Food habits, gut morphology and pH, and assimilation efficiency of the zebraperch Hermosilla azurea, an herbivorous kyphosid fish of temperate marine waters

Adult zebraperch, Hermosilla azurea, were found to be functional herbivores in that animal matter constituted <0.01% of the total dry weight of stomach contents of fish collected off Santa Catalina Island in southern California waters. The diet of these fish consisted mainly of red algae (88.2% by dry wt) and also small amounts of brown (7.8%) and green (4.0%) algae. The most important dietary item, the filamentous red algae Polysiphonia spp., was found in >78% of the stomachs and comprised >60% of the contents by dry weight. The digestive tract was long, on average 4.0 times the standard length of the fish, and was composed of the stomach, pyloric caeca, intestine, hindgut chamber with a blind caecum, and rectum. The mean pH of the cardiac stomach was acidic (3.9), whereas that of the intestine was nearly neutral (6.9) and that of the hindgut and blind caecum slightly acidic (6.3 and 6.6, respectively). Algal foods are apparently digested by acid lysis in the stomach and by microbial fermentation in the hindgut. Zebraperch assimilated nutritional constituents from six species of algae with varying degrees of efficiency: carbon (73.7 to 89.7%), nitrogen (72.4 to 84.5%), and protein (71.9 to 94.9%). The fish assimilated these constituents as efficiently or more efficiently from three species of nondietary brown algae as from three species of dietary red and green algae. These results show that zebraperch, like their tropical and subtropical relatives (members of the genus Kyphosus), can digest a wide variety of algae including brown algae containing defensive secondary compounds. Received: 3 November 1997 / Accepted: 19 June 1998     

Effect of nutrient enrichment in the field on the biomass, growth and calcification of the giant clam Tridacna maxima

Nutrients were added separately and combined to an initial concentration of 10 μM (ammonium) and/or 2 μM (phosphate) in a series of experiments carried out with the giant clam Tridacna maxima at 12 microatolls in One Tree Island lagoon, Great Barrier Reef, Australia (ENCORE Project). These nutrient concentrations remained for 2 to 3 h before returning to natural levels. The additions were made every low tide (twice per day) over 13 and 12 mo periods for the first and second phase of the experiment, respectively. The nutrients did not change the wet tissue weight of the clams, host C:N ratio, protein content of the mantle, calcification rates or growth rates. However, ammonium (N) enrichment alone significantly increased the total population density of the algal symbiont (Symbiodinium sp.: C = 3.6 · 10⁸ cell clam⁻¹, N = 6.6 · 10⁸ cell clam⁻¹, P = 5.7 · 10⁸ cell clam⁻¹, N + P = 5.7 · 10⁸ cell clam⁻¹; and C = 4.1 · 10⁸ cell clam⁻¹, N = 5.1 · 10⁸ cell clam⁻¹, P = 4.7 · 10⁸ cell clam⁻¹, N + P = 4.5 · 10⁸ cell clam⁻¹, at the end of the first and second phases of the experiment, respectively), although no differences in the mitotic index of these populations were detected. The total chlorophyll a (chl a) content per clam but not chlorophyll a per cell also increased with ammonium addition (C = 7.0 mg chl a clam⁻¹, N = 13.1 mg chl a clam⁻¹, P = 12.9 mg chl a clam⁻¹, N + P = 11.8 mg chl a clam⁻¹; and C = 8.8 mg chl a clam⁻¹, N = 12.8 mg chl a clam⁻¹; P = 11.2 mg chl a clam⁻¹, N + P = 11.3 mg chl a clam⁻¹, at the end of the first and second phases of the experiment, respectively). The response of clams to nutrient enrichment was quantitatively small, but indicated that small changes in inorganic nutrient levels affect the clam–zooxanthellae association. Received: 2 June 1997 / Accepted: 9 June 1997     

Assimilation efficiency of a temperate-zone intertidal fish (Cebidichthys violaceus) fed diets of macroalgae

