Binding of manganese by Antarctic Phaeocystis pouchetii and the role of bacteria in its release

The growth of Antarctic Phaeocystis pouchetii and associated bacteria in culture, and the binding and release of manganese were investigated using cultures derived from three clonal isolates collected from Prydz Bay, Antarctica, in November and December 1982. The cultured strains accumulated manganese from the culture medium. The concentration of Mn²⁺ in the alga was up to 58 times that of the medium. The Mn²⁺ apparently binds to the mucilage secreted by the cells and gives the alga its characteristic brown colour. Oxidation or complexing of Mn²⁺ by P. pouchetii was also indicated. Bacterial growth on the mucilage is inhibited in rapidly growing cultures by acrylic acid produced in the mucilage. Once the growth of P. pouchetii and production of ccrylic acid slow, bacterial numbers increase, leading to the solubilization of the mucilage and the release of Mn²⁺. In cool-temperate, subpolar and polar seas where P. pouchetii may form dense blooms, the binding and subsequent release of manganese may have important consequences for competing phytoplankton.     

Dissolved organic substances during a Phaeocystis pouchetii bloom in the German Bight (North Sea)

Dissolved organic substances (carbohydrates, organic nitrogen, free amino acids) were measured in the German Bight (North Sea) in June, 1981. During and before this survey, sea foam was observed in the east Frisian coastal water and it accumulated on the nearby beaches to an unusually high extent. In this coastal water area, a large Phaeocystis pouchetii Lagerheim bloom and very high concentrations of dissolved organic matter were found. The above dissolved organic substances were all positively correlated to a highly significant degree with P. pouchetii cell numbers in the bloom area. An influence of salinity (or river water) on this correlation could be excluded. Thus, exudation or decomposition products of P. pouchetii were most likely the cause of the unusually high concentrations of dissolved organic matter in the bloom area off the east Frisian coast, where P. pouchetii blooms have been reported for many years. Ammonia concentrations were very low in the P. pouchetii bloom area; this and the accumulation of dissolved organic substances might lead to speculation that decomposition of dissolved organic matter by bacteria could have been reduced due to antibiotic activity of P. pouchetii.     

Impact of UV-B radiation on uptake of 15N-ammonia and 15N-nitrate by phytoplankton of the Wadden Sea

Natural marine phytoplankton populations from the German Wadden Sea and unialgal cultures of the haptophycean Phaeocystis pouchetii were tested in 1989 under controlled UV-B stress conditions. Assimilation of ¹⁵N-nitrate in phytoplankton consisting mainly of P. pouchetii, or in pure cultures of this alga, was found to be very sensitive to enhanced UV-B dosage in comparison ¹⁵N-ammonia uptake. In contrast, in phytoplankton samples containing Ceratium spp., Coscinodiscus sp., Noctiluca sp. or others, rate of ¹⁵NO₃ ^- uptake was higher and only slightly affected by UV-B irradiance compared to the P. pouchetii sample. UV-B inhibitory effect on uptake of inorganic nitrogen by P. pouchetii was more pronounced under strong white-light conditions and after a UV-B pre-illumination period of several hours than under low white light. Pools of glutamine and alanine decreased after UV-B exposure. Results are discussed with reference to the damaging effects of white light and UV-B on nitrogen metabolism.     

Nutritional value of Phaeocystis pouchetii (Prymnesiophyceae) and other phytoplankton for Acartia spp. (Copepoda): ingestion,egg production,and growth of nauplii

