Production,composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as g C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d^-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.     

Selective particle ingestion by oyster larvae (Crassostrea virginica) feeding on natural seston and cultured algae

I investigated selective particle ingestion by oyster larvae (Crassostrea virginica) feeding on natural seston from Chesapeake Bay and laboratory-cultured algae of different sizes or chemical content. In 15 of 16 experiments with complex natural suspensions as food, small(<150 m) and large (>150 m) larvae selected most strongly for small (2 to 4 m) food particles, but in the presence of a large (>10 m)-cell dinoflagellate bloom, large larvae strongly selected much larger (22 to 30 m) food material (presumably dinoflagellates). When fed simplified mixtures of four cultured algal species (Synechococcus bacillaris, Isochrysis sp., Dunaliella tertiolecta, and Prorocentrum minimum) ranging in size from 1 to 11 m, small larvae preferred 1 m algae while large larvae preferred 11 m algae. In experiments with algal mixtures, and with suspensions of natural particles and added algae, large larvae preferred algal species harvested from exponential-phase cultures over other species from stationary-phase cultures. Larval ingestion rates of the cultured alga Thalassiosira pseudonana were about three times higher for cells with a low carbon:nitrogen ratio (7.2:1) than for high C:N ratio (16.2:1) cells when these cells were offered separately in suspensions of equal concentration. As a result, more algal cells, algal C, and algal N was ingested by larvae fed low C:N cells. However, larvae did not show a significant preference for either type of cell when they were offered in a 1:1 cell mixture. Feeding patterns of C. virginica larvae in natural food suspensions can vary with the composition of these complex suspensions, and ingestion seems dependent not only on the size, but on the growth rate and chemical quality of food particles.     

Sperm morphology, swimming velocity, and longevity in the house sparrow Passer domesticus

Sperm competition exerts strong selection on males to produce spermatozoa with an optimal morphology that maximizes their fertilization success. Long sperm were first suggested to be favored because they should swim faster. However, studies that investigated the relationship between sperm length and sperm competitive ability or sperm swimming velocity yielded contradictory results. More recently, ratios of the different sections of a spermatozoon (the head, midpiece, and flagellum) were suggested to be more crucial in determining swimming velocity. Additionally, sperm ability to remain and survive in the female storage organs may also influence fertilization success, so that optimal sperm morphology may rather maximize sperm longevity than velocity. In this study, we investigated how sperm morphology is related to sperm velocity and sperm longevity in the house sparrow Passer domesticus. Sperm velocity was found to be correlated with head/flagellum ratio. Sperm with small heads relative to their flagellum showed higher swimming velocity. Additionally, shorter sperm were found to live longer. Finally, we found sperm morphological traits to vary substantially within males and the head/flagellum ratio to be unrelated to total sperm length. We discuss the hypothesis that the substantial within-male variation in sperm morphology reflects a male strategy to produce a diversity of sperm from long, fast-swimming to short, long-living sperm to maximize their fertilization success in a context of sperm competition.     

Oxygen consumption in the mullet Liza macrolepis with special reference to swimming velocity

Oxygen consumption of tagged (plastic opercular tag) and untagged mullet Liza macrolepis (Smith), forced to swim up to 22 cm/sec (fish size: 10 cm), increased proportionately to increase in swimming velocity above 5 cm/sec. The tag did not appear to cause any marked metabolic disturbance. The mean routine metabolic rate and the rate at 5 cm/sec were higher than the rates obtained for several higher levels of forced activity, possibly due to excitement and inefficient swimming at lower swimming speeds, as suggested by earlier workers. The interpretation of the elevation of a line drawn through the mean rates of metabolism at various swimming speeds, in relation to the standard metabolic rate, may be of value in quantitative expression of excitability of individual species.     

