The tunicate Salpa thompsoni ecology in the Southern Ocean. I. Distribution, biomass, demography and feeding ecophysiology

Distribution, density, and feeding dynamics of the pelagic tunicate Salpa thompsoni have been investigated during the expedition ANTARKTIS XVIII/5b to the Eastern Bellingshausen Sea on board RV Polarstern in April 2001. This expedition was the German contribution to the field campaign of the Southern Ocean Global Ocean Ecosystems Dynamics Study (SO-GLOBEC). Salps were found at 31% of all RMT-8 and Bongo stations. Their densities in the RMT-8 samples were low and did not exceed 4.8 ind m⁻² and 7.4 mg C m⁻². However, maximum salp densities sampled with the Bongo net reached 56 ind m⁻² and 341 mg C m⁻². A bimodal salp length frequency distribution was recorded over the shelf, and suggested two recent budding events. This was also confirmed by the developmental stage composition of solitary forms. Ingestion rates of aggregate forms increased from 2.8 to 13.9 μg (pig) ind⁻¹ day⁻¹ or from 0.25 to 2.38 mg C ind⁻¹ day⁻¹ in salps from 10 to 40 mm oral-atrial length, accounting for 25–75% of body carbon per day. Faecal pellet production rates were on average 0.08 pellet ind⁻¹ h⁻¹ with a pronounced diel pattern. Daily individual egestion rates in 13 and 30 mm aggregates ranged from 0.6 to 4.8 μg (pig) day⁻¹ or from 164 to 239 μg C day⁻¹. Assimilation efficiency ranged from 73 to 90% and from 65 to 76% in 13 and 30 mm aggregates, respectively. S. thompsoni exhibited similar ingestion and egestion rates previously estimated for low Antarctic (~50°S) habitats. It has been suggested that the salp population was able to develop in the Eastern Bellingshausen Sea due to an intrusion into the area of the warm Upper Circumpolar Deep Water     

Silica uptake kinetics of Halichondria panicea in Kiel Bight

Besides diatoms Demospongiae are the most important consumers of dissolved silica in the sea. They can play an important role for the silica budget especially in the shallow water areas of the Baltic Sea. The dependence of the silica uptake rate on the silica concentration of the seawater was measured for the sponge Halichondria panicea (Pallas, 1766). The sponges were collected in Kiel Bight. The uptake conformed to Michaelis–Menten kinetics with a half-saturation constant of 46.41 μM and a saturated uptake rate of 19.33 μmol h⁻¹ g⁻¹ ( p < 0.01). In the red algae zone of Kiel Bight the sponges depend on silica supply from the surrounding waters and may be silica-limited rather than food-limited in growth. Because of the much faster uptake of silica by diatoms and their lower saturation point, as well as the difference in spatial distribution of the two main silica consumers, a competition for silica between sponges and diatoms seems unlikely. Received: 21 June 1997 / Accepted: 15 July 1997     

Effects of mussel (Perna canaliculus) biodeposit decomposition on benthic respiration and nutrient fluxes

Suspension-feeding bivalves increase the quantity and quality of sedimenting organic matter through the production of faeces and pseudofaeces that are remineralised in coastal sediments and thus increase sediment oxygen demand and nutrient regeneration. Bivalves are intensively cultivated worldwide; however, no bivalve biodeposit decay rates are available to parameterise models describing the environmental effects of bivalve culture. We examined sediment biogeochemical changes as bivalve biodeposits age by incubating coastal sediments to which we added fresh mussel (Perna canaliculus) biodeposits and measured O₂ and nutrient fluxes as well as sediment characteristics over an 11-day period. Biodeposits elevated organic matter, chlorophyll a, phaeophytin a, organic carbon and nitrogen concentrations in the surface sediments. Sediment oxygen consumption (SOC) increased significantly (P=0.016) by ∼1.5 times to 1,010 μmol m⁻² h⁻¹ immediately after biodeposit addition and remained elevated compared to control cores without additions for the incubation period. This increase is in the range of observed in situ oxygen demand enhancements under mussel farms. To calculate a decay rate for biodeposits in sediments we fitted a first-order G model to the observed increase in SOC. The significant model fit (P=0.001, r ²=0.72) generated a decay rate of 0.16 day⁻¹ (P=0.033, SE=0.05) that corresponds to a half-life time of 4.3 day. This decay rate is 1–2 orders of magnitude higher than published decay rates of coastal sediments without organic enrichment but similar to rates of decaying zooplankton faecal pellets. NH₄⁺ release increased rapidly on the day of biodeposit addition (P=0.013) and reached a maximum of 144 μmol m⁻² h⁻¹ after 5 days which was 3.6 times higher compared to control cores. During this period NH₄⁺ release was significantly (P<0.001 to P=0.043) higher in the cores with biodeposit additions than in control cores.     