Ingestion and egg production by Acartia hudsonica (5°C) and A. tonsa (10°C) from Narragansett Bay, Rhode Island, USA, was measured in the laboratory over a range of concentrations of the prymnesiophyte Phaeocystis pouchetii and the diatom Skeletonema costatum. Both Acartia species reproduced well when fed unialgal diets of S. costatum. Egg production by females fed P. pouchetii, either as gelatioous colonies (primarily >200 m in diameter) or solitary cells (3 to 5 m), was not significantly different than that of starved females. Acartia spp. fed selectively on S. costatum in multialgal treatments. The presence of P. pouchetii did not reduce ingestion of S. costatum; the availability of S. costatum did not increase ingestion of P. pouchetii. Nauplii of A. hudsonica grew equally well on separate diets of P. pouchetii and Isochrysis galbana, two flagellates of similar size. Adult Acartia spp. reproduced poorly when fed these phytoplankton, suggesting that particle size may be more important than food quality in describing the responses. Grazing by Acartia spp. does not directly impact the dynamics of P. pouchetii, but may indirectly contribute to blooms of this prymnesiophyte by removal of competing phytoplankton.     

Two new UV-absorbing mycosporine-like amino acids from the sea anemoneAnthopleura elegantissima and the effects of zooxanthellae and spectral irradiance on chemical composition and content

Many tropical cnidarians living in shallow water contain a class of ultraviolet-A (UV-A, 320 to 400 nm) and ultraviolet-B (UV-B, 280 to 320 nm) absorbing compounds known as mycosporine-like amino acids (MAAs). These compounds may provide protection from the deleterious effects of solar UV radiation. Using a novel application of reverse-phase high performance liquid chromatography, we find that the temperate sea anemoneAnthopleura elegantissima (collected in 1988 from Bodega Bay, California, and in 1991 from Santa Barbara, California) contains four major MAAs: shinorine, porphyra-334, and two new compounds, mycosporine-taurine and mycosporine-2 glycine. Analysis of zooxanthellate (containing zooxanthellae) and naturally apozooxanthellate (lacking zooxanthellae) specimens acclimated in the presence and absence of UV for 28 d in the spring of 1988 suggests that this anemone, unlike some other anthozoans, does not regulate the concentration of its MAAs in response to UV radiation. The presence of similar concentration of MAAs in apozooxanthellate and zooxanthellate specimens indicates that symbiosis with algae is not required for these compounds to be present in the anemone. The total concentration of MAAs in the zooxanthellae is only about 12% of that in their host's tissues.     

Effects of UV-B irradiation on growth and survival of Antarctic marine diatoms

Growth rate, survival, and stimulation of the production of UV-B (280 to 320 nm) absorbing compounds were investigated in cultures of five commonly occurring Antarctic marine diatoms exposed to a range of UV-B irradiances. Experimental UV-B exposures ranged from 20 to 650% of the measured peak surface irradiance at an Antarctic coastal site (0.533 J m^-2 s^-1). The five diatom species (Nitzschia lecointei, Proboscia alata, P. inermis, Thalassiosira tumida and Stellarima microtrias) appear capable of surviving two to four times this irradiance. In contrast to Phaeocystis cf. pouchetii, another major component of the Antarctic phytoplankton, the concentrations of pigments with discrete UV absorption peaks in diatoms were low and did not change significantly under increasing UV-B irradiance. Absorbance of UV-B by cells from which pigments had been extracted commonly greatly exceeded that of the pigments themselves. Most of this absorbance was due to oxidisable cell contents, with the frustule providing the remainder. Survival of diatoms did not correlate with absorption by either pigments, frustules or oxidisable cell contents, indicating that their survival under elevated UV-B irradiances results from processes other than screening mechanisms.     

Phaeocystis pouchetii blooms in the East Frisian coastal waters (German Bight,North Sea)

