Optical characterization of primary productivity across a coastal front

Methods for the remote estimation of phytoplankton biomass and production rates using multiplatform sampling strategies are essential for the better understanding of oceanic bio-geochemical cycles. Recent advances in remote sensing of ocean color have made synoptic estimation of phytoplankton biomass attainable. While considerable success has been achieved in the estimation of plant biomass, the synoptic estimation of phytoplankton rates of production has been inadequate. Rapid shipboard estimates of the vertical distribution of primary productivity, on mesoscale spatial scales and event-time scales, are needed to provide both surface validation and data for the development of bio-optical models linking production to the optical characteristics of the water column. This study details the primary productivity and optical properties of a frontal region in July 1985 along 35°50N in the Southern California Bight which is shown to be consistent with the concurrent high-performance liquid-chromatography pigment-analysis. We describe here a quasi-synoptic shipboard bio-optical sampling strategy across a frontal region as an example of time-corrected data for assessing phytoplankton production in highly variable ocean regions.     

Spring and summer growth rates of subarctic Pacific phytoplankton assemblages determined from carbon uptake and cell volumes estimated using epifluorescence microscopy

In order to determine whether phytoplankton growth rates were normal or depressed, total plant carbon (g l^–1) and in situ production rates (g C l^–1 d^–1) were measured for phytoplankton assemblages at Weathership Station P (50°N; 145°W) and at 53°N; 145°W in the subarctic Pacific in May and August 1984. Plant carbon, estimated from cell volumes determined using epifluorescence microscopy, was distributed as follow: 28% in the <2 m fraction, 38% in the 2 to 5 m size fraction, and the remainder in size classes >5 m. Carbon-specific growth rates (k), as doublings d^–1, were calculated for the phytoplankton assemblages as a whole at each sampling depth down to 100 m for three days in May and for four days in August. The populations in the upper part of the euphotic zone showed average doubling rates of 1 d^–1 and thus appeared to be growing at rates normally expected for the prevailing conditions of light and temperature. The low chlorophyll concentrations (0.3 to 0.4 mg chl a m^–3) characteristically found in this oceanic region do not seem to be due to very slow growth of algal populations.Contribution No. 1695 of the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Sample support and related scale issues in composite sampling

Composite sampling offers the great promise of efficiency and economy in environmental decision making. However, if careful attention is not paid to matching the support of the sample to that required for making the desired decision, the promise is unfulfilled. Obviously this attention must be applied in the design phase of a composite sampling strategy. Less obvious is the potential for alteration of the design sample support during sample collection and assay. The consequences of not paying attention to these aspects of sample design and assay are discussed in this issue paper illustrated with a series of examples taken from the authors consulting experience.     

Interactions between nutritional status and long-term responses to ultraviolet-B radiation stress in a marine diatom

Influences of nutritional status on the photoinhibitory effects of ultraviolet-B radiation (UVBR: 290 to 320 nm) on the specific growth rates (_obs) and biomass of Phaeodactylum tricornutum were determined using nutrient-replete batch cultures and nutrient-limited continuous cultures. P. tricornutum cultures were exposed to UVBR doses representative of current mid-latitude and ozone-depletion intensities. Specific growth rates and biomass were inhibited from 2 to 16% by UVBR during nutrient-replete growth. However, no effect of UVBR on _obs or biomass was detectable when nutrient limitation exceeded the potential for limitation by UVBR. Thus, a competitive interaction appears to occur between macronutrient stress and UVBR stress, such that _obs and biomass will be determined by the most limiting factor. Our results suggest that measurable decreases in phytoplankton _obs and biomass from UVBR are most likely in nutrient-rich areas of the ocean, while these parameters may not be appropriate for measuring UVBR stress in regions of nutrient limitation.     