Phytoplankton growth and microzooplankton grazing rates in a restricted Mediterranean lagoon (Bizerte Lagoon,Tunisia)

Phytoplankton growth and microzooplankton grazing were investigated in the restricted Bizerte Lagoon in 2002 and 2004. The 2002 study, carried out at one station from January to October, showed significant seasonal variations in phytoplankton dynamics. High growth rates (0.9–1.04 day⁻¹), chlorophyll a (Chl a) concentrations (6.6–6.8 μg l⁻¹) and carbon biomass (392–398 μg C l⁻¹) were recorded in summer (July), when several chain-forming diatoms had intensively proliferated and dominated the carbon biomass (74%). In 2004, four stations were studied during July, a period also characterized by the high proliferation of several diatoms that made up 70% of the algal carbon biomass. In 2004, growth rates (0.34–0.45 day⁻¹) and biomass of algae (2.9–5.4 μg Chl a l⁻¹ and 209–260 μg C l⁻¹) were low, which may be related to the lower nutrient concentrations recorded in 2004. Microzooplankton >5 μm were mainly composed of heterotrophic dinoflagellates and ciliates. Microzooplankton biomass peaked during summer (2002 320–329, 2004 246–361 μg C l⁻¹), in response to the enhanced phytoplankton biomass and production. The grazer biomass was dominated by ciliates (71–76%) in July 2002 and by heterotrophic dinoflagellates (52–67%) in July 2004. Throughout the year and at different stations, microzooplankton grazed actively on phytoplankton, removing 26–58% of the Chl a and 57-84% of the primary production. In 2002, the highest grazing impact was observed on the large algae (>10 μm) during the period of diatom dominance. These results have a significant implication for carbon export to depth. Indeed, the recycling of most of the diatom production by the microbial food web in the upper water column would reduce the flux of material to the seafloor. This should be considered when modeling the carbon cycling in coastal environments and under conditions of diatom dominance. During both studies, ciliates had higher growth rates (0.5–1.5 day⁻¹) and a higher carbon demand (165–470 μg C l⁻¹ day⁻¹) than dinoflagellates (0.1–0.5 day⁻¹, 33–290 μg C l⁻¹ day⁻¹). Moreover, when grazer biomass was dominated by ciliates (in July 2002), herbivory accounted for 71–80% of the C ingested by microzooplankton while it accounted only for 14–23% when dinoflagellates dominated the grazer biomass (in July 2004). These results suggest that, in contrast to findings from open coastal waters, ciliate species of the restricted Bizerte Lagoon were more vigorous grazers of the large algae (diatoms) than were dinoflagellates.     

Exposure of the urban population to mercury in Changchun city, Northeast China

An assessment of exposure to mercury in Changchun city has been undertaken. We estimated Hg exposure to members of the general population based on currently available information and our research. We also studied the Hg concentrations in scalp hair of adults. Adults have an estimated intake of all Hg species via all routes of 6.780 μg day⁻¹ (excluding dental amalgam), which equates to an absorbed dose of 1.718 μg day⁻¹. Fish consumption was the most important exposure route (12% of intake, 43% of absorbed dose). Furthermore, air, cereals and vegetables were important exposure routes, and these exposure were estimated for absorbed dosed at 0.296, 0.209 and 0.318 μg day⁻¹, respectively. The mean Hg concentration in hair was 0.448 μg g⁻¹ (range 0.092–10.463 μg g⁻¹). Hg concentration in the hair of males was 0.422 μg g⁻¹ (0.105–2.665 μg g⁻¹), and was 0.474 μg g⁻¹(0.092–10.463 μg g⁻1) in the hair of females. Neither place of residence nor age had any significant effect on hair Hg concentrations.     

