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Thirteen years ago in a review that appeared in the American Zoologist, we presented the first survey of the chemical and ecological bioactivity of Antarctic shallow-water marine invertebrates. In essence, we reported that despite theoretical predictions to the contrary the incidence of chemical defenses among sessile and sluggish Antarctic marine invertebrates was widespread. Since that time we and others have significantly expanded upon the base of knowledge of Antarctic marine invertebrates' chemical ecology, both from the perspective of examining marine invertebrates in new, distinct geographic provinces, as well as broadening the evaluation of the ecological significance of secondary metabolites. Importantly, many of these studies have been framed within established theoretical constructs, particularly the Optimal Defense Theory. In the present article, we review the current knowledge of chemical ecology of benthic marine invertebrates comprising communities along the Western Antarctic Peninsula (WAP), a region of Antarctica that is both physically and biologically distinct from the rest of the continent. Our overview indicates that, similar to other regions of Antarctica, anti-predator chemical defenses are widespread among species occurring along the WAP. In some groups, such as the sponges, the incidence of chemical defenses against predation is comparable to, or even slightly higher than, that found in tropical marine systems. While there is substantial knowledge of the chemical defenses of benthic marine invertebrates against predators, much less is known about chemical anti-foulants. The sole survey conducted to date suggests that secondary metabolites in benthic sponges are likely to be important in the prevention of fouling by benthic diatoms, yet generally lack activity against marine bacteria. Our understanding of the sensory ecology of Antarctic benthic marine invertebrates, despite its great potential, remains in its infancy. For example, along the WAP, community-level non-consumptive effects occur when amphipods chemically sense fish predators and respond by seeking refuge in chemically-defended macroalgae. Such interactions may be important in releasing amphipods from predation pressure and facilitating their unusually high abundances along the WAP. Moreover, recent studies on the sensory biology of the Antarctic keystone sea star Odontaster validus indicate that chemotactile-mediated interactions between conspecifics and other sympatric predatory sea stars may have significant ramifications in structuring community dynamics. Finally, from a global environmental perspective, understanding how chemical ecology structures marine benthic communities along the WAP must increasingly be viewed in the context of the dramatic impacts of rapid climatic change now occurring in this biogeographic region.

Sensitivity (chemoreception) to different amino acids was studied in six invertebrate species: Serolis polita, Glyptonotus antarcticus, Abyssochromene plebs, Waldeckia obesa, Odontaster validus, and Sterechinus neumayeri. The sensitivity was estimated by the changes in basic metabolism (respiration rate). Starvation increased the sensitivity in all the species. The metabolism rates increased in the presence of L-glutamic acid in G. antarcticus, A. plebs, O. validus, and St. neumayeri. The serine and arginine amino acids had a significant impact on the metabolism of the necrophagous species S. polita and W. obesa. The chemical information may be mediated by means of L-glutamic acid via glutamate receptors, which can be blocked by kynurenic acid, as occurs in the experiments with G. antarcticus and A. plebs.

Chemical investigation of the lipophilic extract of the Antarctic soft coral Alcyonium grandis led us to the finding of nine unreported sesquiterpenoids, 2-10. These molecules are members of the illudalane class and in particular belong to the group of alcyopterosins, illudalanes isolated from marine organisms. The structures of 2-10 were determined by interpretation of spectroscopic data. Repellency experiments conducted using the omnivorous Antarctic sea star Odontaster validus revealed a strong activity in the lipophilic extract of A. grandis against predation.

The present study surveyed the palatability of all sponge species that could be collected in sufficient quantities in a shallow-water area along the western Antarctic Peninsula. Of 27 species assayed, 78% had outermost tissues that were significantly unpalatable to the sympatric, omnivorous sea star Odontaster validus. Of those species with unpalatable outer tissues, 62% had inner tissues that were also unpalatable to the sea stars. Sea stars have often been considered as the primary predators of sponges in other regions of Antarctica, and their extra-oral mode of feeding threatens only the outermost sponge tissues. The observation that many of the sponges allocate defenses to inner tissues suggests the possibility that biting predators such as mesograzers, which could access inner sponge layers, may also be important in communities along the Antarctic Peninsula. In feeding bioassays with extracts from 12 of the unpalatable species in artificial foods, either lipophilic or hydrophilic extracts were deterrent in each species. These data indicate an overall level of chemical defenses in these Antarctic sponges that is comparable to, and slightly greater than, that found in a previous survey of tropical species.