Some experiments on phosphate assimilation by coastal marine plankton

A study of phosphate assimilation by coastal marine plankton revealed that both phytoplankton and microheterotrophs incorporated radioactive phosphorus (³³P). Size fractionation of the particulate matter (using 1 m pore diameter Nucleopore^® membrane filters), antibiotic treatment (using garamycin), and independent estimaties of photoautotrophic (¹⁴CO₂ uptake) and heterotrophic (³H-glucose uptake) activities were employed to separate phyto- and bacterioplankton phosphate uptake. Results indicated that phytoplankton ³³P-uptake was best estimated by the fraction of particulate matter retained on the 1 m membrane filters. Usually, less than 10% of the phytoplankton (based on chlorophyll a measurements) passed the 1 m pore-diameter filters, whereas about 90% of the heterotrophic activity passed. At least 50% of the ³³P-uptake was associated with the <1 m fraction. It may be possible to resolve the phytoplankton and bacterial contributions to ³³P-uptake by comparing the percent of total ³³P-uptake with the percent of total ³H-glucose uptake associated with the >1 m fraction.     

Excretion of ammonia by zooplankton and its potential contribution to nitrogen requirements for primary production in the Catalan Sea (NW Mediterranean)

Excretion of ammonia by mesozooplankton (>200 m zooplankton) and its potential contribution to the nitrogen requirement for phytoplankton growth has been estimated for different hydrographical situations along a transect across the Catalan Sea (Northwestern Mediterranean). The nitrogen excreted as ammonia was estimated from mesozooplankton biomass and specific excretion rates. Nitrogen requirements of phytoplankton were estimated by means of carbon fixation rates and C:N ratios of <200 m particulate organic matter. Minimum C:N ratios and maximum primary production, zooplankton biomass, phytoplankton nitrogen requirements, and nitrogen excretion of zooplankton occurred near the Catalan density front. On average, the nitrogen regenerated by the mesozooplankton accounted for 43% of the nitrogen requirements of the phytoplankton. The specific excretion rates of ammonia and the percentage of phytoplanktonnitrogen requirements supplied by excreted nitrogen were higher at coastal stations. In some coastal and frontal stations, the ammonia excreted exceeded the phytoplanktonnitrogen demand. Bacteria competing for nutrient supply and the possible uncoupling between rate processes and standing stocks of phyto- and zooplankton could explain the apparent excess of regenerated ammonia.     

Patterns of photosynthetic carbon metabolism in light-limited phytoplankton

Patterns of phytoplankton carbon (C) metabolism were examined in å combined laboratory and field study to assess the influence of light conditions on ¹⁴C assimilation into photosynthetic end-products. Laboratory studies with three species representing distinct size classes and taxonomic groups tested the influence of low light on patterns of C flow. Prorocentrum mariae-lebouriae (dinoflagellate) and Ditylum brightwellii (diatom) showed decreased movement of photoassimilated ¹⁴C into protein following a shift to low light ¹⁴C assimilation into lipids and photosynthetic pigments increased in low light and was paralleled by increased chl a per cell. The proportion of ¹⁴C fixed into protein returned to the pre-shift level upon return to initial light conditions. Monochrysis lutheri (chrysophyte) did not show this pattern of reduced % ¹⁴C protein. Incubations of 12 and 24 h demonstrated significant rearrangements in labeling patterns at night, wherein ¹⁴C flow into protein in darkness was favored. % ¹⁴C protein at night was lower for M. lutheri than for the other species, suggesting some interspecific differences in the low light response. Measurements of ¹⁴C assimilation in phytoplankton assemblages from Chesapeake Bay demonstrated movement of a higher proportion of photo-assimilated C into protein in samples collected in the surface mixed layer than in those below the pycnocline. In comparison, phytoplankton collected below the pycnocline fixed a higher proportion of ¹⁴C into lipids, photosynthetic pigments, and low molecular weight metabolites, as was observed in low light laboratory cultures. A comparison of 12- and 24-h incubations for measuring patterns of C flow into photosynthetic end-products confirmed the inadequacy of short-term measurements, as significant changes in ¹⁴C allocation occurred in the dark phase of the photocycle. Together, these results suggest that ¹⁴C assimilation into photosynthetic end-products can be a useful measure of adaptive state in changing light conditions, but point out some difficulties in applying this approach in situ.     

