Nature and role of newly described symbiotic associations between a sea anemone and gastropods at bathyal depths in the NW Atlantic

The hormathiid sea anemone Allantactis parasitica was found living as an epibiont on numerous species of gastropods at depths of 725-1100 m along the continental slope of eastern Canada. The proportion of bathyal gastropods hosting 1-6 sea anemones reached 72.5% in a single trawl. Although A. parasitica was occasionally found on other substrata (i.e. empty shells, pebbles), laboratory trials confirmed that they preferably associate with living gastropods. Settlement of planula larvae occurred significantly more often on the shells of live bathyal gastropods than on all other substrata present in the tanks. Juvenile sea anemones (∼ 1 mm diameter) readily moved from the mud or other inert substrata onto shells of burrowed bathyal gastropods. Conversely, larvae, juveniles and adults of A. parasitica never associated with any shallow-water gastropods when given the opportunity. Trials exposing predatory sea stars (Leptasterias polaris) from shallow and bathyal depths to bathyal gastropods (Buccinum undatum) with epibiotic A. parasitica, and to asymbiotic bathyal and shallow-water B. undatum, revealed adaptive behaviours in both prey and predator. Shallow-water gastropods (devoid of epibionts) reacted defensively to L. polaris, whereas bathyal gastropods relied mostly on their epibionts to protect them, thus falling prey to L. polaris when the epibionts were removed. L. polaris from bathyal depths typically ignored symbiotic gastropods, but they consistently preyed on asymbiotic ones, while L. polaris from shallow areas initially attempted to prey on all gastropods, but learned to avoid those harbouring sea anemones. Furthermore, living as epibionts afforded sea anemones a means to escape one of their own predators, the sea star Crossaster papposus. The mutualistic relationship between hormathiid sea anemones and bathyal gastropods from the NW Atlantic may have evolved in response to predation pressure.     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Plant defences and the role of epibiosis in mediating within-plant feeding choices of seagrass consumers

Within-plant variation in susceptibility to herbivory can significantly influence the ecological and evolutionary consequences of plant–herbivore interactions. Seagrasses are marine angiosperms characterised by substantial intra-individual differences in multiple traits, such as nutrients, chemical and structural defences and epibiotic load, all of which can strongly influence herbivore preferences. We quantified the within-plant feeding choices of the two main consumers of the temperate seagrass Posidonia oceanica––the fish Sarpa salpa and the sea urchin Paracentrotus lividus––and determined the plant traits that explained their foraging strategies. We found strong within-plant heterogeneity in both seagrass susceptibility to herbivory and chemical composition, but different consumers exhibited contrasting feeding choices. S. salpa preferred the most nutritious and chemically defended younger leaves, suggesting a full adaptation to consuming this macrophyte and a greater impact of this herbivore on the plant. In contrast, P. lividus consistently preferred the older leaves covered by epibionts, probably attenuating the relative impact of this consumer and helping to explain the weak effects usually recorded for this echinoid in undisturbed meadows. Artificial diet experiments showed that morphology and fine-scale structural defences were the primary determinant of urchin feeding choices, with nutrient content and chemical defences being of secondary importance. Epibiosis did not strongly influence fish feeding, but it did have a strong ‘shared-doom’ effect on urchin consumption. This effect was driven by a distinct preference towards a mixed diet that included both host tissues and their epibiotic community.     