Irradiance and temperature regulation of oxygenic photosynthesis and O2 consumption in a hypersaline cyanobacterial mat (Solar Lake, Egypt)

 Short-term effects of temperature and irradiance on oxygenic photosynthesis and O₂ consumption in a hypersaline cyanobacterial mat were investigated with O₂ microsensors in a laboratory. The effect of temperature on O₂ fluxes across the mat–water interface was studied in the dark and at a saturating high surface irradiance (2162 μmol photons m⁻² s⁻¹) in the temperature range from 15 to 45 °C. Areal rates of dark O₂ consumption increased almost linearly with temperature. The apparent activation energy of 18 kJ mol⁻¹ and the corresponding Q ₁₀ value (25 to 35 °C) of 1.3 indicated a relative low temperature dependence of dark O₂ consumption due to mass transfer limitations imposed by the diffusive boundary layer at all temperatures. Areal rates of net photosynthesis increased with temperature up to 40 °C and exhibited a Q ₁₀ value (20 to 30 °C) of 2.8. Both O₂ dynamics and rates of gross photosynthesis at the mat surface increased with temperature up to 40 °C, with the most pronounced increase of gross photosynthesis at the mat surface between 25 and 35 °C (Q ₁₀ of 3.1). In another mat sample, measurements at increasing surface irradiances (0 to 2319 μmol photons m⁻² s⁻¹) were performed at 25, 33 (the in situ temperature) and 40 °C. At all temperatures, areal rates of gross photosynthesis saturated with no significant reduction due to photoinhibition at high irradiances. The initial slope and the onset of saturation (E _k = 148 to 185 μmol photons m⁻² s⁻¹) estimated from P versus E _d curves showed no clear trend with temperature, while maximal photosynthesis increased with temperature. Gross photosynthesis was stimulated by temperature at each irradiance except at the lowest irradiance of 54 μmol photons m⁻² s⁻¹, where oxygenic gross photosynthesis and also the thickness of the photic zone was significantly reduced at 40 °C. The compensation irradiance increased with temperature, from 32 μmol photons m⁻² s⁻¹ at 25 °C to 77 μmol photons m⁻² s⁻¹ at 40 °C, due to increased rates of O₂ consumption relative to gross photosynthesis. Areal rates of O₂ consumption in the illuminated mat were higher than dark O₂ consumption at corresponding temperatures, due to an increasing O₂ consumption in the photic zone with increasing irradiance. Both light and temperature enhanced the internal O₂ cycling within hypersaline cyanobacterial mats. Received: 30 November 1999 / Accepted: 11 April 2000     

Feeding of Clausocalanids (Calanoida,Copepoda) on naturally occurring particles in the northern Gulf of Aqaba (Red Sea)

A total of 12 feeding experiments were conducted in the northern Gulf of Aqaba during spring (March/April) and autumn (September/October) 2002 at the Marine Science Station (MSS) in Aqaba. Females of three species of clausocalanids were selected: Clausocalanus farrani, C. furcatus and Ctenocalanus vanus. Natural occurring particle (NOP) larger than 5 μm were investigated as food source. The ambient chlorophyll a concentration at sampling depth (∼70 m) ranged between 0.15 and 1.00 μg chl a l⁻¹ and NOP concentrations ranged between 1.78 and 14.0 × 10³ cells l⁻¹ during the sampling periods. The division of particles into five size classes (5–10, 10–20, 20–50, 50–100 and >100 μm) revealed that most of the particles were found in the size classes below 50 μm (81–98%), while most of the natural occurring carbon (NOC) was concentrated in the size classes larger than 20 μm (70–95%). Ingestion rates were food density dependent rather than size dependent ranging between 0.02 and 1.65 × 10³ NOP ind⁻¹ day⁻¹ and 0.01 and 0.41 μg NOC ind⁻¹ day⁻¹, respectively, equivalent to a body carbon (BC) uptake between 0.4 and 51.8% BC day⁻¹. The share of the size classes to the total ingestion resembled in most cases the size class composition of the natural particle community.     