Benthic response to sedimentation events during autumn to spring at a shallow water station in the Western Kiel Bight

Seasonal variations in bacterial populations (total number, biomass, biomass-spectrum, number of dividing cells) as well as in concentrations and decomposition rates of particulate organic material were followed in a sandy mud sediment of the Western Kiel Bight (Baltic Sea; FRG). The strong seasonal variations observed could be traced back to the effect of certain ecological situations and events in the sediment from which the input of the phytoplankton blooms in autumn and spring, respectively, the accumulation of organic material during winter, and the spring development of the benthic fauna turned out to be the most important. Bacterial carbon net production following the breakdown of the phytoplankton blooms ranged between 9 g (autumn) and 16 g (spring) per g of dry weight sediment per day. The consequences of shifts in the size composition of the bacterial populations as well as the importance of the measurement of enzymatic decomposition rates of particulate organic material in sediments are demonstrated and discussed in relation to the events mentioned above.Publication No. 420 of the Joint Research Program at Kiel University (Sonderforschungsbereich 95 der Deutschen Forschungsgemeinschaft)     

Evaluation of Public Housing Lead Risk Assessment Data

A unique data set from lead risk assessments performed on 67 public housing developments from across the United States was made available for analyzes. The data set includes results of lead analysis from 5906 dust wipes and from 1222 soil samples. A total of 487 dwelling units in these developments, as well as associated common areas, were sampled, all by the same team of inspectors. The number of dwelling units within a development that were sampled reflected the guidelines then in force, the 1990 Interim HUD Guidelines, rather than those specified in the 1995 Guidelines. Median dust lead loadings for floors, 151gm^–2 (14gft^–2), and window sills, 936gm^–2 (87gft^–2), were much less than former HUD limits of 1076gm^–2 (100gft^–2) and 5380gm^–2 (500gft^–2), respectively and are only about one-third of the recently established limits of 431gm^–2 (40gft^–2) and 2690gm^–2 (250gft^–2). In contrast, the median lead loading for window troughs, 8560gm^–2 (795gft^–2), was almost identical to the HUD clearance limit of 8610gm^–2 (800gft^–2). There was a strong positive correlation between floor and window trough lead loading values for samples from the same dwelling units and those from common areas of the housing developments. Door threshold samples, which may reflect conditions exterior to the dwelling unit, were collected from 53 dwelling units. Median lead loading levels of these samples were more than ten times higher than those in floor samples from the same dwelling units, were about the same as window sill samples and about one-half of levels in window trough samples. Composite sample results, simulated by averaging results from four samples within a dwelling unit, revealed that in order to have the same rate of excedence of standards, the composite standards would have to be reduced, for example, from the single sample value of 1076gm^–2 (100gft^–2) to 527gm^–2 (49gft^–2) for floor samples and from the single sample value of 8610gm^–2 (800gft^–2) to 5160gm^–2 (479gft^–2) for window troughs. For this public housing data set, the portion of the units in developments containing more than 225 units which exceeded the established limit for window samples was the same when using either the full data set or a random one-half of the data set. This suggests that, for this data set, the number of dwelling units sampled was excessive . Thus, the required increase in the number of dwelling units to be sampled specified in the 1995 Guidelines for developments with more than 225 dwelling units, may not have been necessary if this data set is representative of public housing developments in the United States.     

Hydrobiological parameters during an annual cycle in the Arcachon Basin

Temperature, salinity, nutrients and phytoplankton biomass were monitored on a weekly to bimonthly frequency at six stations in the Bay of Arcachon from July 1984 to July 1985. This particular period appears to have differed from the last ten years in displaying higher amplitudes of both temperature and salinity (1.5° to 24.5°C and 20 to 34.5) at high tide. However, although in spring both temperature and salinity were normal (14°C, 28), an important phytoplankton spring bloom occurred, with maximum chlorophyll levels reaching 15 g l^-1 in April. Utilization of nutrients was high, particularly for nitrate and silicate, the concentrations of which decreased, respectively, from 10–15 to 0.1–2 M and 10–20 to 0.25–3 M from February to May. Exhaustion of nitrate was observed in May, except in areas subjected to river input. In contrast, silicate increased throughout the study area from May to July.     