Members of the HSP70 gene family comprising the inducible (HSP70) genes and GRP78 (glucose-regulated protein 78 kDa) were identified in an Antarctic sea star (Odontaster validus) and an Antarctic gammarid (Paraceradocus gibber). These genes were surveyed for expression levels via Q-PCR after an acute 2-hour heat shock experiment in both animals and a time course assay in O. validus. No significant up-regulation was detected for any of the genes in either of the animals during the acute heat shock. The time course experiment in O. validus produced slightly different results with an initial down regulation in these genes at 2°C, but no significant up-regulation of the genes either at 2 or 6°C. Therefore, the classical heat shock response is absent in both species. The data is discussed in the context of the organisms’ thermal tolerance and the applicability of HSP70 to monitor thermal stress in Antarctic marine organisms.

The loss of the shell in nudibranch gastropods has been related to the acquisition of chemical defensive strategies during evolution, such as the use of natural products to deter predation. In the present study, we investigated the origin, location, and putative role of granuloside (1), a homosesterterpene lactone, recently isolated from the Antarctic nudibranch Charcotia granulosa Vayssiere, 1906. Several adults, egg masses, and its bryozoan prey, Beania erecta Waters, 1904, were chemically analyzed by chromatographic and spectroscopic techniques. Light- and transmission electron microscopy of the mantle revealed complex glandular structures, which might be associated with the storage of defensive compounds in analogy to mantle dermal formations described in other nudibranchs. Although preliminary in situ repellence bioassays with live specimens of the nudibranch showed avoidance against the Antarctic generalist sea star predator Odontaster validus, the specific role of the terpene granuloside requires further investigation. The egg masses do not present granuloside, and the glandular structures are absent in the trochophore larvae. Our results suggest that C. granulosa synthesizes granuloside de novo in early stages of its ontogeny, instead of obtaining it from the prey. Considering the wide geographic area inhabited by this slug, this may be advantageous, because natural products produced by the slug will not be affected by fluctuant food availability. Overall, the Antarctic sea slug C. granulosa seems to possess defensive strategies that are similar, in terms of production and storage, to nudibranchs from other regions of the world. This species is one of the few cladobranchs investigated so far that present de novo biosynthesis of a defensive compound.

UNSPECIFIC IMMUNE RESPONSE OF ANTARCTIC ECTOTERMIC ANIMALS UNDER POLAR TEMPERATURES. This review presents data on nonspecific immune system of Antarctic metazoans poekiloterms. The phagocytosis, inflammation and wound repair were analysed in Antarctic animals, the big head fish (Notothenia coriiceps = N. neglecta ), Antarctic starfish (Odontaster validus) and in Antarctic sea urchin (Sterechinus neumayeri).

Chemical cues released from dead or injured organisms constitute important signals informing nearby animals about a feeding possibility. The ability to detect the signal, evaluate its meaning and locate its source can help organisms to exploit food resources efficiently, which is especially important to animals living in environments with limited food supply. Experiments were carried out to study the behavioral responses of several Antarctic benthic invertebrates to fish (Notothenia corriceps) blood. Necrophagous species such as sea stars Odontaster validus and Lysasterias sp., amphipod Waldeckia obesa and nemertean Parborlasia corrugatus responded to fish blood with changes in their behavior. The behavior common to all these species was locomotion directed towards the stimulus source. Behavioral components consistent with food consumption were observed in O. validus and P. corrugatus. The reaction of herbivorous limpets Nacella concinna to fish blood depended on the animal size. Large (>10 mm) limpets showed no behavioral response, whereas small ones (

The response of scavengers to a feeding cue at Adelaide Island, West Antarctic Peninsula was investigated using a baited video camera system. Fourteen experimental deployments, each lasting 72 h were conducted at two contrasting sites during the winter and summer of 2005. The rate of bait consumption varied between sites but not between seasons, and was low in comparison with studies at lower latitudes and greater depths. At the Hangar Cove site, the nemertean Parborlasia corrugatus was out-competed at the bait and displaced by the lysianassid amphipod Cheirmedon femoratus during winter. However, C. femoratus did not feed on the bait during summer, allowing P. corrugatus to monopolise the feeding opportunity. At the South Cove site the asteroid Odontaster validus dominated the bait in both seasons but sporadic feeding by the nototheniid fish Notothenia coriiceps considerably affected consumption rates during two of the six deployments. Scavengers were attracted to the bait in very high numbers and opportunistic necrophagy seems to be a successful strategy in an environment that is intensely disturbed by ice.

