Micro-zooplankton and its abundance relative to the larger zooplankton and other seston components

Micro-zooplankton populations in the upper 100 m were sampled from 5 marine environments in the northeast Pacific Ocean extending from slope waters off San Diego to an oceanic site near Isla Guadalupe, and their abundance related to that of the larger zooplankton, phytoplankton (as estimated from chlorophyll a), and detritus. The micro-zooplankton and other components of the seston were subdivided into 3 fractions on the basis of size in the deck-mounted collecting unit of a seawater pumping system. Through the euphotic zone at each site, the Protozoa, of which ciliates were the dominant forms, accounted for 95% or more of the total micro-zooplankton numbers. Their biomass, as volume, was estimated to be 13 to 28% of that of the total micro-zooplankton. The standing stock of micro-zooplankton over the euphotic zone at the various sites, in terms of dry weight, was estimated to be 14 to 34% (average 23%) of that of the phytoplankton crop. Micro-zooplankton volumes in the upper 100 m were 21 to 26% of those for the larger zooplankton sampled over the same depth.     

Release of dissolved organic matter from natural populations of marine phytoplankton

A new method is presented for measuring the rate of release of dissolved organic matter (DOM) produced by natural populations of marine phytoplankton. The method has been field-tested using natural populations of estuarine, coastal, and oceanic phytoplankton. Problems associated with the necessity for long incubation times, high initial activity of the inorganic ¹⁴C added to the sample, and self-absorption are overcome. Improved sensitivity is obtained by utilizing a large portion of the sample filtrate, Results from sampling at different times of day and for different lengths of incubation suggest that these problems should be reexamined in terms of both the percent and rate of relase of DOM from natural populations of phytoplankton. The rate of release of DOM from natural populations of marine phytoplankton decreases seaward. Phytoplankton in Georgia (USA) estuaries release <1 to 40 mg C m^-3 day^-1, in southeastern USA coastal waters from 0 to 19 mg C m^-3 day^-1, and in the westernmost Sargasso Sea from 0 to 2 mg C m^-3 day^-1. The percent of photoassimilated carbon released as DOM increases seaward. The percents are less than 7% in the Georgia estuaries, generally less than 13% in southeastern USA surface coastal waters, generally less than 21% for these coastal waters below the surface, and less than 44% for the westernmost Sargasso Sea. The results of this study and of other studies suggest that the measurement of primary productivity in estuarine and nearshore coastal waters is not seriously in error because of the absence of measurements on the rate of release of DOM from such phytoplankton. However, the estimation of primary-productivity values for oceanic waters is underestimated in some cases by about 50%.Contribution No. 595 from the Department of Oceanography, University of Washington, Seattle, Washington, USA and Contribution No. 224 from the University of Georgia Marine Institute, Sapelo Island, Georgia, USA.Extracted in part from a dissertation submitted to the Graduate Faculty of the University of Georgia in partial fulfillment of the requirements for the degree of Doctor of Philosophy.     

Switching between herbivory and carnivory by the planktonic marine copepod Calanus pacificus

Adult females of the omnivorous copepod Calanus pacificus, collected from the plankton off La Jolla, California, USA (June, 1978), fed disproportionately on the prey in greatest relative abundance when given mixtures of diatoms (Thalassiosira fluviatilis) and copepod (C. pacificus) nauplii as food. This switch from herbivorous to carnivorous behavior may be significant in nature during the decline of phytoplankton blooms. More generally, the widespread omnivorous habit among pelagic animals suggests a responsive and flexible trophic organization which contributes to the resiliency of planktonic communities in a dynamic physical environment.Contribution No. 1223 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Physioecology of zooplankton. III. Effects of phytoplankton concentration,temperature, and body size on the metabolic rate ofCalanus pacificus