We studied assimilation efficiencies of the temperate-zone intertidal fish Cebidichthys violaceus (Girard, 1854) fed in the laboratory on each of the following species of macroalgae: Spongomorpha coalita (Chlorophyta), Ulva lobata (Chlorophyta), Iridaea flaccida (Rhodophyta) and Porphyra perforata (Rhodophyta). Together, these 4 algae make up over 75% of the natural summer diet of C. violaceus. Assimilation efficiency was calculated by proximate organic analysis of food and feces; the amount of ash in food and feces was used as a standard. Depending on the algal species, the fish assimilated 43 to 81% of the protein, 21 to 44% of the lipid, 45 to 62% of the carbohydrate and 31 to 52% of all three classes of organic material combined. These data are the first results showing that a temperate-zone marine fish can assimilate macroalgal constituents. Protein, carbohydrate and total organic material were absorbed more efficiently from rhodophytes than from chlorophytes. Conversely, lipid was absorbed more efficiently from chlorophytes than from rhodophytes. These results are compared with previous work showing that C. violaceus in nature eats more chlorophytes than rhodophytes, but in laboratory preference tests prefers rhodophytes to chlorophytes.     

The trophic status of herbivorous fishes on coral reefs

Estimates of feeding rates, alimentary tract structure and temporal patterns of food processing obtained from twelve species of nominally herbivorous fishes on the northern Great Barrier Reef were compared. These included members of the families Acanthuridae, Scaridae and Kyphosidae. Based on an analysis of diet and short-chain fatty acid (SCFA) profiles from a previous study we initially partitioned the twelve species into four dietary categories, as follows: (a) Category 1: herbivores with a diet of macroscopic brown algae and high SCFA profiles in the hindgut region (Naso unicornis, Kyphosus vaigiensis); (b) Category 2: herbivores feeding on turfing and filamentous red and green algae with moderate SCFA profiles in the hindgut region (N. tonganus, K. cinerascens, Zebrasoma scopas, Acanthurus lineatus); (c) Category 3: zooplankton feeders with moderate SCFA profiles (N. vlamingii, N. brevirostris); (d) Category 4: species feeding on detrital and sedimentary materials with low levels of SCFA (Chlorurus microrhinos, Scarus schlegeli, Ctenochaetus striatus, A. olivaceus). The purpose of this comparison was to determine whether measures of feeding activity, alimentary tract structure, and food processing were concordant with diet. A dichotomy in feeding rates was observed. Species with a diet of algae and zooplankton (categories 1–3) had slower feeding rates than those feeding on detrital aggregates and sediment (category 4). The pattern of food processing also followed the same dichotomy with species of categories 1–3 retaining food in the alimentary tract overnight and commencing the feeding day with substantial amounts of food in the intestine and hindgut. Category-4 species commenced the feeding day with empty alimentary tracts suggesting a rapid turnover of gut contents. Within the herbivorous and zooplankton-feeding species neither alimentary tract structure nor food processing mode were predicted by diet or SCFA profiles. A hindgut fermentation chamber was present in K. vaigiensis but not in N. unicornis, a species with high levels of SCFA in the hindgut region and a diet of brown macroscopic algae. In contrast N. vlamingii, with a diet dominated by animal matter, retained large amounts of food material in a hindgut chamber over the entire feeding cycle. In tropical perciform fishes, herbivory and fermentation are not associated with the alimentary tract structures that characterise herbivorous terrestrial vertebrates. Estimates of the abundance of the different groupings of nominally herbivorous fishes indicated that the dominant elements in the reef grazing and browsing fauna were consumers of detrital and sedimentary materials. These could not be classified as herbivores. Members of this group were dominant in all habitats investigated. Explicitly herbivorous taxa were a minority component in all habitats investigated.Communicated by G.F. Humphrey, Sydney     

Feeding and predation impact of two chaetognath species, Eukrohnia hamata and Sagitta gazellae, in the vicinity of Marion Island (southern ocean)