This research was supported by a University of Glasgow Faculty Scholarship to KMD, a Collaborative Gearing Scheme grant from the Natural Environmental Research Council and the British Antarctic Survey (CGS-77) and an ASSEMBLE infrastructure access grant to DMB. Baited underwater cameras are becoming a popular tool to monitor fish and invertebrate populations within protected and inshore environments where trawl surveys are unsuitable. Modelling the arrival times of deep-sea grenadiers using an inverse square relationship has enabled abundance estimates, comparable to those from bottom trawl surveys, to be gathered from deep-sea baited camera surveys. Baited underwater camera systems in the shallow water environments are however, currently limited to relative comparisons of assemblages based on simple metrics such as MaxN (maximum number of fish seen at any one time). This study describes a stochastic simulation approach used to model the behaviour of fish and invertebrates around a BUC system to enable absolute abundance estimates to be generated from arrival patterns. Species-specific models were developed for the tropical reef fishes the black tip grouper (Epinephelus fasciatus) and moray eel (Gymnothorax spp.) and the Antarctic scavengers; the asteroid (Odontaster validus) and the nemertean worm (Parbolasia corrugatus). A sensitivity analysis explored the impact of input parameters on the arrival patterns (MaxN, time to the arrival of the first individual and the time to reach MaxN) for each species generated by the model. Sensitivity analysis showed a particularly strong link between MaxN and abundance indicating that this model could be used to generate absolute abundances from existing or future MaxN data. It in effect allows the slope of the MaxN vs. abundance relationship to be estimated. Arrival patterns generated by each model were used to estimate population abundance for the focal species and these estimates were compared to data from underwater visual census transects. Using a Bland-Altman analysis, baited underwater camera data processed using this model were shown to generate absolute abundance estimates that were comparable to underwater visual census data. Postprint

One characteristic pattern found in the marine Antarctic shallow environments is the unusually high proportion of species with protected and pelagic lecitotrophic development modes. However, species with planktotrophic development generally appear as the most conspicuous types of organisms in these environments. The Antarctic shallow benthos is considered as one of the most disturbed in the world, mainly due to the action of ice, thus one could hypothesize that such an environment should favor organisms with high dispersal capability. In order to test this general hypothesis, for two consecutive summers (2004–2005) and at two locations, we quantified the abundance and size distribution of most echinoderms present along bathymetric transects. Our results show the predominance of broadcasters (i.e., Sterechinus neumayeri and Odontaster validus) at a location where disturbances are common, while brooders (e.g., Abatus agassizii) only occurred at shallower depths of the least disturbed location. These results not only corroborate the hypothesis that local disturbance is an important factor generating these ecological patterns, but also suggest how ice-related disturbances could represent a major selecting agent behind the patterns of species diversity at an evolutionary scale in Antarctica.

The sea stars Granaster nutrix and Neosmilaster georgianus are conspicuous members of benthic communities along the Antarctic Peninsula. An analysis of the proximate composition of somatic body components of nonreproductive adults indicates the nutrient storage organs (pyloric caeca) are rich in both protein (60.7 and 60.6% mean dry wt, respectively) and lipid (25.4 and 29.8% mean dry wt, respectively). Body-wall tissues, while containing small inconspicuous skeletal ossicles, are also comprised of significant levels of organic matter (33.5 and 55.7% mean dry wt, respectively), attributable primarily to protein. Both the pyloric caeca and body-wall tissues are relatively rich in energy (mean energy levels=24.8 and 26.5 kJ g−1 dry wt; 8.4 and 14.1 kJ g−1 dry wt, respectively). Despite the availability of these nutrients and energy neither sea star is preyed upon by the sympatric omnivorous sea star Odontaster validus, a common predator of other Antarctic sea stars. Laboratory feeding bioassays indicate that O. validus rejects live intact individuals and body-wall tissues of both sea star species while readily consuming dried krill. Alginate food pellets containing a krill powder and tissue level concentrations of organic methanol extracts of body-wall tissues were also rejected by O. validus. Moreover, the copious mucus released from the body wall of N. georgianus was deterrent in O. validus food pellet bioassays. Thus, both sea stars evidently possess defensive secondary metabolites that defend against predation and are likely to play a role in mediating materials and energy transfer in the Antarctic benthos.