In 1982 and 1983, blooms of the planktonic alga Phaeocystis pouchetii were studied in the East Frisian coastal waters off Norderney, FRG. Morphological variability of the colonies, population development, effects on inorganic nutrient and oxygen budget, and the role in the seasonal plankton bloom sequence were investigated. in both years two separate Ph. pouchetii blooms occurred in late spring and early summer that were formed by the globosa-type exclusively. The pouchetii-type was only present during the interval between the first and second blooms in 1982. Growth conditions for Ph. pouchetii seem to be most favourable after a breakdown of a diatom bloom when the silicate reserves have been exhausted, phosphate concentrations are relatively low, but a rich supply of dissolved inorganic nitrogen is available. Contrary to diatom blooms, Ph. pouchetii blooms result in a high production of particulate organic carbon (POC). Another characteristic is the release of great amounts of dissolved organic substances, which cause extensive sea foam formation during the peaks and the subsequent breakdown of the blooms. Due to increased assimilation during mass development and intense disintegration after the breakdown, variations of the oxygen content of the water are significant. Presumably the intensity of blooms has increased in the past decade as indicated by sea foam formation which has actually been excessive in some years.     

Daily and seasonal variations of optimum quantum yield and phenolic compounds in Cystoseira tamariscifolia (Phaeophyta)

Effects of solar radiation on phenolic compound concentrations and photosynthetic activity, estimated as in vivo chlorophyll fluorescence, in the brown alga Cystoseira tamariscifolia (Hudson) Papenfuss were analyzed in southern Spain from June 1988 to July 2000. Annual and diurnal variations of optimum quantum yield were negatively correlated with incident irradiance. Optimum quantum yield decreased as irradiance increased at noon, and yield values recovered in the afternoon suggesting a dynamic photoinhibition. The annual and daily fluctuations of phenolic compound concentration in the tissue of C. tamariscifolia showed contrasting patterns. There was an annual cycle of phenolic compound concentration in the apical thallus, which was positively correlated with incident irradiance. The increase in phenolic compounds, however, was twofold greater in the first half of the year than the decrease during the second half of the year. In contrast to the annual cycle, there appeared to be a negative correlation between phenolic compound concentration and irradiance in the summer months while no specific relationship was observed in the fall–winter months. Loss of phenolic compounds from the tissue to the surrounding water was increased as irradiation dosage increased. This suggests that the decrease of phenolic compounds during the diurnal cycle might be regulated by the exudation of these compounds at high irradiances in the field. Collectively, our results suggest that, like dynamic photoinhibition, the rapid synthesis and turnover time of phenolic compounds in the tissue of C. tamariscifolia might serve as photoprotective mechanisms against high irradiances.     

A study of mucus from the solitary coral Fungia fungites (Scleractinia: Fungiidae) in relation to photobiological UV adaptation

The ultraviolet (UV)-absorbance spectrum (300 to 360 nm) of mucus obtained from Fungia fungites (L. 1758, collected in Tahiti in 1991) after being exposed to air for up to 5 min was measured, and UV-absorbing compounds were demonstrated to be present in the mucus, with a peak at 332 nm. The concentration of these UV-absorbing compounds was at a maximum in the first 2 min of secretion and decreased thereafter. Concentration was significantly related to the weight of the coral. Also, as corals were adapted to bathymetric levels of UV radiation, mucus concentration of UV-absorbing compounds decreased significantly with increasing depth.     

Light harvesting in the green alga Ostreobium sp., a coral symbiont adapted to extreme shade

Ostreobium sp. (Chlorophyta: Siphonales) can be found as green bands within the skeletal material of a number of stony corals in the Indo-Pacific and Caribbean regions. Many of these corals also contain symbiotic dinoflagellates in the overlaying coral polyps that effectively screen out all the typical photosynthetically active radiation from the algae in the green bands below. Ostreobium sp., nevertheless, grows photosynthetically. Its action spectrum and absorption spectrum have been shown to extend much further into the near infra-red compared to other green algae. In the present study, carried out in 1987, fluorescence excitation and emission spectra were measured in Ostreobium sp. and compared to spectra obtained from the green alga Ulva sp. and the brown alga Endarachne sp. Xanthophylls, probably siphonein and an unidentified xanthophyll probably related to siphonaxanthin, are photosynthetically active in Ostreobium sp., and can sensitize Photosystem II fluorescence at 688 nm and Photosystem I (PS I) fluorescence at 718 nm. The fluorescence emission spectra of Ostreobium sp. measured at 25° C and 77 K were not remarkably different from those of the green alga Ulva sp. Absorbance changes induced by light were measured in Ostreobium sp. from 670 to 750 nm and were like those normally seen in green plants except that, in addition to the minimum expected for the reaction-center chlorophyll of PS I (P700) at 703 nm, another minimum was seen at 730 nm. It is possible that this spectrumreflects the functioning of a reaction center of Photosystem I that has adapted to function in light highly enriched in far-red wavelengths.CIW-DPB Publication No. 1021     