In situ measurements of photosynthetic irradiance responses of two Red Sea sponges growing under dim light conditions

Photosynthetic responses to irradiance by the photosymbionts of the two Red Sea sponges Theonella swinhoei (Gray) and Clionavastifica (Hancock) growing under dim light conditions were measured in situ (in September 1997) using a newly developed underwater pulse amplitude modulated (PAM) fluorometer. Relative rates of photosynthetic electron transport (ETR) were calculated as the effective quantum yield of photosystem II (Y ) multiplied with the photosynthetic photon flux (PPF). Photosynthesis versus irradiance (P-I ) curves, obtained within minutes, showed that individual specimens of both sponges, growing under very low light conditions, feature lower light saturation points as well as lower maximal ETRs than individuals growing under higher light. Evaluations of such curves using low irradiances of the actinic light source (20 to 130 μmol photons m⁻² s⁻¹) showed a general decrease in Y, with a shoulder from the lowest irradiance applied till 20 to 30 μmol photons m⁻² s⁻¹. Point measurements yielded ETRs close to what could be estimated from the P-I curves. These point measurements also revealed good correlations between the diurnally changing ambient irradiances (1 to 50 μmol photons m⁻² s⁻¹) and average ETR values for both species. Further analysis showed that although Y values varied considerably between the different point measurements, they did not decrease significantly with light under these very low irradiances. Therefore, PPF rather than Y seems to determine the in situ diel photosynthetic performance at the low ambient irradiances experienced by these sponges. Received: 22 November 1997 / Accepted: 8 April 1998     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

Total mercury and methylmercury concentrations in fish from the Mojana region of Colombia

Total mercury (T-Hg) and methylmercury (MeHg) concentrations have been measured in the muscle tissue of 16 fish species consumed in the Mojana region of Colombia. T-Hg analysis was performed by cold-vapor atomic-absorption spectroscopy (CV-ASS) and MeHg analysis by gas chromatography with electron-capture detection. Higher T-Hg and MeHg concentrations were detected in carnivorous species (T-Hg = 0.371 ± 0.172 (μg g⁻¹ fresh wt, MeHg = 0.346 ± 0.171 μg g⁻¹ fresh wt) than in non-carnivorous fish (T-Hg = 0.155 ± 0.108 μg g⁻¹ fresh wt, MeHg = 0.146 ± 0.102 μg g⁻¹ fresh wt). In the different species mercury was present almost completely as the methylated form, with percentages between 80.5 and 98.1% (mean 92.0 ± 3.4%). In 13.5% of fish-tissue samples T-Hg concentrations exceeded the maximum level recommended by the World Health Organization for human consumption (Hg = 0.5 μg g⁻¹ fresh wt). Although mean T-Hg concentrations in all fish samples (0.269 ± 0.181 μg g⁻¹ fresh wt) did not exceed this limit, risk assessment suggested that the consumption of 0.12 kg fish day⁻¹ could increase the risk of mercury poisoning of the inhabitants of this region.     

How size relates to oxygen consumption,ammonia excretion,and ingestion rates in cold (<Emphasis Type="Italic">Enteroctopus megalocyathus</Emphasis>) and tropical (<Emphasis Type="Italic">Octopus maya</Emphasis>) octopus species