Quorum sensing control of phosphorus acquisition in Trichodesmium consortia

Colonies of the cyanobacterium Trichodesmium are abundant in the oligotrophic ocean, and through their ability to fix both CO₂ and N₂, have pivotal roles in the cycling of carbon and nitrogen in these highly nutrient-depleted environments. Trichodesmium colonies host complex consortia of epibiotic heterotrophic bacteria, and yet, the regulation of nutrient acquisition by these epibionts is poorly understood. We present evidence that epibiotic bacteria in Trichodesmium consortia use quorum sensing (QS) to regulate the activity of alkaline phosphatases (APases), enzymes used by epibionts in the acquisition of phosphate from dissolved-organic phosphorus molecules. A class of QS molecules, acylated homoserine lactones (AHLs), were produced by cultivated epibionts, and adding these AHLs to wild Trichodesmium colonies collected at sea led to a consistent doubling of APase activity. By contrast, amendments of (S)-4,5-dihydroxy-2,3-pentanedione (DPD)—the precursor to the autoinducer-2 (AI-2) family of universal interspecies signaling molecules—led to the attenuation of APase activity. In addition, colonies collected at sea were found by high performance liquid chromatography/mass spectrometry to contain both AHLs and AI-2. Both types of molecules turned over rapidly, an observation we ascribe to quorum quenching. Our results reveal a complex chemical interplay among epibionts using AHLs and AI-2 to control access to phosphate in dissolved-organic phosphorus.     

Ability of Monaco shrimp Lysmata seticaudata (Decapoda: Hippolytidae) to control the pest glass anemone Aiptasia pallida (Actiniaria: Aiptasidae)

The present work highlights the ability of wild and cultured Monaco shrimp Lysmata seticaudata to control the glass anemone Aiptasia pallida. Starved shrimp ingested the highest percentages of glass anemones [85.7% and 89.3% for wild (W) and cultured (C) shrimp, respectively]. The absence of symbiotic zooxanthellae in glass anemones did not influence the shrimps feeding rate, with shrimp offered aposymbiotic anemones displaying the same feeding percentages (67.3% and 70.7% for W and C shrimp, respectively) as those offered sea anemones with symbiotic zooxanthellae (70.0% and 74.4% for W and C shrimp, respectively). Shrimp offered larger sea anemones had the lowest feeding percentages (33.0% and 36.3% for W and C shrimp, respectively), along with shrimp offered an alternative food (27.3% and 36.0% for W and C shrimp, respectively). There were no significant differences in the percentage of glass anemones ingested by cultured and wild Monaco shrimp in the same feeding trial.     

Effects of feeding regime on growth rate in the Mediterranean Sea anemone Actinia equina (Linnaeus)

Polyps of Actinia equina are the most common sea anemones in the rocky intertidal zone of the Mediterranean coast of Israel, where they occur in one of the southernmost populations of this species in the northern hemisphere. We examined effects of feeding rate on polyp growth at ambient sea temperature for this population. Under laboratory conditions, polyps were left unfed, or were fed with brine shrimp (Artemia) once every 2 weeks, once a week, or twice a week. Of the four experimental treatments, only feeding twice a week resulted in polyp growth; under all other regimes, the sea anemones lost body mass. We conclude that a high rate of feeding is required at sea temperatures in the eastern Mediterranean, where these sea anemones may have high metabolic rates relative to more northern populations.     

CHANGES IN THE ULTRASTRUCTURE OF SYMBIOTIC ZOOXANTHELLAE (SYMBIODINIUM SP., DINOPHYCEAE) IN FED AND STARVED SEA ANEMONES MAINTAINED UNDER HIGH AND LOW LIGHT1

The ultrastructure of symbiotic dinoflagellates (Symbiodinium sp., zooxanthellae) in the sea anemone Aiptasia pallida Verrill was examined in well-fed or starved (up to 120 days) anemones maintained under two light levels (5 and 50 μmol · m^?2· s^?1). Cell size of zooxanthellae was not affected by feeding history; however, both light and feeding history affected the relative cell volume of chloroplasts, lipids, and vacuoles. Stereological analysis of transmission electron micrographs showed that algae in low-light starved anemones had 10 times as much lipid (17.4% of cell volume) as those in well-fed anemones under the same light conditions (1.8%). The lipid content of algae from anemones in high light increased from 15.4% in well-fed anemones to 30.1% in starved anemones. The starch content of zooxanthellae in low-light anemones was law (4.1%) and not affected by feeding history, while the starch content of zooxanthellae in high-light anemones was greater (10.7%), with some differences among groups. Algal photoacclimation to low light included an increase in chloroplast relative volume from 17% (in well-fed high-light anemones) to 33% in well-fed low-light anemones. Starvation of the host resulted in a significant decrease in chloroplast volume in zooxanthellae in anemones at both light levels. Morphometry provides quantitative confirmation of biochemical and physiological data on zooxanthellae, because the changes in zooxanthellae with starvation of the host are consistent with other indicators of nutrient limitation of zooxanthellae of A. pallida held without food for long periods of time.     