Phytoplankton sinking rates in oligotrophic waters off Hawaii,USA

The sinking rates in two size fractions of natural phytoplankton were measured over much of the photic zone in a subtropical environment. At 24, 40 and 71 m, sinking rates (± SD) of the 3 to 20 m fractions were 0.72±0.05, 0.83±0.05, and 0.34±0.04 m · d^-1, respectively, and rates for the 20 to 102 m fraction were 1.50±0.21, 1.65±0.14, and 0.95±0.22 m · d^-1, respectively. At all depths sampled, the 20 to 102 m size fraction was observed to sink significantly (P<0.01) faster than the 3 to 20 m fraction. considering vertical variability, both size fractions were observed to have significantly (P<0.01) lower sinking rates at 71 m than at more shallow depths. The finding of lower sinking rates in the region which lies just above the subsurface chlorophyll maximum provides empirical support for the hypothesis that variations in phytoplankton buoyancy may be related to the maintenance of this feature which is typical in oceanic environments. Analysis of sinking rate traces describes the distributions of specific sinking velocities for each size fraction and their variation.Oceanic Institute Contribution No. 162     

Effects of copper and cadmium on growth,swimming and predator avoidance in Eurytemora affinis (Copepoda)

In this study we demonstrate the sensitivity of swimming behavior and predator-escape responses of nauplii of the estuarine copepod Eurytemora affinis to sublethal doses of Cu and Cd. Behavior was generally altered at metal doses below those affecting growth rates or survival of the copepods. Swimming velocities of Cu-dosed nauplii were different from controls at all concentrations of Cu tested (10–50 g l^-1 total Cu) after 24- to 48-h exposure, whereas development rate of nauplii was significantly reduced only after 96 h at 25 g l^-1. The 96 h LC50 for Cu was approximately 30 g l^-1 Cu. Naupliar swimming velocity was also affected by Cd. Swimming speeds were reduced after 24 h at 130 g l^-1, and development was slowed after 48 h at 116 g Cd l^-1. The 96-h LC50 was >120 g l^-1. Little is known of the adaptive role of specific motile behaviors in the success of larval copepods. We investigated the relationship of swimming speed to predator — prey interactions of the nauplii using both real and simulated predators. Nauplii exposed to Cu for 24 h were observed to be generally hyperactive, a condition which could increase their encounter frequency with predators. Reduced numbers of escape responses of nauplii to a simulated predator, another indication of increased vulnerability to predation, were observed only after 48-h exposure to Cu. Nevertheless, feeding rates of non-dosed larval striped bass on dosed nauplii (24 h at 25 g Cu l^-1) were significantly higher than on control nauplii. Feeding rates of larval mysid shrimp, however, were not higher on similarly dosed nauplii; 24 h exposure of nauplii to >30 g Cu l^-1 did result in increased predation by mysids.Contribution No. 272 of the US EPA Environmental Research Laboratory, Narragansett, Rhode Island 02882, USA     

Species-specific grazing rates of heterotrophic dinoflagellates in oceanic waters,measured with a dual-label radioisotope technique

A dual-isotope method was developed to measure grazing rates and food preferences of individual species of heterotrophic dinoflagellates from natural populations, collected from the Slope, Gulf Stream, and Sargasso Sea and from a transect from Iceland to New England, in 1983. The isotope method measures the grazing rates of microzooplankton which cannot be separated in natural populations on the basis of size. Tritiated-thymidine and ¹⁴C-bicarbonate were used to label natural heterotrophic and autotrophic food, respectively. Nine oceanic dinoflagellate species in the genera Protoperidinium, Podolampas, and Diplopsalis fed on both heterotrophic and autotrophic food particles with clearance rates of 0.4 to 8.0 l cell^-1 h^-1, based on ³H incorporation, and 0.0 to 28.3 l cell^-1 h^-1, based on ¹⁴C incorporation. Two dinoflagellate species, Protoperidinium ovatum and Podolampas palmipes, fed only on ³H-labelled food particles. Several species of dinoflagellates fed on bacteria (<1 m) which had been prelabelled with ³H-thymidine. The clearance rates of heterotrophic dinoflagellates and ciliates were similar and within the range of tintinnid ciliate clearance rates reported in the literature. As heterotrophic dinoflagellates and ciliates can have comparable abundances in oceanic waters, we conclude that heterotrophic dinoflagellates may have an equally important impact as microheterotrophic grazers of phytoplankton and bacteria in oceanic waters.Partially supported by a grant from the National Science Foundation, OCE-81-17744     