The scaling of metabolic rates with body mass is one of the best known and most studied characteristics of aquatic animals. Herein, we studied how size is related to oxygen consumption, ammonia excretion, and ingestion rates in tropical (Octopus maya) and cold-water (Enteroctopus megalocyathus) cephalopod species in an attempt to understand how size affects their metabolism. We also looked at how cephalopod metabolisms are modulated by temperature by constructing the relationship between metabolism and temperature for some benthic octopod species. Finally, we estimated the energy balance for O. maya and E. megalocyathus in order to validate the use of this information for aquaculture or fisheries management. In both species, oxygen consumption and ammonia excretion increased allometrically with increasing body weight (BW) expressed as Y = aBW ^b . For oxygen consumption, b was 0.71 and 0.69 for E. megalocyathus and O. maya, respectively, and for ammonia excretion it was 0.37 and 0.43. Both species had low O/N ratios, indicating an apparent dependence on protein energy. The mean ingestion rates for E. megalocyathus (3.1 ± 0.2% its BW day⁻¹) and O. maya (2.9 ± 0.5% its BW day⁻¹) indicate that voracity, which is characteristic of cephalopods, could be independent of species. The scope for growth (P = I − (H + U + R) estimated for E. megalocyathus was 28% higher than that observed in O. maya (320 vs. 249 kJ day⁻¹ kg⁻¹). Thus, cold-water cephalopod species could be more efficient than tropical species. The protein and respiratory metabolisms of O. maya, E. megalocyathus, and other octopod species are directly dependent on temperature. Our results offer complementary evidence that, as Clarke (2004) stated, the metabolic response (R and U) cannot be determined mechanistically by temperature, as previously proposed (Gillooly et al. 2002).     

Diel cycles of activity,metabolism, and ammonium concentration in tropical holothurians

Movement rate, oxygen consumption, and respiratory tree ammonium concentration were measured in situ in the holothurians Pearsonothuria graeffei and Holothuria edulis in the Agan-an Marine Reserve, Sibulan, Philippines (9°20′30″N, 123°18′31″E). Measurements were made both day and night for both species during April–July 2005. P. graeffei had significantly higher movement rate during the day than at night (1.14 and 0.27 m h⁻¹, respectively; three-way ANOVA, P < 0.05) while H. edulis had higher movement rate at night compared to the day (0.83 and 0.07 m h⁻¹, respectively), spending the daylight hours sheltering under coral. More than 80% of H. edulis had movement rate of zero during the day. Oxygen consumption of P. graeffei was significantly higher during the day than at night (1.61 and 0.83 μmol O₂ g⁻¹ h⁻¹, respectively; two-way ANCOVA, P < 0.05), but the reduction at night was not as pronounced as the reduction in movement. H. edulis had a 75% reduction in oxygen consumption during the day compared to night (0.51 and 1.96 μmol O₂ g⁻¹ h⁻¹, respectively), matching this species’ reduced movement rates during the day. Ammonium concentration in water withdrawn from the respiratory trees of P. graeffei during the day (12.0 μM) was three times higher than in respiratory tree water sampled at night (4.3 μM) and 15 times higher than ambient seawater (0.8 μM; three-way ANOVA, P < 0.05). Ammonium concentration in the respiratory tree water of H. edulis was six times higher at night (14.6 μM) than during the day (2.2 μM) and 16 times higher than that of ambient seawater (0.9 μM). Even though H. edulis and P. graeffei are found within the same coral reef environment, they may affect different substrates and reef organisms due to their different habitats and distinct but opposite diel cycles.     

Redwood of the reef: growth and age of the giant barrel sponge Xestospongia muta in the Florida Keys