Molecular evidence of digestion and absorption of epibiotic bacterial community by deep-sea crab Shinkaia crosnieri

The hydrothermal vent crab Shinkaia crosnieri is considered to obtain nutrition from the epibiotic bacteria found on the setae, but previous studies have not shown how nutrients can be transferred from the epibionts to the host. In this study, microscopic observations of S. crosnieri intestinal components detected autofluorescent setae fragments and pigmentation derived from the digestion of epibionts in a dye-stained epibiont tracer experiment. An in vitro digestion experiment with epibiotic populations using an intestinal extract demonstrated the degradation of epibiotic cells by digestive enzymes. A phylogenetic analysis showed that many of the bacterial 16S ribosomal RNA gene sequences obtained from the intestine were closely related to the sequences of the epibionts, thus they were probably derived from the epibionts. A stable isotope tracer experiment also indicated that ¹³C assimilated by the epibionts provided a carbon (nutrition) source for the host. Both activity measurements and isotope studies showed that chemosynthetic metabolism by the gut microbial components were inactive. Together with the feeding behaviour of living S. crosnieri, these results indicate that S. crosnieri ingests the epibionts using maxillipeds and assimilates them via its digestive organs as a nutrient source. The results of this study elucidate the mechanism of nutritional transfer in ectosymbiosis between chemosynthetic bacteria and deep-sea invertebrates.     

Flexibility of nutritional strategies within a mutualism: food availability affects algal symbiont productivity in two congeneric sea anemone species

Mutualistic symbioses are common, especially in nutrient-poor environments where an association between hosts and symbionts can allow the symbiotic partners to persist and collectively out-compete non-symbiotic species. Usually these mutualisms are built on an intimate transfer of energy and nutrients (e.g. carbon and nitrogen) between host and symbiont. However, resource availability is not consistent, and the benefit of the symbiotic association can depend on the availability of resources to mutualists. We manipulated the diets of two temperate sea anemone species in the genus Anthopleura in the field and recorded the responses of sea anemones and algal symbionts in the family Symbiodiniaceae to our treatments. Algal symbiont density, symbiont volume and photosynthetic efficiency of symbionts responded to changes in sea anemone diet, but the responses depended on the species of sea anemone. We suggest that temperate sea anemones and their symbionts can respond to changes in anemone diet, modifying the balance between heterotrophy and autotrophy in the symbiosis. Our data support the hypothesis that symbionts are upregulated or downregulated based on food availability, allowing for a flexible nutritional strategy based on external resources.     

Ammonia flux,physiological parameters,and <Emphasis Type="Italic">Symbiodinium</Emphasis> diversity in the anemonefish symbiosis on Red Sea coral reefs

Despite the ecological importance of anemonefish symbioses, little is known about how nutritional contributions from anemonefish interact with sea anemone physiology and Symbiodinium (endosymbiotic dinoflagellate) genetic identity under field conditions. On Red Sea coral reefs, we measured variation in ammonia concentrations near anemones, excretion rates of anemonefish, physiological parameters of anemones and Symbiodinium, and genetic identity of Symbiodinium within anemones. Ammonia concentrations among anemone tentacles were up to 49% above background levels, and anemonefish excreted ammonia significantly more rapidly after diurnal feeding than they did after nocturnal rest, similar to their excretion patterns under laboratory conditions. Levels of 4 physiological parameters (anemone protein content, and Symbiodinium abundance, chlorophyll a concentration, and division rate) were similar to those known for laboratory-cultured anemones, and in the field did not depend on the number of anemonefish per anemone or depth below sea surface. Symbiodinium abundance varied significantly with irradiance in the shaded reef microhabitats occupied by anemones. Most anemones at all depths harbored a novel Symbiodinium 18S rDNA variant within internal transcribed spacer region 2 (ITS2) type C1, while the rest hosted known ITS2 type C1. Association with Symbiodinium Clade C is consistent with the symbiotic pattern of these anemones on other Indo-Pacific reefs, but the C1 variant of Symbiodinium identified here has not been described previously. We conclude that in the field, anemonefish excrete ammonia at rapid rates that correlate with elevated concentrations among host anemone tentacles. Limited natural variation in anemonefish abundance may contribute to consistently high levels of physiological parameters in both anemones and Symbiodinium, in contrast to laboratory manipulations where removal of fish causes anemones to shrink and Symbiodinium to become less abundant.     