Influence of irradiance on the nutritional value of two phytoplankton species fed to larval Japanese scallops (Patinopecten yessoensis)

Japanese scallop (Patinopectin yessoensis Jay) larvae grew faster and were larger after 18 d when fed a diet of high-light(HL)-grown Chaetoceros simplex or HL Pavlova lutheri relative to diets of the same phytoplankton species grown at low light (LL). When provided as saturating rations to larval scallop, these diets could be ranked: HL C. simplex>LL C. simplex>HL P. lutheri>LL P. lutheri. In both phytoplankton species, HL-grown cells contained more of the short-chain saturated fatty acid (FA), 16:0 than LL-grown cells. Scallop growth rates were a significant function of the amounts (mg g^-1 dry wt) and the proportions (as percentage of total FAs) of the FAs 14:0 and 14:0+16:0 (total saturated FAs) in their diet. The proximate biochemical composition of HL- versus LL-grown phytoplankton showed no significant differences in protein, total lipid, carbon, carbohydrate or nitrogen per cell which were consistently associated with the greater nutritional value of HL cells. In spite of this high variability in proximate composition, the larval growth rate was a significant function of the average carbon content, nitrogen content and cell volume of the phytoplankton cells. Increased amounts of the essential polyunsaturated FAs 20:5 3 and 22:6 3 in the phytoplankton were negatively correlated with larval scallop growth rates. Thus HL-grown phytoplankton cells were nutritionally superior to LL-grown cells. This nutritional superiority seems to be determined by the fatty acid composition of the cells which, in turn, is controlled by variation in irradiance. The general tendency of predator FA profiles to resemble that of their prey was not observed in larvae fed P. lutheri. The much greater amounts of 18:4 3, 20:5 3, and 22:6 3 FA in P. lutheri relative to C. simplex were not evident in the scallop larvae fed these cells.     

Nutrient cycling studies in Carmarthen Bay: phytoplankton production,nitrogen assimilation and regeneration

The results from a series of five cruises (August 1979–January 1981) to examine rates of primary production, nitrogen assimilation and regeneration in Carmarthen Bay, S. Wales, are presented. Chlorophyll, primary production and regeneration were distributed irregularly throughout the bay, with the highest concentrations and rates being found around the shallow periphery. Nitrate was the dominant form of nitrogen assimilated by the phytoplankton, apart from periods of high biomass during the summer when ammonium assumed a more important role. The rates of ammonium regeneration from microheterotrophs were greater than the ammonium demand over much of the year, indicating that the bay was a net exporter of ammonium and was a physically dominated open system. During the summer, the rates of ammonium regeneration and assimilation were similar, indicating efficient recycling of nutrients.     

Size classes and major taxonomic groups of phytoplankton at two locations in the subarctic pacific ocean in May and August, 1984

In order to assess the relative importance of the pico- and nanoplankton fractions, the composition of entire phytoplankton communities at Weathership Station P (50°N; 145°W) and at 53°N; 145°W were studied in May and August, 1984, using epifluorescence, scanning electron, and inverted light microscopy. The biomass of major taxa within five size classes was estimated from cell volume and cell concentration. For both months, approximately twothirds of the total phytoplankton carbon were contributed by cells<5 m. In May, 16% of plant biomass was contributed by cells<2 m, and in August 39%. (In both months 90% of plant carbon<2 m was contributed by the bluegreen coccoid Synechococcus spp.) Cells 2 to 5 m contributed about 39% to total plant carbon; they were mostly flagellates in May and nonmotile coccoids in August. The remaining one-third of algal carbon was composed of dinoflagellates, cryptomonads, other flagellates and diatoms, all >5 m. Very little difference between taxa was observed with respect to vertical stratification. Small taxonomic changes were observed in the community between May and August, and within each month.Contribution No. 1694 of the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Phosphorus dynamics associated with phytoplankton blooms in eutrophic Mikawa Bay,Japan

Changes in the phosphorus components of the particulate matter in seawater were studied in the eutrophicated waters of Mikawa Bay, Japan, during summer 1981. The contents of particulate phosphorus and hot-water extractable intracellular phosphorus displayed remarkable changes associated with phytoplankton blooms caused by wind-induced or upwelling-associated nutrient enrichment from the bottom water layers. Nanoplankton <10 m accounted for much of the particulate phosphorus (70 to 79% in June and July, and 44 to 78% in August and September); the contribution of large-sized phytoplankton >25 m varied from 9 to 49%, the peak values being attained under red-tide conditions. The capacity for phosphorus storage in cells was low in nanoplankton cells, high in large phytoplankton species. Differences in rates of phosphorus storage and growth between nanoplankton and large phytoplankton accounted for fluctuations in particulate phosphorus which were closely associated with fluctuations in phytoplankton blooms in Mikaw Bay.     