The predation impact of the two chaetognaths Eukrohnia hamata and Sagitta gazellae on mesozooplankton standing stock were investigated in three depth layers during two 24 h stations occupied in the vicinity of Marion Island in late austral summer (April/May) 1986. The zooplankton community at both stations was dominated by small copepods (Oithona spp., Microcalanus spp.), which accounted for >95% of total zooplankton abundance. Chaetognaths comprised <2% of total zooplankton abundance. E. hamata constituted >95% of the total chaetognath stock. The general trend in both species was decreasing abundance with increasing depth, which appeared to be correlated to the distribution of copepods (r ² = 0.45; P <0.05). Gut-content analysis showed that copepods (mainly Oithona spp., Calanus spp. and Rhincalanus gigas) and ostracods were the main prey of both species, accounting for 87 and 61% of the total number of prey in E. hamata and S.␣gazellae stomachs, respectively. In the guts of S.␣gazellae, pteropods (Limacina spp.) and chaetognaths were also well represented. The mean number of prey items (NPC) for E. hamata ranged from 0.02 to 0.06 prey individual⁻¹ which corresponds to an individual feeding rate (Fr) of between 0.05 and 0.12 prey d⁻¹. For S.␣gazellae, the NPC values were higher, varying between 0.04␣and 0.20 prey individual⁻¹, or between 0.15 and 0.76 prey d⁻¹. The daily predation impact of the two chaetognaths was estimated at between 0.3 and 1.2% of the copepod standing stock or between 7 and 16% of the daily copepod production. Predation by S. gazellae on chaetognaths accounted for up to 1.6% of the chaetognath standing stock per day. Received: 26 November 1996 / Accepted: 31 October 1997     

Effects of host-tissue homogenate of the scleractinian coral Plesiastrea versipora on glycerol metabolism in isolated symbiotic dinoflagellates

Symbiotic dinoflagellate algae (Symbiodinium sp.) isolated from the scleractinian coral Plesiastrea versipora and incubated in homogenized host tissue released 4 to 7 times as much glycerol (14 to 46 nmol glycerol/10⁶ algae) as those incubated in seawater (3 to 6 nmol glycerol/10⁶ algae) after 4 h incubation in the light. During this period, no release of triglycerides was detected. Intracellular glycerol increased 2- to 3-fold in algae incubated in host homogenate, but remained unchanged in algae incubated in seawater at a concentration of 0.82 ± 0.47 nmol glycerol/10⁶ algae. In each incubation condition, intracellular triglyceride levels increased. However, in algae incubated in host homogenate, the intracellular levels of triglycerides reached only about 75% of the amount reached in algae incubated in seawater (max. 18.55 ± 2.40 nmol glycerol/10⁶ cells). Host homogenate did not stimulate the release of glycerol from algae during dark incubation. These data show that the glycerol released by algae incubated in host-tissue homogenate was derived from increased synthesis of glycerol or from diversion of some glycerol or other photosynthetic intermediates from incorporation into algal triglyceride stores, and did not come from existing stores. Received: 20 December 1996 / Accepted: 9 January 1997     

Role of cold resistance and burial for winter survival and spring initiation of an Ulva spp. (Chlorophyta) bloom in a eutrophic lagoon (Veerse Meer lagoon, The Netherlands)

In the eutrophic Veerse Meer lagoon (The Netherlands) large amounts of free-floating thalli from Ulva spp. are present from May to October. In winter however, no algae seem to occur in the lagoon. Sexual reproduction appears to be negligible, as spore formation and germling growth are observed only sporadically. Results of a field survey showed that in winter, viable Ulva biomass is present buried in the sediment of the shallow parts of the lagoon. Freezing experiments demonstrated that the algae are able to survive temperatures of −5 °C for 2 weeks when kept in darkness. In spring, the buried Ulva thalli are liberated out of the sediment to initiate a bloom. A field experiment indicates that bioturbation by the lugworm Arenicola marina does not stimulate the release of the thalli. Burial and winter survival can explain the rapid increase in Ulva biomass in spring and suggest that the initial spring biomass is one of the major factors determining the maximal biomass in summer. Received: 28 October 1997 / Accepted: 20 January 1998     

Characterization of barnacle (Balanus eburneus and B. cenatus ) adhesive proteins