Weight-specific rates of oxygen consumption of actively feeding copepodite stages ofCalanus pacificus Brodsky were measured under various combination of phytoplankton concentration and temperature. The rate decreased logarithmically with a logarithmic increase in dry body weight of copepods, and the relationship between these variables was described using a log-transformed allometric equation. The body-size dependence of the metabolic rate was independent of changes in food concentration and temperature, but the metabolic level increased linearly with a logarithmic increase in temperature and was not significantly affected by changes in food concentration. Respiration rates measured in this study forC. pacificus were about twice as high as rates reported for unfed closely related species of the same genus. An analysis of the metabolic cost of feeding processes suggests that metabolic models derived from feeding models may be of little ecological value at present.Contribution No. 1129 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Photoadaption in marine phytoplankton: Response of the photosynthetic unit

Some species of phytoplankton adapt to low light intensities by increasing the size of the photosynthetic unit (PSU), which is the ratio of light-harvesting pigments to P700 (reaction-center chlorophyll of Photosystem I). PSU size was determined for 7 species of marine phytoplankton grown at 2 light intensities: high (300 E m^-2 s^-1) and low (4 E m^-2 s^-1); PSU size was also determined for 3 species grown at only high light intensity. PSU size varied among species grown at high light from 380 for Dunaliella euchlora to 915 for Chaetoceros danicus. For most species grown at low light intensity, PSU size increased, while the percentage increase varied among species from 13 to 130%. No change in PSU size was observed for D. euchlora. Photosynthetic efficiency per chlorophyll a (determined from the initial slope of a curve relating photosynthetic rate to light intensity) varied inversely with PSU size. In contrast, photosynthetic efficiency per P700 was enhanced at larger PSU sizes. Therefore, phytoplankton species with intrinsically large PSU sizes probably respond more readily to the rapid fluctuations in light intensity that such organisms experience in the mixed layer.Contribution No. 1180 from the Department of Oceanography, University of Washington, Seattle, Washington, USA     

Pathways of arsenic uptake and incorporation in estuarine phytoplankton and the filter-feeding invertebrates Eurytemora affinis,Balanus improvisus and Crassostrea virginica

Arsenic uptake from water and from phytoplankton was followed in the copepod Eurytemora affinis and the barnacle Balanus improvisus collected from the Patuxent River estuary, Chesapeake Bay, eastern coast of the USA in 1987, and in the oyster Crassostrea virginica obtained from a hatchery on the shore of Chesapeake Bay in 1987. Dissolved arsenic was readily taken up by phytoplankton and by shell material of B. improvisus and C. virginica; however, no dissolved arsenic was incorporated into the invertebrate tissues. When E. affinis, B. improvisus and C. virginica were fed phytoplankton containing elevated arsenic contents, significant arsenic incorporation occurred. Juvenile B. improvisus incorporated relatively more arsenic than adults of all three species. Compared to the 100 to 200% increase in arsenic content by phytoplankton exposed to dissolved arsenic, the 25 to 50% increase in these invertebrate species via trophic transfer is relatively small. Even though the trophic pathway for arsenic transfer is the major one for higher trophic levels within an ecosystem, the potential for direct arsenic impact to trophic levels other than phytoplankton appears to be minimal.     

Persistent blooms of surf diatoms along the Pacific coast,USA

Productivity was studied in two diatom species, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which form persistent dense blooms in the surf zone along the Pacific coast of Washington and Oregon, USA. Past observations have shown that surf-diatom standing stock usually declines in summer along with concentrations of nitrate and ammonium. Using the ¹⁴C method, photosynthetic rates in natural surf samples were measured monthly for one year (October 1981 through September 1982) at a study site on the Washington coast. Also measured were temperature, salinity, dissolved nutrients, particulate carbon and nitrogen (used as estimates of phytoplankton C and N), and chlorophyll a. Assimilation numbers (P _max) were higher in summer (5 to 8 g C g^-1 chl a h^-1) than in winter (3 to 4gC). Specific carbon incorporation rates (µ_max) showed no obvious seasonality, mostly falling within the range of 0.09 to 0.13 g C g^-1 C(POC) h^-1. The discrepancy between the seasonal trends for chlorophyll-specific and carbon-specific rates reflects a change in the carbon-to-chlorophyll ratio. Because of seasonal differences in daylength and light intensity, daily specific growth rates () are thought to be higher in summer than in winter. Neither ammonium enrichment assays nor particulate carbon-to-nitrogen ratios provided convincing evidence for nitrogen limitation during summer, and the observed changes in diatom abundance cannot be explained on this basis. Both the high diatom concentrations and their seasonal variations probably are due mainly to factors other than growth rates; two factors considered important are diatom flotation and seasonal changes in wind-driven water transport. C. armatum usually dominates the phytoplankton biomass in the surf zone, and evidence suggests that this species is strongly dominant in terms of primary production.Contribution No. 1391 of the School of Oceanography, University of Washington, Seattle, Washington, USA     