Swimming pattern as an indicator of the roles of copepod sensory systems in the recognition of food

The roles of copepod sensory systems in the recognition of food were investigated using the Bugwatcher, a video-computer system designed to track and describe quantitatively the swimming patterns of aquatic organisms. The swimming behavior of the copepodPseudocalanus minutus in the presence of phytoplankton is characterized by a decrease in average swimming speed and an increase in pause behaviors compared to its swimming behavior in filtered seawater. Copepods exposed to chemosensory stimulation alone (filtered phytoplankton exudate) exhibited an increase in average swimming speed and an increase in the number of burst swimming behaviors. When exposed to a novel, non-food chemosensory stimulus (morpholine), no change in swimming behavior was observed unless the copepods had been conditioned to this odor in the presence of phytoplankton. Copepods exposed to mechanosensory stimulation alone (plastic spheres) exhibited a decrease in swimming speed and an increase in pause behaviors. When exposed to both forms of stimulation simultaneously (phytoplankton exudate and plastic spheres), a further decrease in swimming speed and increase in pause behaviors occurs, yielding a swimming pattern similar to that found in the presence of phytoplankton. This analysis of swimming pattern indicates that both chemoreception and mechanoreception contribute to the recognition of food inP. minutus.Contribution No. 406 of the US EPA Environmental Research Laboratory; Narragansett, Rhode Island 02882, USA     

Isotopic characterization of atmospheric nitrogen inputs as sources of enhanced primary production in coastal Atlantic Ocean waters

Current estimates indicate that atmospheric nitrogen deposition is responsible for 26 to over 70% of new nitrogen (N) input to North Carolina estuaries and coastal waters. Concentrations of N in coastal rainfall events in a 2-yr period (August 1990 to 1992) ranged from 0.7 to 144 M for NO ₃ ^- and 0.5 to 164 M for NH ₄ ⁺ . The ¹⁵N values of the NO ₃ ^- and the NH ₄ ⁺ were determined in 15 rain events. NH₄ ⁺ values averaged-3.13 (range:-12.5 to+3.6), while NO ₃ ^- plus dissolved organic N fractions had an average ¹⁵N of+1.0 (range:-2.0 to+4.7). The uptake of this isotopically light N into particulate N, in parallel with primary productivity and biomass (as chlorophyll a) determinetions, was examined in microcosm and mesocosm bioassays. As phytoplankton productivity and biomass increased with added rainwater N, the ¹⁵N of particulate N decreased. To investigate the effects of significant atmospheric N loading with stable isotope tracers, we measured the ¹⁵N of the>1 m fraction from surrounding coastal waters. Owing to the episodic nature of atmospheric deposition and the great variation in N loading with each event, a simple assessment of the atmospheric contribution was not possible. During a period in which rainfall inputs were significant and frequent (August 1992), ¹⁵N values were several more negative than during periods of drought (Fall 1990). These experiments and observations emphasize the contribution of atmospheric nitrogen deposition to new production in coastal waters.     

The distribution of heavy metals in deposited urban dusts and sediments, Coventry, England

Samples of urban dusts and sediments have been collected and categorised into three compartment those which act mainly as sources, those undergoing transport and those which have been deposited. The sediments were fractionated into <63m to match similar sized lacustrine sediments and <2mm to represent natural and anthropogenic fluvially transported sediments. Preliminary results of total heavy metals analysis show that the <2mm fraction of the source group exhibits the greatest variabilty in metal concentrations. Overall, however, the <63m fraction dominates regardless of compartment. Some heavy metal concentrations decrease through the sourcetransportdeposit process, possibly due to selective onward transport of finer material or other anthropogenic processes unique to the urban environment (e.g. street cleansing). Sitespecificity is indicated in a comparison with other similar studies.     