Phosphate acquisition in the giant clam-zooxanthellae symbiosis

The effect of phosphate on the giant clam Tridacna gigas and on its symbiotic dinoflagellate Symbiodinium sp. was compared with that on cultured Symbiodinium sp. originally isolated from the same clarn species. Incubation of whole clams in elevated phosphate (10 M) reduced their capacity for phosphate uptake, but the uptake capacity of the clam's zooxanthellae population was not influenced. In addition, there was no change in the zooxanthellae density and the N:P ratio, of these algae., On the other hand, cultured zooxanthellae were influenced by the phosphate regimen of their culture medium. Compared with controls (0 M P), addition of 10 M phosphate to the culture medium caused an increase of 100% in cell density and decreases of 50% in the N:P ratio, and 80% in the phosphate-uptake capacity of the zooxanthellae. Zooxanthellae freshly isolated from the clams exhibited properties similar to those of zooxanthellae cultured in the absence of phosphate. These results demonstrate that the zooxanthellae population of T. gigas have limited access to the inorganic phosphate in sea water and the phosphate reserves within the animal host.     

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

The effect of low concentrations of phenol and ammonia on egg production rates,fecal pellet production and egg viability of the calanoid copepod Acartia clausi

Estimates of daily fecundity, hatching success and fecal pellet production are reported for Acartia clausi females exposed for 10 d to low levels of phenol and ammonia. Copepods were collected in 1991 and 1992 from a southern coastal area of the Mar Grande of Taranto (southern Italy). A reduction in egg numbers and fecal pellet production was observed for females after 8 d of exposure to 500 g l^-1 phenol concentration. Ammonia (120 g l^-1) produced a significant increment in egg production, but hatching success was reduced by about 50% after nine exposure days. A. clausi was more sensitive to ammonia than phenol at high concentrations (24-h LC₅₀ phenol-32.26 mg l^-1; 24-h LC₅₀ ammonia=0.91 mg l^-1). At low concentrations, only long-term exposure to phenol determined a reduction in fecundity.     

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     

Zooplankton standing stock and community size structure within the epipelagic zone: a comparison between the central Red Sea and the Gulf of Aden

Standing stock and size composition of the zooplankton comunity (>100 m) were studied in four depth strata of the upper 200 m of the water column during a Meteor cruise to the central Red Sea and Gulf of Aden in spring 1987. The central Red Sea was divided into a northern area of higher salinity and a less saline southern part. Both areas exhibited significant differences in zooplankton abundance and standing stock. The latter increased by the ratio 1:2:3 from the northern central Red Sea to its southern part and further south to the Gulf of Aden. For size structure analysis samples were fractionated into three size classes (100 to 300, 300 to 500, 500 to 5000 m). In the central Red Sea the smallest size was dominant whereas in the Gulf of Aden the largest size fraction played a greater relative role than in the central Red Sea. This shift in size structure of the zooplankton community from the Red Sea to the Gulf of Aden is apparently primarily related to ecosystem difference between both areas, leading to a change in species composition. In addition, size reduction of individual species common to both seas may be of some significance in the extreme environment of the Red Sea.     

Dinoflagellate luminescence increases susceptibility of zooplankton to teleost predation

The burglar alarm theory of bioluminescence was investigated by determining predation rates of a nocturnal teleost predator,Porichthys notatus, on nonluminescent kelp mysids illuminated by dinoflagellate flashes, between the fall and spring of 1989/1990. Mysids (Holmesimysis costata) were placed in aquaria containing varying concentrations (0 to 40 cells/ml) of the dinoflagellatePyrocystis fusiformis and a single midshipman fish. Controls usedP. fusiformis during their luminescence-inhibited day phase. Mysid swimming movements readily stimulated dinoflagellate luminescence. Flashes and prey strikes were observed simultaneously by image-intensifying and infrared video cameras on a splitscreen monitor. Predation rates increased at dinoflagellate concentrations of 3 to 15 cells/ml and decreased below controls at levels>20 cells/ml. Videotape analysis showed that at low concentrations (2 to 5 cells/ml), strike success rates exceeded 75% if prey were previously illuminated by a flash, but dropped below 50% at higher cell densities. Increased predation was attributed to luminescence revealing prey position. The decrease at higher concentrations was considered to be due to greater flash frequency providing a more diffuse and confusing target. The study demonstrates the effects of secondary luminescence on zooplankton predation at normally encountered dinoflagellate concentrations.     