The colonial ascidian Distaplia cylindrica occurs both as scattered individual colonies or in “gardens” of colonies in fine-grained soft substrata below 20 m depths off Anvers Island along the Antarctic Peninsula. Individual colonies, shaped as tall rod-like cylinders and anchored in the sediments by a bulbous base, may measure up to 7 m in height. D. cylindrica represent a considerable source of materials and energy for prospective predators, as well as potential surface area for fouling organisms. Nonetheless, qualitative in situ observations provided no evidence of predation by sympatric predators such as abundant sea stars, nor obvious biofouling of colony surfaces. Mean energy content of whole-colony tissue of D. cylindrica was relatively high for an ascidian (14.7 kJ g−1 dry wt), with most of this energy attributable to protein (12.7 kJ g−1 dry wt). The sympatric omnivorous sea star Odontaster validus consistently rejected pieces of D. cylindrica colonies in laboratory feeding assays, while readily ingesting similarly sized alginate food pellets. Feeding deterrence was determined to be attributable to defensive chemistry, as colonies of D. cylindrica are nutritionally attractive and lack physical protection (conspicuous skeletal elements or a tough outer tunic), and O. validus display significant feeding-deterrent responses to alginate food pellets containing tissue-level concentrations of organic extracts. In addition, high acidity measured on outer colony surfaces (pH 1.5) as well as homogenized whole-colony tissues (pH 2.5) are indicative of surface sequestration of inorganic acids. Agar food pellets prepared at tissue levels of acidity resulted in significant feeding deterrence in sea stars. Thus, both inorganic acids and secondary metabolites contribute to chemical feeding defenses. D. cylindrica also possesses potent antifoulant secondary metabolites. Tissue-level concentrations of hydrophilic and lipophilic extracts caused significant mortality in a sympatric pennate diatom. Chemical feeding deterrents and antifoulants are likely to contribute to the abundance of D. cylindrica and, in turn, play a role in regulating energy transfer and community structure in benthic marine environments surrounding Antarctica.

The punctate terebratulid brachiopod Liothyrella uva is the most common brachiopod species in Antarctica. Whole brachiopods, either live or freeze dried and ground into a powder and suspended in alginate, were unpalatable to the sympatric macropredators Odontaster validus (an abundant, omnivorous sea star) and Notothenia coriiceps (an abundant, omnivorous, epibenthic fish). The unpalatability of these ground tissues coupled with that of lipophilic extracts of whole L. uva presented in alginate pellets to O. validus, suggests an involvement of chemical defenses. Several isolated brachiopod tissues were also unpalatable to O. validus after being freeze dried, ground and suspended in alginate, but only the pedicle was unpalatable in such preparations to both O. validus and N. coriiceps. This observation is consistent with the Optimal Defense Theory since the pedicle is the only tissue not protected inside the brachiopod shell. There was, however, no correlation between the energetic content and unpalatability of any of the individual tissues. Organic extracts of tissues involved in feeding (lophophore and intestine–stomach) had relatively strong antimicrobial activity when assayed against several strains of Antarctic bacteria. However, the lophophore was palatable to both macropredators, suggesting nonoverlapping chemical defenses are involved in protection against predators and pathogens.

The common Antarctic nudibranch Austrodoris kerguelenensis (Bergh) contains diterpene diacylglycerides only present in its external body parts. These compounds provide a chemical defense against sympatric predators, such as the seastar Odontaster validus Koehler. Bioassays conducted with O. validus revealed that live nudibranchs, mantle tissue and Et2O extract of the A. kerguelenensis mantle deterred feeding by the seastar. Further bioassays testing organic fractions of the Et2O mantle extract showed that the diterpene diacylglycerides, as well as corresponding monoacylglycerides and monoacylglycerides of regular fatty acids, were responsible for the feeding deterrence in O. validus. We suggest that A. kerguelenensis derives the bioactive diacylglycerides by de novo biosynthesis rather than by sequestration from its sponge diet, since the mollusk does not contain active metabolites in the viscera, and neither the active compounds nor precursors were detected in the sponge diet. Furthermore, A. kerguelenensis did not show a strong chemodetection or feeding preference for its main diet, hexactinellid sponges, in Y-maze and food choice experiments, respectively.

The Antarctic limpet, Nacella concinna, has the ability to escape predation by the predatory sea star, Neosmilaster georgianus. When the sea star comes into contact with the limpet, the limpet undergoes characteristic behaviors that include extension of its pallial tentacles, raising its shell in a mushroom-like fashion, rotation, and flight. These behaviors were not observed until the sea star physically contacted the limpets and were not induced by the omnivorous sea star, Odontaster validus, which occupies the same habitats as Ne. georgianus. Crude hydrophilic extracts from Ne. georgianus, but not from O. validus, induced the escape behaviors, indicating that this response is chemical as well as tactile. These putative escape behaviors displayed by Na. concinna may contribute to avoidance of predation by Ne. georgianus.