Effects of epibiotic algae on the survival, biomass and recruitment of mussels, Mytilus L. (Bivalvia: Mollusca)

Habitats composed of living ‘ecosystem engineers’, such as mussels, are subject to direct and indirect interactions with organisms that live among them. These interactions may affect the presence and structure of habitat, and hence, the associated taxa. We examined the direct effects of epibiotic algae on the survival, biomass and recruitment of mussels (Mytilus L.) on the west coast of Ireland. A field experiment showed that the presence of epibiotic fucoid algae reduced the likelihood of survival of mussels during storms. We also found that the strength of attachment of mussels did not increase in the presence of epibionts. Another in situ experiment revealed that the presence of ephemeral epibiotic algal mats had no effect on the biomass of host mussels, suggesting no effect on mussel growth or production. The abundance of small mussels (< 5 mm) covaried with cover of epibiotic algae at one location, but experimental results suggest no direct effect of the algae on recruitment unless through the action of chemical cues which linger after the removal of algae. We have identified a negative direct interaction between epibiotic algae and mussels on exposed rocky shores, which may often be characterised by bottom-up regulation. It is thought that positive interactions may be more important on more sheltered or sedimentary shores where top-down processes are more likely to dominate.     

Methane-Carbon Flow into the Benthic Food Web at Cold Seeps – A Case Study from the Costa Rica Subduction Zone

Cold seep ecosystems can support enormous biomasses of free-living and symbiotic chemoautotrophic organisms that get their energy from the oxidation of methane or sulfide. Most of this biomass derives from animals that are associated with bacterial symbionts, which are able to metabolize the chemical resources provided by the seeping fluids. Often these systems also harbor dense accumulations of non-symbiotic megafauna, which can be relevant in exporting chemosynthetically fixed carbon from seeps to the surrounding deep sea. Here we investigated the carbon sources of lithodid crabs (Paralomis sp.) feeding on thiotrophic bacterial mats at an active mud volcano at the Costa Rica subduction zone. To evaluate the dietary carbon source of the crabs, we compared the microbial community in stomach contents with surface sediments covered by microbial mats. The stomach content analyses revealed a dominance of epsilonproteobacterial 16S rRNA gene sequences related to the free-living and epibiotic sulfur oxidiser Sulfurovum sp. We also found Sulfurovum sp. as well as members of the genera Arcobacter and Sulfurimonas in mat-covered surface sediments where Epsilonproteobacteria were highly abundant constituting 10% of total cells. Furthermore, we detected substantial amounts of bacterial fatty acids such as i-C15∶0 and C17∶1ω6c with stable carbon isotope compositions as low as −53‰ in the stomach and muscle tissue. These results indicate that the white microbial mats at Mound 12 are comprised of Epsilonproteobacteria and that microbial mat-derived carbon provides an important contribution to the crab''s nutrition. In addition, our lipid analyses also suggest that the crabs feed on other ¹³C-depleted organic matter sources, possibly symbiotic megafauna as well as on photosynthetic carbon sources such as sedimentary detritus.     