Cements from two species of barnacles, Balanuseburneus Gould and Balanus crenatus Bruguiére, were analyzed to identify the number and nature of proteins present. B. crenatus cement was composed mainly of a small peptide cross-linked into aggregates ranging in size from ˜ 3 kD to 40–50 kD. These aggregates could be reduced with 2-mercaptoethanol (2-ME) in the presence of sodium dodecyl sulfate (SDS) but only after incubation at 40 °C for 12 to 24 h and only when the cement was recently formed. Reductive alkylation of cysteine residues with 4-vinylpyridine produced a water-soluble peptide of less than 5 kD. By comparison, the cement of B. eburneus could be dissolved only partially in SDS and 2-ME when heated at 100 °C for 10 min. Five major proteins were identified by SDS-PAGE: 7, 22, 36 and 58 kD bands for which N-terminal sequence and amino acid compositions are presented; and a 52 kD band for which sequence data are given. A minor protein band of ˜ 80 kD has the same N-terminus as the 36 kD band. CNBr digests of individual proteins produced peptides for which sequence and composition data are also presented. The study was conducted during 1993 to 1995. In general, the proteins identified from B. crenatus cement are similar to those characterized from B. eburneus, and they are different in composition and sequence than those previously reported from Mytilus edulis. Received: 7 August 1996 / Accepted: 8 October 1996     

Low molecular-weight factor from Plesiastrea versipora (Scleractinia) that modifies release and glycerol metabolism of isolated symbiotic algae

When symbiotic dinoflagellate algae (Symbiodinium sp., isolated from the coral Plesiastrea versipora) were incubated with NaH¹⁴CO₃ in the light in seawater, they released 22.69±9.16 nmol carbon/10⁶ algae. Release of photosynthetically fixed carbon was stimulated more than six-fold for algae incubated in host-tissue homogenate (148.54±97.03 nmol C/10⁶ algae) and more than four-fold (102.00±49.16 nmol C/10⁶ algae) for algae incubated in a low molecular weight fraction (≤1 000 M _r ) prepared from host homogenate. Soluble released ¹⁴C-labelled products, as determined by chromatography and autoradiography, were the same when algae were incubated in either host homogenate or the low molecular weight fraction. After 4 h incubation in the light (300 mol photons m⁻² s⁻¹),␣intracellular␣glycerol increased in algae incubated with the low molecular weight fraction (an increase of 0.39 to␣0.67 nmol glycerol/10⁶ algae) compared with little or no increase in algae incubated in seawater (0 to 0.12 nmol glycerol/10⁶ algae). Partial inhibition of triglyceride synthesis (up to 51%) was also observed when algae were incubated in the low molecular weight fraction. All these effects are the same as those observed when algae were incubated in host homogenate. These data indicate that the “host release-factor” activity of P.␣versipora is a compound of low molecular weight. Received: 13 February 1997 / Accepted: 24 October 1997     

Sediment processing in Capitella spp. (Polychaeta: Capitellidae): strain-specific differences and effects of the organic toxicant fluoranthene

The effects of the widespread polycyclic aromatic hydrocarbon (PAH) fluoranthene on sediment-processing rates in the infaunal polychaete Capitella spp. were investigated by comparing five populations of this deposit-feeding species complex: Capitella sp. I from New York, USA; Capitella sp. M from Milos, Greece; Capitella sp. S from Sylt, Germany; and two unidentified Capitella populations from salmon farm sediments –Capitella population K from Kilmelford, Scotland and Capitella population C from Cranford, Ireland. Replicate worms from each strain were exposed to 0, 10, and 95 μg (g dry wt sediment)⁻¹ fluoranthene (=μg/g fluoranthene) for a period of 16 days. Initial and final wet and dry weights (mg) of worms and worm-specific growth rates (WSGRs) were calculated. Sediment processing was measured as the sum of the total dry weight of pellets produced during the experiment, and we estimated size-specific processing rates (SSPRs) as a measure of sediment processed per mg worm dry weight per day. The five populations of Capitella spp. differed significantly in body size, WSGR, and sediment-processing rates. Capitella sp. I grew faster than all of the other populations. Capitella population C and Capitella population K from fish farm sediments, with the largest body lengths (up to 52.0 ± 27.2 mm), had the highest processing rates, whereas the small Capitella sp. S (up to 17.1 ± 5.6 mm) had the lowest. There were also significant differences in SSPR among populations with Capitella sp. I having a higher SSPR (about 12 × body wt/day) than Capitella population C (about 5 × body wt/day) and Capitella population K (3 × body wt/day). The fluoranthene concentrations used in the present study, while representing moderately to highly contaminated conditions, had only marginal effects on sediment-processing and growth rates of all of the Capitella populations examined. Processing of contaminated sediment by Capitella spp. may be important in the remediation of PAH-contaminated sediment. Received: 16 January 2000 / Accepted: 28 August 2000     