Evidence for multiple recruitment-cohorts of Puget Sound Dungeness crab,Cancer magister

While sampling intertidally in Puget Sound, Washington, USA, for juvenile Dungeness crab (Cancer magister) in 1984, we found evidence of two distinct cohorts of the same year-class based on sizes of first-instar juveniles (J1) and the spatial/temporal patterns of settlement. In 1988, three distinct cohorts were observed to settle in Puget Sound and its approaches. Settlement of one cohort occurred during May in the Strait of Juan de Fuca and in those areas of Puget Sound closest to the Strait. J1 individuals of this cohort were large (x=7.4 mm carapace width, CW) and comparable in both size and timing of settlement to populations along the Washington coast (e.g. Grays Harbor and Willapa Bay). Initial settlement density of the May cohort was as high as 215 crabs/m² in intertidal eelgrass beds along the Strait of Juan de Fuca and decreased to <2 crabs/m² within Puget Sound and the Strait of Georgia. A second cohort apparently originated in Hood Canal (a deep inland fjord), its size upon settlement in June was significantly smaller (J1 x= 5.3 mm CW) than the May cohort, and it was limited to Hood Canal and areas of Puget Sound close to the mouth of Hood Canal. A third cohort, which settled in late July and August, was the smallest of the three cohorts (J1 x= 4.8 mm CW), and was widely distributed around Puget Sound from Seattle in the south to the USA/Canadian border in the north. We hypothesize that most juvenile recruitment in Hood Canal and Puget Sound originates from parental stocks endemic to their respective basins (Hood Canal and Puget Sound cohorts), but that, on occasion, oceanographic conditions allow substantial influx of Pacific Ocean Dungeness crab larvae (oceanic cohort) through the Strait of Juan de Fuca into Puget Sound. Tracking of spatial/temporal settlement patterns and comparison of J1 sizes proved useful for estimating the probable sources and dispersion of Dungeness crab larvae. Differences in size and time of settlement between various larval cohorts of C. magister may prove useful as biomarkers for tracing circulation patterns within and between inland waters of Washington and the Pacific Ocean. Causes of smaller size and later settlement of the Puget Sound cohort relative to oceanic conspecifics of the same year-class are discussed.Contribution No. 856, School of Fisheries, University of Washington, Seattle     

Production and respiration of overall plankton and ultraplankton communities at the entrance to the Gulf of Finland in the Baltic Sea

Primary productivity and respiration of the overall plankton community and of ultraplankton (organisms passing through a 3-m Nuclepore filter) were studied at the entrance to the Gulf of Finland during the growth season in 1982. Data of the respiration measurements from previous years are also presented. During the development of a diatom spring bloom, the algal component could be successfully separated from the bacterial component by size fractionation with a 3-m Nuclepore filter and thus the algal respiration could be approximated, being on the order of 10 to 20% of the gross production. After the phytoplankton spring maximum, bacteria played an important role in mediating the energy flow from phytoplankton exudates to higher trophic levels. Maximum values of 1 230 and 740 mg O₂ m^-2 d^-1 were recorded for overall and for ultraplankton respiration, respectively, during late July. High productivity values coupled with low phytoplankton biomass and low inorganic nutrient values were also recorded in late July, indicating effective nutrient regeneration and rapid turnover of the plankton community. During late summer, a considerable fraction (over 30%) of phytoplankton production was released as exudates, suggesting that much of the energy is channeled to higher trophic levels via bacterial pathways rather than by direct herbivorous grazing during this season. The summer development of phytoplankton community structure and functioning is strongly controlled by hydrographic conditions, i.e. by nutrient inputs via upwelling and by water temperature. A carbon budget for late summer indicated that bacteria may contribute only up to 50% of the overall respiration of the plankton community, which suggests that heterotrophs other than bacteria play an important role in nutrient regeneration. The present study stresses the importance of energy flow via the phytoplankton exudatebacteria-micrograzer pathway in relatively oligotrophic, brackish water ecosystems.     

“Copper” granules in the barnacle Balanus balanoides

Balanus balanoides (L.) collected from an area with high heavy-metal run-off contained two different types of granule within the parenchyma cells of the prosoma. X-ray microprobe analysis shows one to be the familiar zinc granule made up of concentric layers and giving major peaks for phosphorus and zinc, and the other to be homogeneous and giving peaks for sulphur and copper. This latter granule is designated the copper granule. Whilst zinc granules are known to be composed of inorganic phosphate, various tests on copper granules in sections and in a granule-rich pellet have shown that the copper is probably complexed with organic matter. The tests also demonstrated the relatively, insoluble (inert) nature of these granules. Although zinc and copper granules were present together in the prosoma, atomic absorption analyses of whole bodies (prosoma+thorax) have shown the level of zinc (50.28 g/mg dry weight) to be much higher than that of copper (3.75 g/mg dry weight).