic capacity or the CO 2 -saturated photosynthetic rate (Vmax), chlorophyll, and nitrogen are closely linked leaf traits that determine C 4 crop photosynthesis and yield. Accurate, timely, rapid, and non-destructive approaches to predict leaf photosynthetic traits from hyperspectral reflectance are urgently needed for high-throughput crop monitoring to ensure food and bioenergy security. Therefore, this study thoroughly evaluated the state-of-the-art physically based radiative transfer models (RTMs), data-driven partial least squares regression (PLSR), and generalized PLSR (gPLSR) models to estimate leaf traits from leaf-clip hyperspectral reflectance, which was collected from maize (Zea mays L.) bioenergy plots with diverse genotypes, growth stages, treatments with nitrogen fertilizers, and ozone stresses in three growing seasons. The results show that leaf RTMs considering bidirectional effects can give accurate estimates of chlorophyll content (Pearson correlation r=0.95), while gPLSR enabled retrieval of leaf nitrogen concentration (r=0.85). Using PLSR with field measurements for training, the cross-validation indicates that V max can be well predicted from spectra (r=0.81). Here, the integration of chlorophyll content (strongly related to visible spectra) and nitrogen concentration (linked to shortwave infrared signals) can provide better predictions of V max (r=0.71) than only using either chlorophyll or nitrogen individually. This study highlights that leaf chlorophyll content and nitrogen concentration have key and unique contributions to V max prediction.

59 BASIC BIOLOGICAL SCIENCES↗

Chlorophyll Algorithms for Ocean Color Sensors - OC4, OC5 and OC6

A high degree of consistency and comparability among chlorophyll algorithms is necessary to meet the goals of merging data from concurrent overlapping ocean color missions for increased coverage of the global ocean and to extend existing time series to encompass data from recently launched missions and those planned for the near future, such as PACE, OLCI, HawkEye, EnMAP and SABIA-MAR. To accomplish these goals, we developed 65 empirical ocean color (OC) chlorophyll algorithms for 25 satellite instruments using the largest available and most globally representative database of coincident in situ chlorophyll a and remote sensing reflectances. Excellent internal consistency was achieved across these OC ‘Version -7’ algorithms, as demonstrated by a median regression slope and coefficient of determination (R (sup 2)) of 0.985 and 0.859, respectively, among 903 pairwise comparisons of OC-modeled chlorophyll. SeaWiFS and MODIS-Aqua satellite-to-in situ match-up results indicated equivalent, and sometimes superior, performance to current heritage chlorophyll algorithms.

ocean color↗

De novo design of proteins housing excitonically coupled chlorophyll special pairs

Natural photosystems couple light harvesting to charge separation using a ‘special pair’ of chlorophyll molecules that accepts excitation energy from the antenna and initiates an electron-transfer cascade. To investigate the photophysics of special pairs independently of the complexities of native photosynthetic proteins, and as a first step toward creating synthetic photosystems for new energy conversion technologies, we designed C 2 -symmetric proteins that hold two chlorophyll molecules in closely juxtaposed arrangements. X-ray crystallography confirmed that one designed protein binds two chlorophylls in the same orientation as native special pairs, whereas a second designed protein positions them in a previously unseen geometry. Spectroscopy revealed that the chlorophylls are excitonically coupled, and fluorescence lifetime imaging demonstrated energy transfer. The cryo-electron microscopy structure of a designed 24-chlorophyll octahedral nanocage with a special pair on each edge closely matched the design model. The results suggest that the de novo design of artificial photosynthetic systems is within reach of current computational methods.

