Filter-feeding ethology of benthic invertebrates (ascidians). V. Influence of temperature on pumping,filtration and digestion rates and rhythms in Phallusia mamillata

The influence of temperature has been studied simultaneously on the pumping, filtration, and digestion rates of Phallusia mammillata (Cuvier, 1815). Eighteen experiments were made between 7° and 25°C on 5 individuals. The average velocities of the water current varied between 3.37 and 9.65 cm sec^-1 (maximum 34.90 cm sec^-1). No recognizable rhythm emerged; pumping was continuous except at 7°C, where it soon ceased. Above 20°C, the curves were irregular and reflected the high sensitivity of the ascidian. The pumping rate was highest at 15°C (mean=5,788 ml h^-1 g^-1 dry weight of organs). At 10°C, the mean was 3,560; at 20°C, 2,629 ml h^-1 g^-1 dry weight of organs. At 20°C, the coefficients of variation displayed higher values, indicating a more irregular pumping at this temperature. Although there was no filtration rhythm, the variability of the results was higher at 20°C and above. As for pumping, maximum values were observed at 15°C (mean=4,286 ml h^-1 g^-1 dry weight of organs) decreasing with lower and higher temperatures, such decreases being more marked at the higher temperatures. Means were 352 ml h^-1 g^-1 dry weight of organs at 7°C; 2,935 at 10°C; 1,995 at 20°C; 973 at 25°C. The mean temperature coefficients for the filtration rates were: Q₁₀ for 7° to 15°C=11.86, Q₁₀ for 10° to 20°C=0,66, Q₁₀ for 15° to 25°C=0.22. The filtering efficiency was fairly constant throughout an experiment; the pumping and filtration curves were in fact almost parallel. The filtering efficiency of the branchial sac was high (75 to 85%), with constant values at 10° and 15°C; it became smaller (59%) at 20°C, with a higher coefficient of variation. The digestion rate also displayed maximum values at 15°C (mean=5.47 mg of albumin equivalent 24 h^-1 g^-1 dry weight of organs). It was lower at 10°C (mean=3.60 mg) and reached its minimum at 20°C (mean=1.71 mg). The higher temperature affected the percentage of food utilization, which showed smaller values at 20°C (59%) than at 10°C (89%) and 15°C (87%).     

Filter-feeding ethology of benthic invertebrates (Ascidians). III. Recording of water current in situ — Rate and rhythm of pumping

Water current in 4 species of ascidians (Ascidia mentula Müller, 1776; Phallusia mammillata Cuvier, 1815; Styela plicata (Lesueur, 1823) and Microcosmus sabatieri Roule, 1885) has been recorded for the first time in situ with hot film probes. The mathematical treatment of probe calibration has allowed the evaluation of the characteristics of currents (minimal, maximal and average velocities) and the calculation of pumping rates. The method utilized yields qualitative and quantitative results: Pumping is continuous, and no particular rhythm shows up in the curves obtained. The velocity of the current does not change in the course of a recording, but it can vary from one experiment to another in the same individual; these variations are related to the variability of the diameter of the cloacal siphon rather than to a change in the rhythm of the ciliary activity. The average velocities are 3.42 cm sec^-1 for A. mentula, 3.92 for P. mammillata, 3.30 for S. plicata and 3.79 for M. sabatieri. The pumping rates are fairly constant for the same individual. The means, expressed as ml x h^-1 x g^-1 organ dry weight are 3422 for A. mentula, 2070 for P. mammillata, 4892 for S. plicata and 3084 for M. sabatieri.This work was supported by the C.N.R.S. (L.A. No. 117) and C.N.E.X.O. (Convention No. 76/1497).     

Mathematical analysis of the effects of particle retention efficiency on determination of filtration rate

Morphological, ecological, and behavioral aspects of filtration of small, inefficiently retained particles by benthic as well as planktonic filter-feeders are of current as well as historical interest. The exponential model applied to determination of filtration rate was originally conceived for filter-feeders clearing suspensions of large, efficiently retained cells. Therefore, applicability of this model and its assumptions to filtration of small, inefficiently retained particles merits critical re-examination. Derivation of the exponential model historically used in calculation of filtration rate is reviewed and shown to be inappropriate when filtration efficiency is less than 100%. Box models are constructed for filter-feeders that act either as sieves or aerosol filters in suspensions of small inefficiently filtered cells. The models predict a complex or double exponential decline in cell concentration that cannot easily be translated into filtering or pumping rates. The models further predict that apparent filtration rate will decline over time because of physical limitations, rather than behavioral changes, with respect to filtration efficiency. Compartmental analysis of filtration of inefficiently retained particles in a turbidostat system predicts a similar artifact in determination of filtration rate.     