Zooplankton growth rates: the influence of size in nauplii of tropical marine copepods

Growth rates and development times were determined for nauplii of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total naupliar development time was short: 3 to 4 d inshore and 4 to 5 d offshore. Mean instantaneous growth rates (g) ranged from as high as 0.90 d⁻¹ for Parvocalanuscrassirostris to as low as 0.41 d⁻¹ for Corycaeus spp. In general, nauplii of cyclopoid copepods appeared to grow more slowly than those of calanoids of the same size. Naupliar growth rates were significantly related to body size (r ²= 0.43 to 0.50), but were unrelated to chlorophyll concentration in any measured size-fraction. This suggests that nauplii are generally not limited by resources, but are growing at their maximum temperature and size-dependent rates. Received: 30 May 1997 / Accepted: 13 May 1998     

Impact of the polychaete Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon fluoranthene

Polychaetes belonging to the genus Capitella are often present in high numbers in organic-rich sediments polluted with, e.g., oil components, and Capitella spp. may have a great impact on the biogeochemistry of these sediments. We examined the influence of Capitella sp. I on microbial activity in an organic-rich marine sediment contaminated with the polycyclic aromatic hydrocarbon, fluoranthene. Capitella sp. I were added to microcosms (10 000 ind m⁻²) and the impact of a pulse-sedimentation of fluoranthene-contaminated sediment (3 mm layer) was studied for a period of 12 d after sedimentation. The sediment oxygen uptake and total sediment metabolism (TCO₂ production) increased in cores with worms (71 to 131%), whereas the anaerobic activity, measured as sulfate reduction rate 12 d after sedimentation, was lower compared to cores without worms. The effect of fluoranthene on sulfate reduction was most pronounced in the presence of worms, with a 34% reduction versus 16% in cores without worms. The reduced sulfur pools in cores with worms were smaller than in cores without worms, suggesting that the reduced anaerobic activity was caused by increased oxidation of the sediment, which may favor O₂ and other electron-acceptors (e.g. NO₃ ⁻, Fe³⁺, Mn⁴⁺) in organic matter decomposition. The sediment oxygen uptake and TCO₂ production did not show significant changes due to fluoranthene treatment, indicating that these parameters were either less sensitive to fluoranthene stress or recovered more rapidly (i.e. within 48 h) than sulfate reduction rates. Bioturbation by Capitella sp. I altered the depth profile of fluoranthene such that fluoranthene was found in deeper sediment layers (down to 2 cm) where diffusional loss and microbial breakdown probably are reduced relative to surface layers. In cores without worms, fluoranthene was found down to 1 cm, with 75% remaining in the upper 5 mm. Received: 5 December 1996 / Accepted: 11 February 1997     

Effect of diet on growth rate and reproductive fitness of Turbo sarmaticus (Mollusca: Vetigastropoda: Turbinidae)

Experiments were conducted to determine the effect of four algal diets (Corallina spp., Gelidium pristoides, Ulva rigida and an equal volume mixture of these seaweeds) on growth and reproduction of the South African gastropod Turbo sarmaticus Linnaeus, 1758. The best growth rate of juveniles (up to 13.8 mm shell length and 34.26 g wet body weight increase in 12 months), reproductive fitness of mature specimens (gonad index up to 33%) and energy reserves (up to 4.76 mg glycogen/100 mg foot tissue) were achieved when T. sarmaticus was fed U. rigida or a mixed diet. In addition, the gonad index of individuals fed these diets was almost twice that of similar-sized field specimens. Juveniles fed Corallina spp. only, grew very little (only 2.4 mm shell length and 4.23 g wet body weight increase in 12 months). The reproductive fitness of adults fed on such a diet was also poor (gonad index <4.5%) and energy reserves were low (<3.5 mg glycogen/100 mg foot tissue). Along the southeastern coast of South Africa, T. sarmaticus has a distinct reproductive cycle with gametogenesis occurring from March/April until August/September, whilst maturity (gonad index = 15%) was maintained until December, after which spawning occurred until March. Received: 6 July 1998 / Accepted: 8 March 1999     

Zooplankton responses to hypoxia: behavioral patterns and survival of three species of calanoid copepods