The growth of animals in most taxa has long been well described, but the phylum Porifera has remained a notable exception. The giant barrel sponge Xestospongia muta dominates Caribbean coral reef communities, where it is an important spatial competitor, increases habitat complexity, and filters seawater. It has been called the ‘redwood of the reef’ because of its size (often >1 m height and diameter) and presumed long life, but very little is known about its demography. Since 1997, we have established and monitored 12 permanent 16 m diameter circular transects on the reef slope off Key Largo, Florida, to study this important species. Over a 4.5-year interval, we measured the volume of 104 tagged sponges using digital images to determine growth rates of X. muta. Five models were fit to the cubed root of initial and final volume estimates to determine which best described growth. Additional measurements of 33 sponges were taken over 6-month intervals to examine the relationship between the spongocoel, or inner-osculum space, and sponge size, and to examine short-term growth dynamics. Sponge volumes ranged from 24.05 to 80,281.67 cm³. Growth was variable, and specific growth rates decreased with increasing sponge size. The mean specific growth rate was 0.52 ± 0.65 year⁻¹, but sponges grew as fast or slow as 404 or 2% year⁻¹. Negative growth rates occurred over short temporal scales and growth varied seasonally, significantly faster during the summer. No differences in specific growth rate were found between transects at three different depths (15, 20, 30 m) or at two different reef sites. Spongocoel volume was positively allometric with increasing sponge size and scaling between the vertical and horizontal dimensions of the sponge indicated that morphology changes from a frustum of a cone to cylindrical as volume increases. Growth of X. muta was best described by the general von Bertalanffy and Tanaka growth curves. The largest sponge within our transects (1.23 × 0.98 m height × diameter) was estimated to be 127 years old. Although age extrapolations for very large sponges are subject to more error, the largest sponges on Caribbean reefs may be in excess of 2,300 years, placing X. muta among the longest-lived animals on earth.     

Production and biomass accumulation of periphytic diatoms growing on glass slides during a 1-year cycle in a subtropical estuarine environment (Bay of Paranaguá, southern Brazil)

Production rates, chlorophyll concentrations and general composition of periphytic diatom communities growing on glass slides were studied in relation to environmental parameters during one seasonal cycle in the Bay of Paranaguá, southern Brazil. Slides were routinely submersed at 1, 2 and 3 m depth and recovered weekly for microscopic examinations, analyses of chlorophyll, cell counts and in situ photosynthetic incubations using the Winkler titration method. Water samples were also collected at surface and bottom layers for determinations of temperature, salinity, nutrients and chlorophyll in the water. The periphytic community was mainly formed by epipelic and epipsammic species, dominated by Navicula phyllepta, Cylindrotheca closterium, Navicula spp. and Amphora sp. Weekly chlorophyll a and cell accumulations on slides varied from <1–32 mg m⁻² and up to 31 × 10⁸cells m⁻², respectively. Photosynthetic rates varied from <1 to 35 mg oxygen mg chlorophyll a ⁻¹ h⁻¹, with higher values in summer. Daily production varied from 5 to 3,600 mg oxygen m⁻² day⁻¹ (<0.01–1.4 g carbon m⁻² day⁻¹). Multiple regression analysis revealed that vertical differences in light conditions and grazing pressure jointly affected the influence of temperature on the seasonal patterns of cell densities and chlorophyll concentrations according to depth. Received: 27 April 2000 / Accepted: 16 August 2000     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

Relationship between filtration activity and food availability in the Mediterranean mussel <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis>

The filtration activity of the Mediterranean mussel, Mytilus galloprovincialis, was assessed under different concentrations and compositions of seston by using a new automated image acquisition and analysis system. This approach allowed for frequent and simultaneous measurements of valve gape and exhalant siphon area. Filtration rates were measured through clearance measurements whereas pumping rates were measured using hot-film probes. The average filtration rate (17.5 l g h⁻¹ DW⁻¹ for a 0.36 g DW mussel) recorded during the present study was higher than those available for Mytilus edulis when standardized to flesh dry weight but almost equivalent (17.5 l h⁻¹ g DW⁻¹ for a 53 mm shell length mussel) to those rates when standardized to shell length. Immediately after the addition of algal cells (Isochrysis galbana; 4.5 μm in size), valve gape, exhalant siphon area and filtration rate increased quickly as mussels reached their maximum filtration activity. These three parameters then gradually decreased until complete closure of the shell. The algal cell concentration inducing this transition was close to 800 cells ml⁻¹ and 0.5 μg Chl a l⁻¹. When algal concentration was maintained above this threshold by successive algal additions, both valve gape and exhalant siphon area remained maximal. Temporal changes in the exhalant siphon area were continuous as opposed to those of valve gape. Therefore, despite the significant correlation between these two parameters, valves and siphon were sometimes dissociated due to a reduction of the area or even a closure of the exhalant siphon while the valves remained open. The velocity of exhaled water tended to be constant irrespective of exhalant siphon area and thus pumping rates were a linear function of exhalant siphon area. Consequently, reductions in exhalant siphon area and pumping rate were almost similar in M. galloprovincialis. Our results thus clearly support the hypothesis that exhalant siphon area constitutes a better proxy of pumping rate than valve gape as already suggested for Mytilus edulis. Finally, the high filtration rates measured during the present study together with the high concentrations of inorganic matter (> 40 mg DW l⁻¹) requested to alter those rates suggest that the studied mussels were well adapted to oligotrophic waters featuring strong hydrodynamism and frequent sediment resuspension events.     