Microplastics in the benthic invertebrates from the coastal waters of Kochi,Southeastern Arabian Sea

This study examined microplastic particles present in the benthic invertebrates Sternaspis scutata, Magelona cinta (deposit feeders) and Tellina sp. (suspension feeder) from the surface sediments of off-Kochi, southwest coast of India. The microplastic particles and thread-like fibres detected in these organisms were identified to be polystyrene by using DXR Raman microscope. Examination of the microplastic particle in Sternaspis scutata by epifluorescent microscopy showed fragmentation marks on the surface suggesting that the microplastic particle was degraded/weathered in nature. The study provides preliminary evidence of the presence of microplastics in benthic fauna from the coastal waters of India. However, further studies are required to understand the sources, distribution, fate and toxicity of the different types of microplastics in benthic invertebrates in order to identify any potential threats to higher trophic level organisms.     

Relative importance of benthic microalgae,phytoplankton, and mangroves as sources of nutrition for penaeid prawns and other coastal invertebrates from Malaysia

Carbon, sulfur, and nitrogen stable isotope ratio techniques were used in 1989 and 1990 to evaluate the relative importance of algae and of mangrove detritus in the nutrition of two penaeid prawn species on the west coast of Peninsular Malaysia. Mangrove detritus was found to contribute to the nutrition of juvenile Penaeus merguiensis de Man living within tidal creeks, but not to adult P. merguiensis and juvenile and adult Parapenaeopsis sculptilis (Heller) captured offshore. Results from radiotracer feeding studies, with refractory ¹⁴C mangrove lignocellulose as the food source, indicated that juvenile P. merguiensis from tidal creeks assimilated mangrove carbon with an efficiency of 13.4%. This did not differ significantly (P=0.05) from the assimilation efficiencies of juvenile and adult P. sculptilis living offshore (10.0 and 10.9%, respectively); these values were significantly higher (P<0.001) than for adult P. merguiensis (2.1%). Thus, the differential contribution of mangrove material to the nutrition of prawns in the tidal creeks and offshore was not related to differences in the prawn's ability to utilize detritus. Instead, our analysis of C stable isotopes in sediments indicated that mangrove detritus was generally more abundant within the tidal creeks than offshore. Juvenile prawns in the tidal creeks may also utilize mangrove material indirectly by feeding on small detritivorous invertebrates. Stable isotope analyses suggest that benthic microalgae constitute the other major dietary component for prawns living in tidal creeks. Prawns offshore were utilizing mainly phytoplankton-based material.     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Influence of various concentrations of orthophosphate on the division rate of an estuarine benthic diatom,Navicula arenaria,in culture

A hydrolysis procedure along with a high-pressure liquid chromatographic procedure is given enabling simple and reliable thymine determinations in the nanogram range in different fractions of sea-water samples taken from three different locations in the Northern Adriatic Sea. The levels corresponded to 1–3 g DNA per liter. From total polyanionic thymine, which had been precipitated as the cetyltrimethylammonium salt, the highest percentage was linked to the particulate fraction, with a definite subsurface minimum at 10 to 15 m. There was a corresponding maximum of a high molecular non-particulate thymine-containing fraction at the corresponding depth. From the bottom at 30 m upwards to about 20 m, a low molecular thymine-containing material has been found. Remarkably these basic features were common to all three locations, one of which was supposed to be in clean water, one near a thickly settled, touristic area, and the other in front of a large river delta coming from industrial hinterland.     