Seasonally recurrent and persistent hypoxic events in semi-enclosed coastal waters are characterized by bottom-water dissolved oxygen (d.o.) concentrations of < 2.0 ml l⁻¹. Shifts in the distribution patterns of zooplankters in association with these events have been documented, but the mechanisms responsible for these shifts have not been investigated. This study assessed interspecific differences in responses to hypoxia by several species of calanoid copepods common off Turkey Point, Florida, USA: Labidocera aestiva (Wheeler) (a summer/fall species), Acartia tonsa (Dana) (a ubiquitous year-round species), and Centropages hamatus (Lilljeborg) (a winter/spring species). Under conditions of moderate to severe hypoxia 24-h survival experiments were conducted for adults and nauplii of these species from August 1994 to October 1995. Experiments on adults used a flow-through system to maintain constant d.o. concentrations. Adults of A. tonsa showed no decline in survival with d.o. as low as 1.0 ml l⁻¹, sharp declines in survival at d.o. = 0.9 to 0.6 ml l⁻¹, and 100% mortality with d.o. = 0.5 ml l⁻¹. Adults of L. aestiva and C. hamatus were more sensitive to oxygen depletion: both species experienced significant decreases in survival for d.o. = 1.0 ml l⁻¹. Nauplii of L. aestiva and A. tonsa showed no significant mortality with d.o. = 1.1 to 1.5 ml␣l⁻¹ and d.o. = 0.24 to 0.5 ml l⁻¹, respectively. In addition, experiments investigating behavioral avoidance of moderate to severe hypoxia were carried out for adults of all three species. None of the three species effectively avoided either severely hypoxic (d.o. < 0.5 ml l⁻¹) or moderately hypoxic (d.o. ≈ 1.0 ml l⁻¹) bottom layers in stratified columns. These results suggest that in␣nearshore areas where development of zones of d.o. < 1.0 ml l⁻¹ may be sudden, widespread, or unpredictable, patterns of reduced copepod abundance in bottom waters may be due primarily to mortality rather than avoidance. Received: 31 August 1996 / Accepted: 24 September 1996     

Kinetic measurements of metal accumulation in two marine macroalgae

 We measured the uptake kinetics of four metals (Cd, Cr, Se and Zn) in two marine macroalgae (the green alga Ulva lactuca and the red alga Gracilaria blodgettii). Metal uptake generally displayed a linear pattern with increasing exposure time. With the exception of Cr, which exhibited comparable uptake rate constants at different concentrations, uptake rate constants of Cd, Se and Zn decreased with increasing metal concentration, indicating that the seaweeds had a higher relative uptake at lower metal concentration. Uptake of Cd and Zn was higher in U. lactuca than in G. blodgettii, whereas uptake of Cr and Se was comparable between the two species. Only Cd and Zn uptake in U. lactuca was significantly inhibited by dark exposure. A decrease in salinity from 28 to 10‰ enhanced the uptake of Cd, Cr, Se and Zn in U. lactuca 1.9-, 3.0-, 3.6-, and 1.9-fold, respectively. In G. blodgettii, Cd uptake increased twofold when salinity was decreased from 28 to 10‰, whereas uptake of Cr and Zn was not significantly affected by salinity change. The calculated depuration rate constants of metals in U. lactuca were 0.01 d⁻¹ for Cd, 0.05 to 0.08 d⁻¹ for Cr, 0.14 to 0.16 d⁻¹ for Se, and 0.12 to 0.15 d⁻¹ for Zn, and were relatively independent of the metal body burden in the algae. The predicted bioconcentration factor was 3 × 10⁴ for Cd, 2 × 10³ for Cr, 40 to 150 for Se, and 1 to 2 × 10⁴ for Zn in U. lactuca. Our kinetic study suggested that U. lactuca would be a good biomonitor of Cr and Zn contamination in coastal waters. Received: 14 September 1998 / Accepted: 29 May 1999     

Seasonal abundance and toxicity of the dinoflagellate Gambierdiscus spp. (Dinophyceae), the causative agent of ciguatera in Tahiti, French Polynesia