Phytoplankton blooms are strongly impacted by microzooplankton grazing in coastal North Pacific waters

Phytoplankton growth and microzooplankton grazing were measured in two productive coastal regions of the North Pacific: northern Puget Sound and the coastal Gulf of Alaska. Rates of phytoplankton growth (range: 0.09–2.69 day⁻¹) and microzooplankton grazing (range: 0.00–2.10 day⁻¹) varied seasonally, with lowest values in late fall and winter, and highest values in spring and summer. Chlorophyll concentrations also varied widely (0.19–13.65 μg l⁻¹). Large (>8 μm) phytoplankton cells consistently dominated phytoplankton communities under bloom conditions, contributing on average 65% of total chlorophyll biomass when chlorophyll exceeded 2 μg l⁻¹. Microzooplankton grazing was an important loss process affecting phytoplankton, with grazing rates equivalent to nearly two-thirds (64%) of growth rates on average. Both small and large phytoplankton cells were consumed, with the ratio of grazing to growth (g:μ) for the two size classes averaging 0.80 and 0.42, respectively. Perhaps surprisingly, the coupling between microzooplankton grazing and phytoplankton growth was tighter during phytoplankton blooms than during low biomass periods, with g:μ averaging 0.78 during blooms and 0.49 at other times. This tight coupling may be a result of the high potential growth and ingestion rates of protist grazers, some of which feed on bloom-forming diatoms and other large phytoplankton. Large ciliates and Gyrodinium-like dinoflagellates contributed substantially to microzooplankton biomass at diatom bloom stations in the Gulf of Alaska, and microzooplankton biomass overall was strongly correlated with >8 μm chlorophyll concentrations. Because grazing tended to be proportionally greater when phytoplankton biomass was high, the absolute amount of chlorophyll consumed by microzooplankton was often substantial. In nearly two-thirds of the experiments (14/23), more chlorophyll was ingested by microzooplankton than was available for all other biological and physical loss processes combined. Microzooplankton were important intermediaries in the transfer of primary production to higher trophic levels in these coastal marine food webs. Received: 12 November 1999 / Accepted: 4 October 2000     

Seasonal variations in the growth rates of euphausiids (<Emphasis Type="Italic">Thysanoessa inermis</Emphasis>, <Emphasis Type="Italic">T. spinifera</Emphasis>, and <Emphasis Type="Italic">Euphausia pacifica</Emphasis>) from the northern Gulf of Alaska

The euphausiids Thysanoessa inermis (Kroyer 1846), Thysanoessa spinifera (Holmes 1900), and Euphausia pacifica (Hansen 1911) are key pelagic grazers and also important prey for many commercial fish species in the Gulf of Alaska (GOA). To understand the role of the euphausiids in material flows in this ecosystem their growth rates were examined using the instantaneous growth rate (IGR) technique on the northern GOA shelf from March through October in 2001–2004. The highest mean molting increments (over 5% of uropod length increase per molt) were observed during the phytoplankton bloom on the inner shelf in late spring for coastal T. inermis, and on the outer shelf in summer for T. spinifera and more oceanic E. pacifica, suggesting tight coupling with food availability. The molting rates were higher in summer and lower in spring, for all species and were strongly influenced by temperature. Mean inter-molt periods calculated from the molting rates, ranged from 11 days at 5°C to 6 days at 8°C, and were in agreement with those measured directly during long-term laboratory incubations. Growth rate estimates depended on euphausiid size, and were close to 0 in early spring, reaching maximum values in May (0.123 mm day⁻¹ or 0.023 day⁻¹ for T. inermis) and July (0.091 mm day⁻¹ or 0.031 day⁻¹ for T. spinifera). The growth rates for E. pacifica remained below 0.07 mm day⁻¹ (0.016 day⁻¹) throughout the season. The relationship between T. inermis weight specific growth rate (adjusted to 5°C) and ambient chlorophyll-a concentration fit a Michaelis–Menten curve (r ² = 0.48) with food saturated growth rate of 0.032 day⁻¹ with half saturation occurring at 1.65 mg chl-a m⁻³, but such relationships were not significant for T. spinifera or E. pacifica.     