Photosynthetic carbon reduction: high rates in the sea-surface microlayer

The ocean is a significant sink for increasing anthropogenic carbon dioxide emissions. The sea-surface microlayer (upper 50 m layer) serves as the primary point for exchange of materials, including carbon dioxide, between the atmosphere and hydrosphere. We determined microalgal standing stocks and activities in microlayer and subsurface water from areas with (S) and without (NS) visible natural surface slicks in Sequim Bay, Washington, USA, in July 1984. Enrichment ratios (microlayer concentration: subsurface bulkwater concentration) were: phytoneuston population abundance, 37 (NS) to 154 (S); total chlorophylls, 1.3 (NS) to 18 (S); particulate carbon fixation, 2 (NS) to 52 (S); and dissolved carbon excretion, 17 (NS) to 63 (S). Photoinhibition of 36 to 89% occurred in phytoneuston under natural summer light-intensities. Slick samples had greater standing stocks and rates of primary production than non-slick samples. The species composition of phytoneuston was distinctly different from that of phytoplankton. These results suggest that phytoneuston form a unique community. Although carbon fixation in summer was less per individual in phytoneuston than in phytoplankton, because of their overall abundance at the air-sea interface, phytoneuston may play an important role in the transfer of CO₂ from the atmosphere into ocean surface-water.     

Interactions of inorganic nitrogen in the uptake and assimilation by marine phytoplankton

The effect of ambient ammonium concentration on the nitrate uptake rate of marine phytoplankton was investigated. These studies consisted of laboratory experiments using unialgal species and field experiments using natural phytoplankton communities. In laboratory experiments, ammonium suppressed the uptake rates of nitrate and nitrite. Approximately 30 min were required for ammonium to exhibit its fully inhibitory effect on nitrate uptake. At high ammonium concentration (>3 g-at/l), a residual nitrate uptake rate of approximately 0.006 h^-1 was observed. When the ambient ammonium concentration was reduced to a value less than 1 g-at/l, the suppressed nitrate uptake rate subsequently attained a value comparable to that observed before the addition of ammonium. A range of 25 to 60% reduction in the nitrate uptake rate of natural phytoplankton communities was observed at ambient ammonium concentrations of 1.0 g-at/l. A mechanism is proposed for the suppression of nitrate uptake rate by ammonium through feedback control of the nitrate permease system and/or the nitrate reductase enzyme system. The feedback control is postulated to be regulated by the level of total amino acids in the cell.Contribution No. 936 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of a dissertation submitted to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

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     

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     

The dynamics of phytoplankton blooms in puget sound a fjord in the Northwestern United States