Lipid‐rich zooplankton subsidise the winter diet of benthivorous Arctic charr (Salvelinus alpinus) in a subarctic lake

相似文献   

Fatty acid signatures of stomach oil and adipose tissue of northern fulmars (<Emphasis Type="Italic">Fulmarus glacialis</Emphasis>) in Alaska: implications for diet analysis of Procellariiform birds

Procellariiforms are unique among seabirds in storing dietary lipids in both adipose tissue and stomach oil. Thus, both lipid sources are potentially useful for trophic studies using fatty acid (FA) signatures. However, little is known about the relationship between FA signatures in stomach oil and adipose tissue of individuals or whether these signatures provide similar information about diet and physiology. We compared the FA composition of stomach oil and adipose tissue biopsies of individual northern fulmars (N = 101) breeding at three major colonies in Alaska. Fatty acid signatures differed significantly between the two lipid sources, reflecting differences in dietary time scales, metabolic processing, or both. However, these signatures exhibited a relatively consistent relationship between individuals, such that the two lipid sources provided a similar ability to distinguish foraging differences among individuals and colonies. Our results, including the exclusive presence of dietary wax esters in stomach oil but not adipose tissue, are consistent with the notion that stomach oil FA signatures represent lipids retained from prey consumed during recent foraging and reflect little metabolic processing, whereas adipose tissue FA signatures represent a longer-term integration of dietary intake. Our study illustrates the potential for elucidating short- versus longer-term diet information in Procellariiform birds using different lipid sources.     

Epibiotic protozoan communities on juvenile southern king crabs (<Emphasis Type="Italic">Lithodes santolla</Emphasis>) from subantarctic areas

In this study, we describe for the first time the composition of epibiotic protozoan communities on juvenile southern king crabs Lithodes santolla. Basibionts were collected in subantarctic bays near Santa Ana Point (Magellan Strait, Chile). Seven epibiotic protozoan ciliates were found: Ephelota gemmipara, Ephelota gigantea, Podophrya fixa, Acineta tuberosa, Zoothamnium duplicatum, Chilodochona quennerstedti and Gymnodinioides sp. The mean number of epibionts per crab was 99 (maximum 897). Both Ephelota species were the most abundant and most widely distributed epibionts on the crab examined. The lengths of basibionts were correlated with the number of epibionts, indicating a relationship between the age of the crab and the state of epibiosis. Epibionts differed in their distribution patterns along the anteroposterior axis of the crab. Less abundant species tended to occupy sites that are not colonized by the predominant Ephelota species. Maxillipeds and pereiopods were most densely colonized by epibionts. The behaviours of epibionts and basibionts related to epibiosis are discussed. The recent studies about invasive lithodids in Antarctic areas enhance the interest for the epibiotic communities colonizing these crustaceans, which can illustrate changes in biodiversity and state of these environments.     

Symbiotic interactions in the Silurian of North America

There are at least 47 different symbiotic pairs of taxa and 16 symbiotic associations in the Silurian of North America. Crinoids are most common host species and they hosted variety of epibiotic and endobiotic symbionts, including Tremichnus, platyceratid gastropods, brachiopods, microconchids, cornulitids, cyclostome bryozoans and favositid tabulates. Eighteen symbiotic pairs contain at least one colonial partner. Stromatoporoids hosted the most diverse fauna of endobiotic symbionts, including cornulitids, lingulids, Chaetosalpinx, Heliocosalpinx and rugosans. Among 16 symbiotic associations of Silurian of North America, 8 are common between North America and Baltica. North American symbiotic associations involving stromatoporoid hosts are the most similar to their Baltic equivalents.     

Effects of anemonefish on giant sea anemones: expansion behavior,growth, and survival

The symbiosis between giant sea anemones and anemonefish on coral reefs is well known, but little information exists on impacts of this interaction on the sea anemone host. On a coral reef at Eilat, northern Red Sea, individuals of the sea anemone Entacmaea quadricolor that possessed endemic anemonefish Amphiprion bicinctus expanded their tentacles significantly more frequently than did those lacking anemonefish. When anemonefish were experimentally removed, sea anemone hosts contracted partially. Within 1–4 h in most cases, individuals of the butterflyfish Chaetodon fasciatus arrived and attacked the sea anemones, causing them to contract completely into reef holes. Upon the experimental return of anemonefish, the anemone hosts re-expanded. The long-term growth rate and survival of the sea anemones depended on the size and number of their anemonefish. Over several years, sea anemones possessing small or no fish exhibited negative growth (shrinkage) and eventually disappeared, while those with at least one large fish survived and grew. We conclude that host sea anemones sense the presence of symbiotic anemonefish via chemical and/or mechanical cues, and react by altering their expansion behavior. Host sea anemones that lack anemonefish large enough to defend them against predation may remain contracted in reef holes, unable to feed or expose their tentacles for photosynthesis, resulting in their shrinkage and eventual death.     