Effects of ovarian fluid on sperm velocity in Arctic charr (Salvelinus alpinus)

Numbers of studies in externally fertilizing fish species provide evidence for an effect of ovarian fluid on sperm motility characteristics such as duration of forward mobility, velocity or percent motile sperm cells. Yet, because of variations among females in the quality of their ovarian fluid, such effects might differ between individuals. Additionally, ovarian fluid from different females could also be expected to affect each ejaculate differently, resulting in cryptic female choice. In this study on Artic charr (Salvelnius alpinus), sperm velocity from several males was measured in the diluted ovarian fluid of several females according to a fully balanced crossing design. This design allowed us to estimate variations among females in the effect of their ovarian fluid on the velocity of sperm from different males, and to detect variations among males in the ability of their sperm to swim in ovarian fluid. Sperm velocity was estimated by computer-assisted sperm analysis. Average velocity was found to vary among females, with some females having constantly higher velocity measurements in their ovarian fluid, and among males, indicating that some males had overall faster sperm in ovarian fluid than others. Moreover, variation in sperm velocity was shown to depend on individual female-male interactions. Our results document that females vary in the effect of their ovarian fluid on sperm velocity and that their ovarian fluid may stimulate sperm velocity according to individual characteristics of males. This latter result suggests a potential mechanism for cryptic female choice.

Feeding,growth and bioluminescence of the heterotrophic dinoflagellate Protoperidinium huberi

Feeding, growth and bioluminescence of the thecate heterotrophic dinoflagellate Protoperidinium huberi were measured as a function of food concentration for laboratory cultures grown on the diatom Ditylum brightwellii. Ingestion of food increased with food concentration. Maximum ingestion rates were measured at food concentrations of 600 g C l^-1 and were 0.7 g C individual^-1 h^-1 (1.8 D. brightwelli cells individual^-1 h^-1). Clearance rates decreased asymptotically with increasing food concentration. Maximum clearance rates at low food concentration were ca. 23 l ind^-1 h^-1, which corresponds to a volume-specific clearance rate of 5.9x10⁵ h^-1. Cell size of P huberi was highly variable, with a mean diameter of 42 m, but no clear relationship between cell size and food concentration was evident. Specific growth rates increased with food concentration until maximum growth rates of 0.7 d^-1 were reached at a food concentration of 400 g C l^-1 (1000 cells ml^-1). Food concentrations as low as 10 g C l^-1 of D. brightwellii (25 cells ml^-1) were able to support growth of P. huberi. The bioluminescence of P. huberi varied with its nutritional condition and growth rate. Cells held without food lost their bioluminescence capacity in a matter of days. P. huberi raised at different food concentrations showed increased bioluminescence capacity, up to food concentration that supported maximum growth rates. The bioluminescence of P. huberi varied over a diel cycle, and these rhythmic changes persisted during 48 h of continuous darkness, indicating that the rhythm was under endogenous control.     

Sloughed mucus and drag-reduction in a school of Atlantic silversides,Menidia menidia

Atlantic silversides, Menidia menidia, were collected by seine-net from the Newport River estuary, North Carolina, USA and maintained in the laboratory. Direct measurement showed that the amount of mucus sloughed off individual silversides was 567 g h^-1 at a swimming speed of 50 cm s^-1. Individual M. menidia were also allowed to swim in a flowtank filled with 0, 1, and 10 ppm of the synthetic drag-reducer Polyox to determine if any changes in tailbeat frequency or amplitude could be correlated with the level of this sloughed material. Tailbeat measurements between seawater controls and either Polyox concentration did not differ significantly. Calculations based on this evidence indicate that the amount of solubilized mucus sloughed off a school of 10 000 M. menidia would be at least two orders of magnitude less than the highest concentration of Polyox tested. We conclude that solubilized mucus in the water column does not confer drag-reduction to schooling M. menidia.     