Large viruses and infected microeukaryotes in Ross Sea summer pack ice habitats

A variety of Ross Sea summer pack ice habitats between 66 and 75°S were examined for viruses 𔒦 nm capsid diameter. Maximum abundances of these viruses likely to infect eukaryotes were 10⁶-10⁷ ml^-1 brine in surface, interior, and bottom habitats and constituted up to 18% of the total (all sizes) viruses. There is abundant ultrastructural evidence for infection of a variety of microheterotrophs and some autotrophs. One station exhibited the classical characteristics of a lower latitude algal bloom with potential viral control. The blooming alga, Pyramimonas tychotreta Daugbjerg 2000, was infected, as were two abundant heterotrophs, Cryothecomonas spp. and an unidentified flagellate, that fed on P. tychotreta. Infections were observed in only one life history stage (multiflagellate cells) of P. tychotreta, suggesting a relationship among virus-induced lysis, life-history stages, physiology, and environmental factors regulating the life cycle. There is good evidence that diatoms are not a likely source of the large viruses, and viruses in general are not a major food source for ice microheterotrophs in summer.     

Mycosporine-like amino acids: possible UV protection in eggs of the sea hare Aplysia dactylomela

We investigated mycosporine amino acid (MAA) involvement as protective sunscreens in spawn of the sea hare Aplysia dactylomela to determine if adult diet and ultraviolet (UV) exposure affected the UV sensitivity of developing embryos. Adults were fed a red alga rich in MAAs (Acanthophora spicifera) or a green alga poor in MAAs (Ulva lactuca). Adults on each diet were exposed for 2 wk to ambient solar irradiance with two types of acrylic filters; one allowed exposure to wavelengths >275 nm (designated UV) and one to wavelengths only >410 nm (designated NOUV). Spawn from each adult group was likewise treated with UV or NOUV and monitored during development for differences in mortality and metabolic rate (measured as oxygen consumption: V˙ _O2). Also recorded were number of eggs or embryos per capsule, times to hatching, hatching success, size at hatching, and V˙ _O2 of adults. Spawn from adults eating red algae was almost twice as rich in MAAs as spawn from adults eating green algae, suggesting that MAA content is diet-related. Although overall quantities of MAAs in the spawn reflected MAA contents of the adult diet, specific MAAs were differentially sequestered in the spawn. Thus, porphyra-334, found in high concentration in Aplysia dactylomela's preferred red algal food, was present in only low concentration in the spawn. Conversely, mycosporine-glycine, in low concentration in red algal food, was the most abundant MAA in the spawn. UV treatment of adults had no effect on quantities of MAAs in the spawn. Adults exposed to UV had significantly higher V˙ _O2s and spawned twice as often. The UV-treated adults produced spawn with significantly higher V˙ _O2s and their embryos developed to hatching sooner. The only significant effect of UV exposure of the spawn was to reduce the percentage of veligers hatching from 71 to 50%. There was no significant effect on hatching time or size of the veligers at hatching, nor on number of eggs per capsule. Received: 13 May 1997 / Accepted: 27 June 1997     

Carbon assimilation and lipid production in phytoplankton in northern Norwegian fjords