Resource acquisition in two intertidal fucoid seaweeds, Fucus serratus and Himanthalia elongata : seasonal variation and effects of reproductive development

Seasonal variations and the effect of reproductive development on resource acquisition by two intertidal fucoid species, the iteroparous Fucus serratus L. and the semelparous Himanthalia elongata (L.) S. F. Gray were examined. The oxygen-exchange characteristics of vegetative apical tissue of both non-fertile and fertile plants and receptacle tissue were compared at monthly intervals throughout reproductive development. Respiratory rates in non-fertile F. serratus varied seasonally between 1.5 and 8.0 μmol g⁻¹ fresh wt h⁻¹; in fertile plants the receptacle had a significantly lower respiratory rate than the vegetative tissue. The respiratory rate of the vegetative button of fertile H. elongata displayed less seasonal variation and was lower than that of the receptacle, which varied from a maximum of 9.5 μmol g⁻¹ fresh wt h⁻¹ at receptacle initiation in October to a minimum of 2.0 μmol g⁻¹ fresh wt h⁻¹ in February. The maximum photosynthetic rate (P _max) of non-fertile plants of both species did not vary in a distinct seasonal manner (∼60 μmol g⁻¹ fresh wt h⁻¹ for F. serratus and ∼12 μmol g⁻¹ fresh wt h⁻¹ for H. elongata). In fertile plants, the P _max of the receptacle tissue was (∼50% lower in F. serratus, and at its peak three times higher in H. elongata, than that of vegetative tissue. The stable carbon-isotope ratio (δ¹³C) did not differ between different tissue types in F. serratus, but values did vary seasonally, being less negative in the summer than in the winter (−13.5‰ compared to −18‰). The receptacle tissue of H. elongata also displayed a distinct seasonal variation in δ¹³C values (−12‰ in summer, −16‰ in winter), whilst the δ¹³C of the vegetative button did not vary seasonally. The rate of uptake of inorganic nitrogen by the vegetative thallus was lower in H. elongata than in F. serratus. The receptacle tissue of F. serratus had lower uptake rates than the vegetative tissue, whilst the uptake rate by H. elongata receptacle tissue was higher than that of the vegetative button. Received: 14 March 1997 / Accepted: 22 April 1997     

Functional responses of copepod nauplii using a high efficiency gut fluorescence technique

To investigate copepod nauplii ingestion rates on phytoplankton, we have adapted the traditional gut fluorescence technique as it can be used with lower gut pigment concentrations. With the improved technique, laboratory experiments were performed to estimate functional responses for nauplii of Calanus helgolandicus and Centropages typicus. Nauplii were raised from eggs to copepodites and the experiments were performed with stages NIV-NV. Gut evacuation rates and ingestion rates were measured on Isochrysis galbana at different concentrations. Specific ingestion rates ranged between 0.038–0.244 μg C μg⁻¹ nauplii C d⁻¹ for C. typicus and 0.041–1.412 μg C μg⁻¹ nauplii C d⁻¹ for C. helgolandicus. Both species showed a type III functional response, reaching a saturation concentration at around 600 μgC l⁻¹ for C. typicus and 800 μgC l⁻¹ for C. helgolandicus. An erratum to this article can be found at