59 BASIC BIOLOGICAL SCIENCES↗

A family of α/β hydrolases removes phytol from chlorophyll metabolites for tocopherol biosynthesis in Arabidopsis

Tocopherol synthesis requires phytyl diphosphate derived from phytol esterified to chlorophyll metabolites. The >600-member Arabidopsis thaliana α/β hydrolase (ABH) gene family contains 4 members that can release phytol from chlorophyll metabolites in vitro; however, only pheophytinase (PPH) affects tocopherol synthesis when mutated, reducing seed tocopherols by 5%. We report the biochemical analysis of 2 previously uncharacterized ABHs, chlorophyll dephytylase 2 (CLD2) and CLD3, and their respective mutants singly and in combinations with pph and cld1 alleles. While all CLDs localized to the thylakoid and could hydrolyze phytol from chlorophylls and Pheophytin a in vitro, CLD3 had the highest in vitro activity and the largest effect on tocopherol synthesis in vivo. The 3 CLDs acted cooperatively to provide phytol for 31% of tocopherols synthesized in light-grown leaf tissue. Dark-induced leaf senescence assays showed PPH is required for 18% of the tocopherols synthesized. Though the cld123 triple mutant had no impact on dark-induced tocopherol content, cld123 in the pph background reduced tocopherol levels by an additional 18%. In seeds, pph and cld123 each reduced tocopherol content by 5% and by 15% in the cld123pph quadruple mutant. VTE7 ( ViTamin E7 ) is an envelope-localized ABH that specifically affects chlorophyll biosynthetic intermediates in vivo and is required for 55% of seed tocopherol synthesis. The introduction of cld123pph into the vte7 background further reduced seed tocopherol levels to 23% of that of the wild type. Our findings demonstrate that phytol provision for tocopherol biosynthesis and homeostasis is a complex process involving the coordinated spatiotemporal expression of multiple ABH family members.

arabidopsis↗

Data from Unique Contributions of Chlorophyll and Nitrogen to Predict Crop Photosynthetic Capacity from Leaf Spectroscopy

The photosynthetic capacity or the CO2-saturated photosynthetic rate (Vmax), chlorophyll, and nitrogen are closely linked leaf traits that determine C4 crop photosynthesis and yield. Accurate, timely, rapid, and non-destructive approaches to predict leaf photosynthetic traits from hyperspectral reflectance are urgently needed for high-throughput crop monitoring to ensure food and bioenergy security. Therefore, this study thoroughly evaluated the state-of-the-art physically based radiative transfer models (RTMs), data-driven partial least squares regression (PLSR), and generalized PLSR (gPLSR) models to estimate leaf traits from leaf-clip hyperspectral reflectance, which was collected from maize (Zea mays L.) bioenergy plots with diverse genotypes, growth stages, treatments with nitrogen fertilizers, and ozone stresses in three growing seasons. The results show that leaf RTMs considering bidirectional effects can give accurate estimates of chlorophyll content (Pearson correlation r=0.95), while gPLSR enabled retrieval of leaf nitrogen concentration (r=0.85). Using PLSR with field measurements for training, the cross-validation indicates that Vmax can be well predicted from spectra (r=0.81). The integration of chlorophyll content (strongly related to visible spectra) and nitrogen concentration (linked to shortwave infrared signals) can provide better predictions of Vmax (r=0.71) than only using either chlorophyll or nitrogen individually. This study highlights that leaf chlorophyll content and nitrogen concentration have key and unique contributions to Vmax prediction.

Biomass Analytics↗

Mapping of chlorophyll a distributions in coastal zones

It is pointed out that chlorophyll a is an important environmental parameter for monitoring water quality, nutrient loads, and pollution effects in coastal zones. High chlorophyll a concentrations occur in areas which have high nutrient inflows from sources such as sewage treatment plants and industrial wastes. Low chlorophyll a concentrations may be due to the addition of toxic substances from industrial wastes or other sources. Remote sensing provides an opportunity to assess distributions of water quality parameters, such as chlorophyll a. A description is presented of the chlorophyll a analysis and a quantitative mapping of the James River, Virginia. An approach considered by Johnson (1977) was used in the analysis. An application of the multiple regression analysis technique to a data set collected over the New York Bight, an environmentally different area of the coastal zone, is also discussed.

Johnson, R. W.↗