The benthic dinoflagellate Gambierdiscus spp. is the primary causative agent of ciguatera fish poisoning in French Polynesia. However, the initiation of ciguatera outbreaks and the factors that control ciguatoxin production in this dinoflagellate are still poorly understood. In this paper, we present some original data concerning the seasonal abundance and toxicity of natural populations of Gambierdiscus spp. based on a long-term survey in a ciguateric site of the island of Tahiti. From February 1993 to December 1997, Gambierdiscus spp. population densities were monitored weekly in the Atimaono barrier reef of Papara in relation to temperature and salinity. Densities peaked at 4992 cells g⁻¹ in October 1994 and constituted ≥1000 cells g⁻¹ on several occasions. A total of 58 blooms were recorded in the area, of which 65% occurred in 1995 and 1996 alone. Our data confirmed the high endemicity of Gambierdiscus spp. in the Papara area. Refined analysis of our raw data by a principal-component analysis of processes (PCAP) revealed a seasonality in cell densities from February 1993 to May 1995. During this period, Gambierdiscus spp. populations tended to reach maximum abundance at the beginning and end of the hot season. In contrast, salinity did not appear to be a determining factor in the seasonal abundance of this dinoflagellate. The noticeable increase in both peak densities and frequency of blooms further noticed in 1995 and 1996 was preceded by unusually high water temperatures in January to April 1994, concomitant with a severe coral-bleaching episode. During the course of the study, a total of 303 × 10⁶ cells obtained from 48 successive blooms was harvested. Toxicity screening revealed that toxin production was maximum from October 1994 through December 1996. No correlation was found between toxicity of these blooms and their biomass nor the seasonal pattern of temperatures. It is suggested that the toxicity of naturally-occurring blooms of Gambierdiscus spp. and, consequently, the severity of ciguatera incidents in a given area, is mainly dependent on the clonal nature of cells which coexist within local populations of this dinoflagellate. Received: 12 October 1998 / Accepted: 15 June 1999     

Megabenthic decapod crustacean assemblages on the Galician continental shelf and upper slope (north-west Spain)

The structure of megabenthic decapod crustacean assemblages on the Galician (north-west Spain) continental shelf (100 to 200 depth) and upper slope (200 to 500 m) was analyzed based on surveys carried out in autumn and spring, from 1980 to 1987. Forty species belonging to 19 families were caught. The portunid crab Polybius henslowii, a species with pelagic phases, was the most abundant species, but displayed strong spatial and temporal fluctuations. Other dominant species were the Norway lobster Nephrops norvegicus, the portunid Liocarcinus depurator, the galatheids Munida intermedia and M. sarsi, and the shrimps Solenocera membranacea, Plesionika heterocarpus, Pasiphaea sivado and Dichelopandalus bonnieri. Total abundance and biomass (average values excluding Polybius henslowii = 255 individuals and 2.06 kg/30 min tow) and species richness and diversity, H′ (6.85 species and H′ = 1.45 per tow) displayed a significant positive correlation with depth, and strong interannual fluctuations. The factors determining community organization were depth and, to a lesser extent, spatial structure. There was clear evidence of bathymetric zonation, differentiating between species characteristic of the slope (D. bonnieri and Pasiphaea sivado), shelf-slope edge (Macropipus tuberculatus, Pontophilus spinosus, Munida sarsi, S. membranacea, Processa spp.) and shelf (L. depurator, Macropodia tenuirostris, Paguridae and Chlorotocus crassicornis). The spatial zonation was related to changes in oceanography and sediment along the continental margin. Goneplax rhomboides, N. norvegicus, C. crassicornis and Alpheus glaber are benthic species which generally exhibit burrowing behaviour, and they were found mainly in the southern area where there are fine sediments due to the outwelling from the Rías Baixas. Different benthopelagic shrimps (Pontophilus spinosus, Plesionika heterocarpus,Processa spp. and Pasiphaea sivado) were typical of the zone just north of Fisterra, characterized by a convergence of water masses bringing about an increase in productivity due to upwelling. The benthic anomuran and brachyuran crabs Munida intermedia, M.␣sarsi, L.␣depurator and Macropipus tuberculatus were characteristic of the northwestern zone between Fisterra and Estaca, where the infauna reaches high biomass despite coarser sediments with a lower concentration of organic material than in the southern area. Lastly, both the Paguridae and Macropodia tenuirostris were species typically found in the waters in the northern shelf. Based on interannual changes in assemblage structure, two periods could be distinguished: between 1980 and 1984, when Polybius henslowii, D. bonnieri and Pasiphaea sivado had abundance peaks; and another period from 1985 to 1987 when L. depurator, Munida intermedia, M.␣sarsi and Macropipus tuberculatus increased in abundance. Received: 21 May 1996 / Accepted: 9 August 1996     