Carbon assimilation and lipid production were studied in phytoplankton in Balsfjorden and Ullsfjorden, northern Norway, during the exponential growth phase of the spring bloom in 1983 (6–7 April). In Balsfjorden, phytoplankton biomass was constant with depth and equivalent to 1.5 g chlorophyll a 1^-1. Phytoplankton biomass in Ullsfjorden varied with depth, with a maximum of ca. 7 g chlorophyll a 1^-1 occurring at 5 to 10 m. Particulate carbon-14 assimilation was about 18 mg C per m^-2 h^-1 in Balsfjorden and about 39 mg C per m^-2 h^-1 in Ullsfjorden over the depth range 4 to 8 m. In Balsfjorden, the percentage of total fixed carbon recovered as total lipid was 14.7 and 20.4% at 4 and 8 m depth, respectively. In Ullsfjorden, the corresponding values were 8.8 and 28.1% at 4 and 8 m, respectively. The percentages of total fixed carbon present as fatty acids were 1.1 and 1.6% at 4 and 8 m, respectively, in Balsfjorden, and 0.8 and 6.4% at 4 and 8 m in Ullsfjorden. The majority of the radioactivity in lipid at both locations and at both depths was present as polar lipid, with small percentages present in triacylglycerols and very small percentages present in free fatty acids. On average, about 18% of the total carbon-14 incorporated into phytoplankton over a 6 to 7 h mid-day period was recovered as total lipid and its percentage tended to increase with depth. The relatively low percentage of incorporated carbon-14 present as fatty acids in total lipid implies that most of the radioactivity is present in glyceryl and/or glucosyl moieties and that measurement of total radioactivity in total lipid does not necessarily give an accurate estimation of lipogenesis in phytoplankton. Fatty acid analyses of total phytoplankton in Balsfjorden and Ullsfjorden in 1983 and of a surface slick at the end of a bloom of Phaeocystis pouchetii in Balsfjorden in May 1980 showed an abundance (more than 40% of the total) of (n-3) polyunsaturates in all cases. C-18 polyunsaturates, especially 18:4 and 18:5, were very abundant (about 30% of the total) in the P. pouchetii surface slick in Balsfjorden in 1980. Both P. pouchetii biomass and C-18 polyunsaturates were more abundant in Ullsfjorden than in Balsfjorden (1983). Lipids from the P. pouchetii surface slick were deficient in C-16 polyunsaturates and relatively deficient in C-20 polyunsaturates, but both these classes of fatty acids were abundant in Balsfjorden and Ullsfjorden in 1983. The phytoplankton in both locations in 1983 was dominated by P. pouchetii and diatoms; Chaetoceros socialis was especially abundant in Balsfjorden. The results are discussed in terms of the fatty acids present in herbivorous zooplankton in northern Norwegian fjords.     

Bacteria on the surface of crustose coralline algae induce metamorphosis of the crown-of-thorns starfish Acanthaster planci

The crustose coralline alga Lithothamnium pseudosorum induces high rates of settlement and metamorphosis of larvae of the coral-eating crown-of-thorns starfish (Acanthaster planci). In cases where crustose coralline algae (CCA) induce metamorphosis of marine invertebrate larvae it is normally assumed that the inductive molecules are produced by the alga, but an alternative is that they originate from bacteria on the plant surface. Bioassays using shards of L. pseudosorum treated with several antibiotics, whereby some shards were reinfected with bacteria from the alga, showed that if bacteria populations are depleted then settlement and metamorphosis of larvae of A. planci are inhibited. This demonstrates that bacteria are necessary for induction and suggests that morphogenic substances are produced by bacteria on the surface of the alga and not directly by the alga itself. However, surface bacteria are not inductive if they are isolated from soluble algal compounds, suggesting either that they require a substrate from the alga to produce the inductive agents or, alternatively but less likely, that compounds from both the alga and bacteria are required. There is no evidence that inductive compounds derive from the alga, since algal cell debris and soluble extracts prepared from the alga do not induce metamorphosis of A. planci. This is the first time that induction of metamorphosis in a marine invertebrate by CCA has been shown to be mediated by bacteria associated with the alga.     