Growth rate of the bathypelagic crustacean Gnathophausia ingens (Mysidacea: Lophogastridae). I. Dimensional growth and population structure

Gnathophausia ingens has 13 instars, each with a distinct range of sizes which does not overlap the sizes of adjacent instars. The intermolt interval, measured in the laboratory at 5.5°, 6.5° and 7.5°C, increases with increasing size and decreases with increasing temperature. At 5.5°C it varies from 166 days for the smallest individuals to 253 days for the oldest. The period of larval development in the marsupium of a female is estimated to be 530 days. The life span of females is estimated to be 2,950 days with the onset of reproduction at 2,400 days. It is sugquested that this species is semelparous. The population structure data suggest that there is low mortality through the first 7 instars, progressively higher mortality from Instar 8 through Instar 11, and slightly lower mortality in the remaining 2 instars. These life-history characteristics appear to be directed toward maximizing absolute fecundity (as opposed to time-specific fecundity) in a stable environment. These characteristics may have been selected for by low available food energy and made possible by the stability of the deep sea.     

The ultrastructure of epidermal mucous cells in marine invertebrates (Nemertini,Polychaeta, Prosobranchia,Opisthobranchia)

The epidermal mucous cells of various nemerteans, polychaetes and gastropods exhibit a highly heterogeneous fine structure, which is not correlated to systematic position or way of living. Formation and extrusion of the mucus secretory substances also follow different patterns. In the majority of species investigated, the mucus inclusions originate in the Golgi area, in Pomatias they form in the rough endoplasmic reticulum. The mucus droplets either coalesce in the cytoplasm or retain their individuality before being extruded.Supported by the Deutsche Forschungsgemeinschaft (Sto 75/2, We 380/4).     

Trophic state,ecosystem efficiency and biodiversity of transitional aquatic ecosystems: analysis of environmental quality based on different benthic indicators

Estuaries and coastal lagoons are characterized by a strong spatial and temporal variability of physicochemical characteristics and productivity patterns. In these environments, the magnitude and direction of the ecological responses to inorganic nutrient increase (i.e. eutrophication) are difficult to predict. In the framework of the project, New Indicators of Trophic state and environmental quality of marine coastal ecosystems and transitional environments (NITIDA), we analysed benthic indicators of trophic state, ecosystem efficiency, and environmental quality in four different transitional environments. The trophic state of the sediments was assessed in terms of quantity and bioavailability of sediment organic C pools; ecosystem efficiency was determined in terms of the prokaryote efficiency in exploiting enzymatycally degraded organic C; environmental quality was determined in terms of meiofaunal diversity. Here, we provide a synopsis of the results obtained and a meta-analysis of the scores assessments obtained using the different ecological indicators of environmental quality and demonstrate that trophic state, ecosystem efficiency, and biodiversity in transitional ecosystems are closely linked. We conclude that the assessment of the environmental quality of transitional ecosystems should be based upon a battery of trophic state indicators and ‘sensors’ of ecosystem functioning, efficiency, and quality.     

Trophic state, ecosystem efficiency and biodiversity of transitional aquatic ecosystems: analysis of environmental quality based on different benthic indicators

Estuaries and coastal lagoons are characterized by a strong spatial and temporal variability of physicochemical characteristics and productivity patterns. In these environments, the magnitude and direction of the ecological responses to inorganic nutrient increase (i.e. eutrophication) are difficult to predict. In the framework of the project, New Indicators of Trophic state and environmental quality of marine coastal ecosystems and transitional environments (NITIDA), we analysed benthic indicators of trophic state, ecosystem efficiency, and environmental quality in four different transitional environments. The trophic state of the sediments was assessed in terms of quantity and bioavailability of sediment organic C pools; ecosystem efficiency was determined in terms of the prokaryote efficiency in exploiting enzymatycally degraded organic C; environmental quality was determined in terms of meiofaunal diversity. Here, we provide a synopsis of the results obtained and a meta-analysis of the scores assessments obtained using the different ecological indicators of environmental quality and demonstrate that trophic state, ecosystem efficiency, and biodiversity in transitional ecosystems are closely linked. We conclude that the assessment of the environmental quality of transitional ecosystems should be based upon a battery of trophic state indicators and 'sensors' of ecosystem functioning, efficiency, and quality.     