This paper describes a quantitative investigation of relationships between the growth of phytoplankton, and climatic and hydrodynamci conditions in temperate fjords with marked tides, as exemplified by Puget Sound, Washington (USA). Algal growth in the open waters of the central basin of the Sound is dominated by a number of intense blooms beginning in late April or May and recurring throughout the summer. Rarely, and only briefly, does nitrate become exhausted. The phytoplankton production rate in the central basin of Puget Sound is about 465 g C m^-2 year^-1. During the springs of 1966 and 1967, oceanographic measurements were carried out at a mid-channel station with sufficient frequency to allow investigation of physical and biological processes with time scales of the order of a day. The principal investigative tool is a numerical model in which the hydrodynamical conditions are represented by an approximate analysis of the gravitational convection mode of circulation. Algal concentration is represented as a continous function of space and time in the model which ascribes changes in phytoplankton density to variations in photosynthetic and respiratory activity, algal sinking, grazing by herbivores, and to mixing and advection. Computations adequately reproduce the principal features of phytoplankton concentrations observed during 75 days and 35 days in the springs of 1966 and 1967, respectively. Numerical experiments assess the relative importance of various processes which govern the level of primary production in Puget Sound. It is concluded that phytoplankton growth is limited by a combination of factors, including vertical advection and turbulence, modulation of underwater light intensity by self-shading and inorganic particulates, sinking of algal cells, and occasional rapid horizontal advection of the population from the area by sustained winds. The high primary productivity of the Sound is due to intensive upward transport of nitrate by the estuarine mechanism. These results should be generally applicable to other temperate fjords because of the largely conventional choice of the biological functions.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. X. Chemical composition of the surf diatom Chaetoceros armatum and its major herbivore,the Pacific razor clam Siliqua patula

The surf diatom Chaetoceros armatum T. West, collected from its natural habitat along the Washington coast (USA), had a large inorganic component in the form of a clay coat (consisting of clay minerals, illite and montmorillonite) surrounding the cell chains: 63% of the dry weight of the natural material collected in November was inorganic. The organic fraction was composed of 67.6% lipid, 29.7% protein, and 1.3% carbohydrate. Culture cells were likewise high in protein and lipid and low in carbohydrate. Traces of chitin found in field samples were probably a contaminant, since chitin was absent from cultured cells. This diatom species serves as a major food source throughout the year for the Pacific razor clam Siliqua patula Dixon, which inhabits these same beaches. Protein constituted 47% and lipid 42% of the dry weight of razor clam tissue. The fatty acid distribution in the diatom lipid resembled that previously reported for other diatom species; similarly, the fatty acid distribution of the clam lipid was similar to that previously reported for other bivalve molluscs. The clam fatty acids differed in chain length and degree of saturation from those extracted from its food source, indicating an active fatty acid metabolism in the clam.Contribution No. 995 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Primary productivity and sizes of pools of organic carbon in the mixed layer of the ocean

We estimated primary productivity and distributions of carbon in the phytoplankton, micro-zooplankton, and suspended and dissolved matter in various areas of the World Ocean to increase our information about the organic carbon cycle in the surface layer of the sea. Primary productivity ranged from about 0.1 gC m^–2 day^–1 in the Gulf of Mexico to 9 gC m^–2 day^–1 in nutrient-rich water off Peru. Phytoplankton carbon ranged from less than 10 g/l in the former to 750 g/l in the latter and in nutrient-rich water off southwest Africa. Micro-zooplankton carbon usually was less than 50 g/l in all waters, and was dominated by ciliates, copepodids, and copepod nauplii in all areas. Concentrations of particulate carbon ranged from 12 g/l off the east coast of South America to 850 g/l off southwest Africa. Concentrations of dissolved organic carbon varied between 0.5 and 1.5 mg/l in all areas except off Peru, where maximum values of 4.5 mg/l were observed. Turnover rates of carbon by small standing crops of micro-flagellates (1 to 5 longest dimension) and dinoflagellates in nutrient-poor waters were lower than those by large standing crops of diatoms and micro-flagellates in nutrient-rich waters. Concentrations of phytoplankton usually accounted for 20 to 55% and micro-zooplankton for 2 to 30% of the particulate carbon in the surface layer of the sea. Concentrations of dissolved organic carbon were not related to concentrations of particulate carbon in most waters except off Peru, where they appear to be directly related.     

Ammonia excretion by zooplankton and its significance to primary productivity during summer

Excretion rates of ammonia have been determined for zooplankton off the coasts of Washington and Oregon (USA). Rates varied from 0.16 to 0.60 g-at NH ₄ ⁺ -N/mg dry weight/day for most planktonic animals, and from 0.02 to 0.06 for jellyfishes. Ammonia concentration in seawater was low in offshore regions. Ammonia released by zooplankton was studied in relation to primary productivity during summer. It was found that, in the Columbia River plume offshore, excreted ammonia contributed about 90% of the total nitrogen requirements of observed production rates. The ammonia-N contribution was 36% in oceanic waters, and was relatively unimportant in the inshore region. The significance of eddy diffusivity in offshore waters and upwelling in inshore waters is also discussed.Contribution No. 747 from the Department of Oceanography, University of Washington, Seattle, USA.     