Effects of diet,ultraviolet exposure,and gender on the ultraviolet absorbance of fish mucus and ocular structures

Ultraviolet (UV) radiation can be damaging to fish skin and ocular components. Coral reef fishes are regularly exposed to potentially harmful radiation. It was recently discovered that tropical marine fishes possess UV-absorbing compounds in their mucus. This experiment demonstrates significant effects of both diet and ultraviolet exposure on the UV-absorbing compounds in the mucus of a tropical wrasse, Thalassoma duperrey. Fish that are exposed to UV radiation increase the UV absorbance of their mucus only if UV-absorbing compounds are provided in their diet. Fish that are protected from UV radiation decrease the UV absorbance of their mucus regardless of diet. Mucus from female T. duperrey absorbed less UV and females had higher rates of skin damage than males. Females sequester UV-absorbing compounds in their pelagic eggs as well as their epithelial mucus, whereas males do not sequester these compounds in the testes. Spectral transmission through the whole eye was not affected by diet or UV manipulations, but corneal tissue transmission decreased significantly in the UV-exposed individuals. These results demonstrate that coral reef fish can adapt to UV exposure, so long as UV-absorbing compounds are available in the diet.Communicated by P.W. Sammarco, Chauvin     

Release of ultraviolet-absorbing compounds by the red-tide dinoflagellateLingulodinium polyedra

We tested the hypothesis that ultraviolet-absorbing compounds known as mycosporine-like amino acids (MAAs) are not only synthesized but also excreted by marine phytoplankton. An experiment was performed with cultures of the marine dinoflagellateLingulodinium polyedra (previously known asGonyaulax polyedra) exposed to visible (photosynthetically available, PAR, 400 to 700 nm) and ultraviolet (UV, 290 to 400 nm) radiation. Absorption properties of both particulate and dissolved organic matter pools (POM and DOM, respectively) showed maxima in ultraviolet absorption at 360 nm. Chromatographic analysis confirmed the presence of MAAs in both pools. Release of organic matter byL. polyedra, as measured spectrophotometrically by changes in UV absorption in the surrounding medium, showed a differential increase at 360 nm in cultures exposed to UV-B + PAR radiation. The changes in absorption in the DOM fraction were inversely proportional to intracellular UV absorption. Photodegradation experiments in which the DOM fraction was exposed to visible and UV-B radiation showed a decrease in absorption with dose. First-order photooxidation decay rates varied between – 0.005 and – 0.26 m² (mol quanta)^–1 and were also a function of the initial optical density (OD). These results indicate that UV-absorbing compounds synthesized by phytoplankton, such as certain dinoflagellates, may be a component of the DOM pool in surface waters of the ocean and contribute to the attenuation of UV radiation in the water column. Photooxidation consumes only 3 to 10% of the daily production of the DOM absorbing between 280 and 390 nm (including MAAs). This suggests that MAAs dissolved in seawater may contribute to the decrease of UV transmission through the water column on a time scale representative of phytoplankton growth (days) and bloom development (weeks).     

High molecular weight algal substances in the sea

Phytoplankton exudates into culture media and extracts from littoral algae are shown to contain significant quantities of extracellular materials having molecular weights in excess of 50,000. A recent development in experimental fluid mechanics (the reduction of turbulent friction coefficients of flowing solutions by dissolved macromolecules) has been utilized as a specific test for high molecular weight compounds. Phytoplankton species in all major algal groups could be found which exude high molecular weight compounds into their culture media; the red alga Porphyridium cruentum typically produced friction reductions as high as 60%. Seaweed samples in the Chlorophyta, Phaeophyta, and Rhodophyta showed major changes in turbulent friction coefficient; in some cases the friction was less than 1/2 that of pure seawater. All samples of the genus Porphyra and Gigartina produced substantial friction changes. From these experiments it is concluded that algal exudates can be a prominent source of high molecular weight compounds in the sea.     