Atmospheric dinitrogen fixation by benthic communities of Tikehau Lagoon (Tuamotu Archipelago, French Polynesia) and its contribution to benthic primary production

Acetylene reduction rates were measured in lagoonal sediments, cyanobacterial mats and limestone surfaces between 1991 and 1995 at many sites, depths and seasons; all the studied substrata contained cyanobacteria. The acetylene reduction/¹⁵N₂ fixation ratio was measured for the different communities and varied between 1.8 and 4.8, depending on substratum. Fixation rates were 1.7 to 7 times higher during daylight compared to night-time rates. N₂ fixation rates ranged from 0.4 to 3.9 mg N m^-2 day^-1 for the lagoonal sediment/mat communities, and the rate was about 2 mg N m^-2 day^-1 for the lagoonal limestone substrata. Total lagoonal benthic N₂ fixation contributed 24.4% of the total nitrogen requirement for the benthic primary production of benthic communities of the lagoon. The input of N₂ fixation by the microbial planktonic communities (including cyanobacteria) of the lagoon, which are highly productive, is unquantified but is likely to be large.     

Ethologie alimentaire d'invertébrés benthiques filtreurs (ascidies). I. Dispositif expérimental. Taux de filtration et de digestion chez Phallusia mammillata

In a study on ascidian feeding ethology, rates of filtration and digestion of the unicellular algae Monchrysis lutheri by Phallusia mammillata (Cuvier, 1815) have been determined experimentally. All results were obtained under constant conditions of temperature, salinity, pH, light, and food, using 5 individuals in 9 experiments of 24 h each. The method of Winter (1969) has been modified by using an automatic system which enables constant food concentrations (±5%) to be maintained. Controls are made by a fluorometer. The rate of filtration is deduced from the quantity of culture food added each hour; this rate varies; a well defined rhythm is not observed, but alternating maxima and minima. The values obtained for, specimens of medium size (10 to 12 cm) vary from 3856 to 4730 ml/h and per gramme organic dry weight, the average being 4380 ml. The quantity of food actually converted by each ascidian, once it has passed the alimentary canal, is calculated from the amount of proteins present in food and faeces. Digestion rate varies from 4.65 to 5.75 mg of albumin equivalent/24 h and per gramme organic dry weight (mean 5.30 mg). The percentage food conversion (filtrated amount=100%) is high: 86 to 93%, with a mean of 90%.     

Metal levels in some benthic organisms of Haifa Bay,mediterranean (Israel)

Benthic organisms revealed enhanced uptake of mercury from polluted surface sediments (in the vicinity of a chlor‐alkali plant) which were enriched in mercury by up to 148 times relative to sediments from an unpolluted area. There is a strong correlation between the concentrations of mercury in the benthic organisms and sediments. No correlation was found between cadmium, lead, copper and zinc levels in the benthic fauna populations and the concentrations of these metals in the sediments. Ranges of concentrations (μg/g wet wt.) in the whole soft tissues varied: Hg, <0.005–3.31; Cd <0.018–1.99; Pb, <0.025–1.56; Cu, 0.242–28.8; and Zn, 9.75–310.2. The three gastropod molluscs show higher metal concentrations than do the bivalves.     

Comparative species richness,composition, and abundance of invertebrates in Caribbean seagrass (Thalassia testudinum) meadows (Panamá)

The results of a year-long study in which epibenthic invertebrates were collected monthly from seagrass (Thalassia testudinum) meadows along the Caribbean coast of Panamá and the Panamá Canal Zone are described in this paper. Differences in species composition and abundance among sites were primarily due to the proximity of surrounding habitats, especially coral reefs, which contain a number of species that utilize the seagrass meadows. In contrast to many previous characterizations of tropical marine habitats, important seasonal fluctuations in both species number and abundance took place at each of the sites. Data on breeding activity among several species of decapod crustaceans indicate year-round reproduction, although considerable seasonal differences occur in the percentage of ovigerous females. These interspecific differences in observed reproductive output may be explained by differences in life-cycle length, a factor not often considered in discussions of seasonal breeding patterns in tropical marine invertebrates. Overall species composition was qualitatively similar to that reported in comparable studies of tropical and subtropical seagrass meadows elsewhere, although caridean shrimp and xanthid orab species were reduced in number and total abundance were much lower than in previous studies.     