Asexual Propagation of Sea Anemones That Host Anemonefishes: Implications for the Marine Ornamental Aquarium Trade and Restocking Programs

Anemonefishes and their host sea anemones form an iconic symbiotic association in reef environments, and are highly sought after in the marine aquarium trade. This study examines asexual propagation as a method for culturing a geographically widespread and commonly traded species of host sea anemone, Entacmaea quadricolor. Two experiments were done: the first to establish whether size or colour morph influenced survival after cutting into halves or quarters; and the second to see whether feeding was needed to maximise survival and growth after cutting. Survival rates were high in both experiments, with 89.3 and 93.8% of the anemones cut in half, and 62.5 and 80.4% cut in quarters surviving in experiments 1 and 2, respectively. Anemones that were cut in half were larger in size, and healed and grew quicker than those cut in quarters. However, even though survival was lower when the individuals were cut in quarters, this treatment produced the greatest number of anemones. Feeding increased oral disc diameter growth and reduced wet weight loss, but did not significantly influence pedal disc diameter. Given that the anemones took up to 56 d to form an off-centre mouth, it is highly likely that feeding may have produced greater effect if the experiment was run for longer. This low technology method of propagation could be used to produce individuals throughout the year and the anemones could then be used to supply the aquarium trade or restock depleted habitats, thus supporting biodiversity conservation in coral reef areas.     

The contrasting diet of Harpagifer antarcticus (Notothenioidei, Harpagiferidae) at two localities of the South Shetland Islands, Antarctica

The diet of Harpagifer antarcticus was studied at two localities of the South Shetland Islands, Antarctica. The analysis of the stomach contents of specimens collected in tide pools at Potter Cove, King George Island, indicated that gammarid amphipods (mainly Gondogeneia sp.) were the main prey of this fish, followed by polychaetes, gastropods and isopods. By contrast, the specimens from Harmony Point, Nelson Island, which were recovered from stomach contents of Antarctic shags Phalacrocorax bransfieldensis feeding at depths of 46–110 m, preyed almost exclusively on the Antarctic krill Euphausia superba. These results are discussed and compared with previous studies. Received: 27 August 1997 / Accepted: 17 November 1997     

Impact of symbiotic algae on sea anemone metabolism: analysis by in vivo 31P nuclear magnetic resonance spectroscopy

High-field pulsed Fourier-transform nuclear magnetic resonance spectroscopy (NMR) was used to quantify the adenylate levels of sea anemones (Aiptasia pulchella) with and without symbiotic dinoflagellates (Symbiodinium sp.). Animals were fed to repletion, then starved in darkness for up to six days before collection of in vivo NMR spectra. The host adenylate ratio of ATP: (ATP + ADP) declined significantly with increasing periods of starvation in both symbiotic and aposymbiotic hosts (P less than 0.05). However, the decline in the animal adenylate ratio was significantly more rapid in animals bearing symbiotic algae (P less than 0.05). This suggests that symbiotic algae in darkness cause more rapid depletion of host energy reserves, possibly by drawing on host pools of organic substrates. In vivo NMR spectroscopy was also used to evaluate the effect on A. pulchella of photosynthesis by zooxanthellae. Symbiotic anemones were fed to repletion, then starved under high irradiance (300 to 320 mu Ein m-2 s-1) or low irradiance (70 to 80 mu Ein m-2 s-1) conditions for up to five days. The host adenylate ratio declined significantly (P less than 0.01) with starvation under both treatments, but no significant difference was detected between treatments (P greater than 0.35). Blotted wet weight of anemones under high and low irradiance declined by 50% over eight days of starvation, but there was no significant difference in the rate of weight loss by anemones in the two treatments. There results suggest that translocation of photosynthate from symbiotic zooxanthellae does not significantly affect host adenylate ratio or have a sparing effect on host biomass during starvation in this symbiotic sea anemone.