Dynamics of benthic vegetation standing-stock,irradiance, and water properties in central Puget Sound

The standing stock of benthic macroalgae, sediment-associated microalgae and eelgrass (Zostera marina L.) was sampled in conjunction with irradiance and water properties from June 1982 through March 1984 to examine the relationship between the dynamics of benthic primary producers and environmental factors in central Puget Sound, Washington, USA. Sediment-associated microalgal standing stock (measured by chlorophylla) peaked in April and August. The seaweed assemblage, dominated by bladed green algae (e.g.Ulva fenestrata) and eelgrass exhibited maximum standing stocks in August. Although water temperature correlated best with changes in standing stock of all vegetation types, solar irradiance appeared to trigger the onset of biomass buildup and autumn die-back by the plants. Seasonal variations in dissolved oxygen reflected the buildup and loss of plant standing-stock. Nutrient concentrations, with the exception of ammonia, exhibited seasonal trends. Most nutrients were in greatest concentration in winter and reached minimum concentration in late spring-summer. Regeneration of nutrients in autumn followed shortly after the autumn dieback of the benthic vegetation. We concluded that irradiance was the primary controlling factor in the system. Nutrient limitation (primarily nitrate) may control standing-stock accumulations from the period May–October when light is not limiting. In contrast to phytoplankton systems in deep portions of Puget Sound, shallow nearshore systems may be more susceptible to the effects of increased inorganic nutrient-loadings from anthropogenic sources.Contribution No. 808, School of Fisheries, University of Washington     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Physioecology of zooplankton. II. Effects of phytoplankton concentration,temperature, and body size on the development and molting rates of Calanus pacificus and Pseudocalanus sp.

Developmental time and stage duration for Calanus pacificus Brodsky and Pseudocalanus sp. and the rate of loss of body carbon by molting for C. pacificus were estimated for copepodite stages cultured under various combinations of phytoplankton concentration and temperature. Mean development time and stage duration for C. pacificus decreased hyperbolically with increasing food concentration, and the minimum time required for reaching a given stage decreased logarithmically with a logarithmic increase in temperature. Low temperature retarded the development of early stages proportionally more than that of late stages, and stage duration increased logarithmically with increasing body weight. Therefore, copepodite development was not isochronal. The rate of loss of body carbon by molting was small, ranging from 0.2 to 2% day^-1. This rate increased hyperbolically with food concentration and was linearly related to the growth rate. The critical food concentration for the rates of development and molting increased with temperature and stage of development, but these rates were less dependent on food concentration than the growth rate. The development rate of Pseudocalanus sp. was higher than that of C. pacificus, and was less influenced by changes in food concentration and temperature. It is postulated that the inverse relationship between temperature and body size results from a differential effect of temperature and body size on the rates of growth and development. That is, with increasing body size the growth rate tends to become temperature-independent, but the development rate remains proportional to temperature. Thus, copepodites growing at low temperature can experience a greater weight increment between molting periods than individuals growing at high temperature, because the growth rate is similar at all temperatures but stage duration is longer at low temperature.Contribution No. 1128 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Impact of curve construction and community dynamics on the species-time relationship

The species-time relationship (STR) describes how the species richness of a community increases with the time span over which the community is observed. This temporal scaling provides insight into theoretical questions on species diversity patterns as well as applied questions on the appropriate time scale for biodiversity assessments. To better understand STRs, we discuss the methods used to construct STRs in the literature and derive the impact of curve construction on STR properties. Using vegetation data from Mount St. Helens, Washington, USA, we illustrate the sensitivity of the STR to construction under colonization-dominated dynamics. This study highlights the importance of considering the type of STR when interpreting, comparing, and applying STRs, particularly in disturbed or successional systems.