The filtration rate of Mytilus edulis and its dependence on algal concentration,measured by a continuous automatic recording apparatus

A new apparatus for long-term, continuous automatic measurements of filtration rates in suspension-feeding organisms is described. As the concentration of algae in the experimental medium is diminished by the filter-feeding activity of the experimental animals, algal suspension is automatically added, thus keeping the algal concentration constant. In this way, accurate determinations of filtration rates in relation to particle concentration are made possible. For determination of filtration rates in the common mussel Mytilus edulis L., individuals of different body size (shell length 8.5 to 56.5 mm) were used. Within the range of 10x10⁶ to 40x10⁶ cells of Dunaliella marina/l, mussels of the same body size filter-out approximately the same amount of algae at high or low concentrations. A low algal concentration is counterbalanced by a corresponding higher filtration rate. Within the range of body size (W=dry weight of tissues) and algal concentrations used, the filtration rate (F) follows the general allometric equation F=a·W ^b, where a and b are constants at specific experimental conditions. At a temperature of 12 °C, the values obtained for a are 2410 at a concentration of 20x10⁶, and 1313 at a concentration of 40x10⁶ Dunaliella cells/l; correspondingly, the filtration rates of a mussel of 1 g dry-tissue weight are 2410 ml/h and 1313 ml/h. b, the slope of the regression line (0.73 to 0.74), is independent of algal concentration. However, examination of all known measurements reveals that, most probably, the general allometric equation is an oversimplification; in large individuals there is a more pronounced decrease in filtration rate. The relationship between filtration rate, body size of mussels, and algal concentrations used is discussed.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung — Abwässer in Küstennähe.     

Ethologie alimentaire d'invertébrés benthiques filtreurs (ascidies). II. Variations des taux de filtration et de digestion en fonction de l'espèce

Rates of filtration and digestion of 4 species of ascidians (Clavelina lepadiformis), Müller, Ciona intestinalis (Linné, 1767), Halocynthia papillosa (Linné, 1767) and Microcosmus sabatieri (Roule, 1885) from a rocky shore at banyuls-sur-Mer, France have been studied using the unicellular algae Monochrysis lutheri as food and the methods previously applied to Phallusia mammillata (Fiala-Médioni, 1973). Eleven experiments of 24 h each were performed under constant conditions of temperature, pH, salinity, oxygen and food concentration. No recognizable feeding rhythm emerged; filtration is irregular, varying around a mean value. Observed filtration rates averaged 2489 ml/h/g organ dry weight in Clavelina lepadiformis, 3515 in Ciona intestinalis, 6349 in Halocynthia papillosa and 6909 in Microcosmus sabatieri. The higher rates are related to larger size and higher complexity of the gills. No pseudo-faeces are formed; a very small part of the faecal material is discharged within 24 h. The mean rates of digestion, in mg albumin equivalent/24 h/g organ dry weight, are: Clavelina lepadiformis, 3.05 mg; Ciona intestinalis, 4.74 mg; Halocynthia papillosa, 9.25 mg; Microcosmus sabatieri, 10.41 mg. The amount of digested algae corresponds to 83–92% (mean=85%) of algae filtered. This high percentage indicates good assimilation of Monochrysis lutheri by ascidians.

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Ethologie alimentaire d'invertébrés benthiques filtreurs (ascidies). II. Variations des taux de filtration et de digestion en fonction de l'espèce

     

Growth and reproductive cycles of the marine fouling ascidians Ciona intestinalis,Styela plicata,Botrylloides violaceus,and Leptoclinum mitsukurii at Aburatsubo-Moroiso Inlet (central Japan)

Experimental cultivations of post-metamorphosis juveniles were repeated in different seasons for Ciona intestinalis (L.) and Styela plicata (Lesueur) (simple ascidians) as well as for Leptolinum mitsukurii (Oka) and Botrylloides violaceus Oka (compound ascidians) at Aburatsubo-Moroiso Inlet near Misaki Biological Station (Japan). Growth in body length of simple ascidians was exponential during juvenile development up to near sexual maturity. C. intestinalis grew to sexual maturity in 1 month in summer and in 2 months in winter; S. plicata, in 2 months in summer and in nearly 5 months in winter. The doubling time decreased approximately twofold in the two simple ascidians with a 10C° increase in environmental temperature. Growth of colonies in compound ascidians was exponential after maturation of the first functional zooids. Sexual reproduction of L. mitsukurii and B. violaceus was continuous throughout the year, forming many generations in a year. Doubling time of colony growth in the two compound ascidians decreased approximately threefold with a 10C° increase in temperature.