Growth and reproduction of the generalist opisthobranch Haloa japonica: effect of algal seasonality on growth rate

The life history of Haloa japonica (Pilsbry), especially the seasonality of growth and reproduction, was investigated from 1992 to 1995 on a flat rocky shore exposed at low tide at Kattoshi, Hakodate Bay, Japan. To estimate the effect of environmental seasonality on growth rate, we conducted two sets of field cage experiments in which algal food was controlled. H. japonica was shown to have an annual life cycle. Spawning occurred from late April to mid-July. Recruitment occurred in June. Simultaneously, large adults disappeared from the habitat. A rapid increase in wet weight of H. japonica was observed in early spring, and coincided with the bloom of the membranous green alga, Monostroma angicava. Under field cage experiments, individuals feeding on both M. angicava and Polysiphonia japonica grew faster than those feeding on only P. japonica. Growth rates of H. japonica feeding on only P. japonica, however, did not vary from February to April. These results suggest that seasonal variation in the algal community limits the growth of H. japonica in early spring at Kattoshi. This factor is relatively more important as a limiting factor on the growth of H. japonica than the physical environment in this season.     

Reproductive biology of the red alga Polysiphonia lanosa (Ceramiales) in the Bay of Fundy, Canada

The purpose of this investigation was to establish reproductive phenology of the dioecious marine floridean red alga Polysiphonia lanosa (L.) Tandy. Specifically, the distribution and frequencies of male and female plants, timing of their sexual maturity, fecundity and success of fertilization were investigated. The presence of cystocarps was taken as an indication of syngamy. Populations of P. lanosa in the Bay of Fundy reproduce in spring, summer and autumn. Both male and female filaments were present in equal abundance at all sampling locations, grew in close proximity, and occurred with equal abundance at all elevations above chart datum. During the growing season most of the mature plants contained reproductive organs with apparently functional gametangia containing gametes the structure and behavior of which conformed to those reported in the literature for other species of the genus. Male and female plants of P. lanosa matured synchronously during late May to early July at all collecting sites. The ratio of spermatia to carpogonia calculated for the sampled population was high, 3000–4800 spermatia:1 carpogonium. At the end of the growing season (late October) the fertilization success was at its maximum of 50 to 90% and at its minimum in July at 24%. The three free-living life-history phases (females, males and tetrasporophytes) were abundant and occurred with equal frequency, indicating that a complete life cycle was common at the studied location. Received: 10 December 1996 / Accepted: 16 January 1997     

Dynamics of a peculiar plant-herbivore relationship: the photosynthetic ascoglossan Elysia timida and the chlorophycean Acetabularia acetabulum

The ascoglossan mollusc Elysia timida Risso, 1818 retains functional chloroplasts from its algal food, the chlorophycean Acetabularia acetabulum (L.). Photosynthates from the plastids are an important source of organic nutrients for the mollusc. Chloroplast exploitation has an ecological function, allowing the ascoglossan to live entirely on an algal diet which is of limited, seasonal availability to other herbivores. Between October 1987 and July 1988, the annual evolution of the molluscan and algal populations was studied in a cove of Mazarrón Bay, southeast Spain. The population density of the mollusc is highly dependent on its food supply, being controlled by the seasonal life cycle of the algal population. During its life cycle, the degree of grazing by the mollusc decreases with increasing algal calcification, the cell walls of the alga progressively calcify, and the eventually highly calcificied stalks are completely resistant to ascoglossan grazing. In contrast, the exploitation of the algal chloroplasts retained by the molluscs increases during the seasonal cycle. The progressively increasing scarcity of food during the seasonal cycle may have led to the retention of symbiotic chloroplasts by E. timida. The developmental strategy of the ascoglossan also changes during the year: when food is abundant (in November, December, January, February and March) it is direct, with no planktonic larval phase, when food is scarce (in October, April, May and June) it is lecithotrophic, with a short planktonic larval phase. Chloroplast retention acts as a buffer, alleviating the effects of annual changes in density, structure and abundance of the alga on the nutritional state of the molluse.