An approach for particle sinking velocity measurements in the 3–400?μm size range and considerations on the effect of temperature on sinking rates

The flux of organic particles below the mixed layer is one major pathway of carbon from the surface into the deep ocean. The magnitude of this export flux depends on two major processes—remineralization rates and sinking velocities. Here, we present an efficient method to measure sinking velocities of particles in the size range from approximately 3–400?μm by means of video microscopy (FlowCAM^?). The method allows rapid measurement and automated analysis of mixed samples and was tested with polystyrene beads, different phytoplankton species, and sediment trap material. Sinking velocities of polystyrene beads were close to theoretical values calculated from Stokes’ Law. Sinking velocities of the investigated phytoplankton species were in reasonable agreement with published literature values and sinking velocities of material collected in sediment trap increased with particle size. Temperature had a strong effect on sinking velocities due to its influence on seawater viscosity and density. An increase in 9?°C led to a measured increase in sinking velocities of ~40?%. According to this temperature effect, an average temperature increase in 2?°C as projected for the sea surface by the end of this century could increase sinking velocities by about 6?% which might have feedbacks on carbon export into the deep ocean.     

Particle-size selection by Pseudopolydora paucibranchiata (Polychaeta: Spionidae) in suspension feeding and in deposit feeding: influences of ontogeny and flow speed

Pseudopolydora paucibranchiata Okuda suspension feeds and deposit feeds at the sediment-water interface, where it is exposed to a variety of particles differing in physical characteristics and nutritional value. In flume experiments (conducted in August 1994 and May 1995) with two sizes of either suspended or deposited beads, I measured particle-size selection separately in each feeding mode. The same influences of palp width and of ambient flow speed were observed in each mode. At velocities 0.74 cm s^-1 there were no relationships between palp width and the proportion of gut contents composed of large beads. At velocities 1.8 cm s^-1 worms with narrower palps ingested relatively fewer large beads (and more small beads) than did worms with wider palps. Palp width and body length were linearly related, and results were similar when analyzed with body length as the independent variable. As flow speed increased, selectivity changed in a worm-size-specific manner: worms with a palp width -1. Assuming that in the field (1) particle size is the principle criterion for selection, and (2) the amount of digestible food component in deposited and suspended particles, respectively, is related to particle surface area and volume, I hypothesize that changes in selectivity as velocity rises can cause juveniles to experience a decreasing profitability of suspension feeding and a simultaneously increasing profitability of deposit feeding. Juveniles could maintain a diet of high food value despite flow variations by adjusting the proportion of time they spend suspension feeding relative to deposit feeding.     

Effect of inorganic phosphorus on the growth rate of diatoms

The effect is studied of different inorganic phosphorus concentrations on changes in growth rates of 7 diatom species isolated from the plankton of the Black Sea. All species examined increase their cell-division rate with increasing phosphorus concentration in the medium. The phosphorus concentrations have been determined above which division rate is not limited by phosphorus content in the medium. The non-limiting concentration amounts to V/V _m =0.9. In the species studied, non-limiting concentrations ranged from 1 to 30 g P/1. The highest values were obtained for relatively large-sized species. The ratio of cell surface to volume tended to be inversely related to the growth-rate-limiting concentration. Minimal values of phosphorus content in cells have been calculated to range from 0.6 to 4.0·10^–17 g.at–P/^–3. Based on a comparison of phosphate levels in the Black Sea with experimentally-derived rate-limiting concentrations, it is concluded that phosphorus does not limit the reproduction rate of phytoplankton in the more productive regions of the Black Sea or in the lower strata of the euphotic zone.     

Brood size and chemical composition of Pareuchaeta norvegica (Crustacea: Copepoda) in Loch Etive,Scotland

Brood size and chemical composition (carbon, nitrogen, and ash) of Pareuchaeta norvegica in Loch Etive, Scotland have been determined. From 23 to 37 eggs were found in each egg mass (mean=30 eggs). The eggs are oval; the mean size of the longer axis is 438 m. Variation in egg mass (as dry weight) is related to increase in dry weight of the maternal body. an increase of 1 mg in maternal body dry weight produces an increase of 0.03 mg dry weight in the egg mass. Carbon content increases with increasing body dry weight both in pre-spawning and spent females, but nitrogen content decreases with increasing body weight. Thus, the carbon:nitrogen ratio increases with increasing body dry weight. The carbon content of an egg mass ranges from 61.4 to 65.3%, and nitrogen content from 9.6 to 13.2% of the total dry weight. Some examples of partitioning of chemical components of the maternal body into the egg mass are also described.