“Seed bank” of coastal planktonic diatoms in bottom sediments of Hiroshima Bay, Seto Inland Sea, Japan

Abundance and temporal distribution of viable (able to germinate) resting stage cells of planktonic diatoms in bottom sediments have been investigated almost monthly during 1989 to 1992 in Hiroshima Bay, western part of Seto Inland Sea, Japan. The abundance of viable resting stages in bottom sediments was enumerated with the extinction dilution method (most probable number method, MPN). In bottom sediments of Hiroshima Bay, dominantly distributed species and/ or genera of the diatom resting stages were Skeletonema costatum, Chaetoceros spp. and Thalassiosira spp. Viable resting stages of these diatoms were densely distributed on the orders of 10³ to 10⁶ (MPN g⁻¹ wet sediments), and persisted in bottom sediments throughout the investigation period. Conversely, vegetative cells of these diatoms fluctuated remarkably in the water column and disappeared sporadically. Survival of the resting stages in a collected sediment sample was also determined with the MPN method, at different storage temperatures (5, 10, 15, 20, 25 °C). The survival test demonstrated that the diatom resting stages could survive in the dark for several months or years in sediments. Resting stages survived longer at the lower storage temperature, and the order of longevity was consistent within three diatoms (Chaetoceros spp. > Thalassiosira spp. > S. costatum) at each storage temperature. The present study suggests that these diatom resting stages in the coastal bottom sediments could serve as a “seed bank”, analogous to those of terrestrial plants. The seed bank would ensure the survival of diatoms within highly fluctuating coastal environments, while it would also be the source of sporadic and autochthonous diatom blooms in coastal waters. Received: 29 November 1996 / Accepted: 16 December 1996     

Cytochrome c used as a model system to probe the structural properties of benthic mucilage

Benthic mucilage, whether native or artificially fragmented to microscopic dimensions by ultrasonic treatment, was mixed with cytochrome c, which was used as a 12 000 Da polycationic model compound. Cytochrome c binding profiles proved to depend on the aggregation state of the mucilage. The native mucus matrix binds cytochrome c quasi hyperbolically with an apparent affinity constant K = 1 × 10⁶  M ⁻¹. As shown by chemical modification of both the mucilage and cytochrome c, the binding expression is dependent on the availability of both the positive charges on the cytochrome c surface and the negative charges within the mucus matrix. The extent of binding is sensitive to the ionic strength of the medium. The ultrasonic-stabilized mucus fragments display a peculiar binding profile, with an apparent low affinity, abruptly entering into a high affinity binding region. The results suggest that, depending on the mucus to polycation ratio, a polymeric reticulus builds up. This reticulus can accommodate molecules of at least 12 000 Da molecular weight. The results are also discussed with respect to biological implications. Received: 23 April 1997 / Accepted: 24 November 1997     

Essential amino acid synthesis and nitrogen recycling in an alga–invertebrate symbiosis

When aseptically-cultured sea anemones, Aiptasia pulchella, were incubated with ¹⁴C-labelled glucose, aspartate and glutamate, radioactivity was incorporated into animal protein. Radioactivity was recovered from all amino acids in the protein hydrolysates of A. pulchella bearing the symbiotic alga Symbiodinium sp., and from all but seven of the amino acids in A. pulchella experimentally deprived of their algae. These data suggest that these seven amino acids (histidine, isoleucine, leucine, lysine, phenylalanine, tyrosine and valine) may be synthesized by the symbiotic algae and translocated to the sea anemone's tissues; and that methionine and threonine, two amino acids traditionally considered as dietary essentials for animals, are synthesized by A. pulchella. Essential amino acid translocation from the symbiotic algae to the animal host is a core element in symbiotic nitrogen-recycling. Its nutritional value to the animal host is considered in the context of the amino acid biosynthetic capacity of the host. Received: 26 October 1998 / Accepted: 28 June 1999