In an attempt to improve the understanding of the natural variability and distribution of trace elements in Antarctic organisms, the concentrations of arsenic, cadmium, cobalt, chromium, copper, manganese, nickel, vanadium and zinc in representative benthic species from two pristine coastal environments were measured and compared with literature data for other uncontaminated coastal ecosystems. Correlations between the elements, differences between the species and between the sampling sites were examined by principal component analysis. Metal accumulation was particularly evident in the tissues of the sea star Odontaster validus, the bivalve mollusc Laternula elliptica and in the red alga Phyllophora antarctica. However, metal accumulation was not the same for all the analytes, but, rather, depended on the organism characteristics. In particular, the soft tissues of Odontaster validus were characterized by high concentrations of cadmium, zinc and copper, those of Phyllophora antarctica by high concentrations of manganese and nickel, and the tissues of Laternula elliptica by high concentrations of all measured elements, particularly in its digestive gland. The Antarctic data as well as those reported for other pristine coastal ecosystems showed remarkably high natural variability in metal content, which must be taken into account when interpreting results from biomonitoring programmes.

The circumpolar nudibranch Tritoniella belli Eliot occurs in abundance in shallow-water benthic communities of McMurdo Sound, Antarctica. Density estimates based on belt transects averaged collectively 0.46 individuals m−2 at three study sites between depths of 6 and 30 m in November 1996. At two of the sites, population densities increased linearly between 18 and 30 m depth (up to 0.7 and 1.15 individuals m−2 at 30 m depth). Individuals at all sites were rare or absent at depths shallower than 12 m. Size frequencies of individuals at the sites were similar, and a pooled analysis revealed a unimodal distribution skewed highly towards juvenile size classes. This suggests both recent recruitment and constant rates of mortality across size classes. The relationship between foot length and wet weight best fits an exponential growth equation, indicative of an allometric growth pattern. Distribution of T. belli in the field suggests that it is a habitat and diet generalist. Potential invertebrate predators include sea anemones and seastars, both of which co-occur in abundance in McMurdo Sound. Laboratory experiments indicate that the sea anemone Isotealia antarctica can capture and ingest T. belli. However, 70% of T. belli that are captured escape from the tentacles or, following ingestion, are rejected from the gastrovascular cavity. The seastars Odontaster validus, Perknaster fuscus, and Acodontaster conspicuus, avoid contact with T. belli, but if forced into contact with mantle tissues, retract their tube-feet. Mucus secreted from the mantle tissues, coated on to the tips of glass rods, and presented to seastar tube-feet, causes significantly longer tube-foot retraction times than control rods. Moreover, pieces of freeze-dried krill coated with mantle mucus are consumed significantly less often than untreated control pieces of krill by a benthic scavenging fish (Pseudotrematomas bernacchi). Employing seastar tube-foot retractions as a bioassay, we found the bioactive compound(s) are soluble in ethyl acetate, indicating they are lipophilic or moderately hydrophilic in nature. Chemical defenses in the mucus of T. belli probably contribute to its high abundance in Antarctic benthic communities.

The effects of temperature on development and viability were measured at 14 levels between -2{deg}C and +3{deg}C on embryos of two asteroids (Odontaster validus and Odontaster meridionalis) and an echinoid (Sterechinus neumayeri) from Signy Island, Antarctica. Development rates were 2 to 10 times slower than those for temperate or tropical echinoderms, with times to hatching up to 240 h. Development rates for the two asteroids differed by 1.15 x, and rates for both species approximately doubled over the experimental temperature range. In O. validus, embryo viability was independent of temperature, but in O. meridionalis viability declined with increasing temperature. Development rates for S. neumayeri were little affected by temperature above +0.2{deg}C, but declined rapidly at lower temperatures. Conversely, the number of nonviable eggs was low and constant below +1.7{deg}C, but rose rapidly at higher temperatures. A window of optimal temperature, between +0.2{deg}C and +1.7{deg}C, has therefore been proposed for development time and embryo viability in this population of S. neumayeri. Spawning trials and field observations of larvae indicated that the time of gamete release and periods of larval development in S. neumayeri coincided with austral summer sea temperatures in the same window. Embryos of O. meridionalis and O. validus are released in winter, when temperatures are constantly below -1.6{deg}C. Comparison of the different strategies suggests that larval food supply and predation during planktonic phases are not the dominant ecological factors for these species.

Activities and characteristics of digestive enzymes of polar and subtropical echinoderms are similar. Specific adaptations to facilitate digestion at low temperature were not observed. Levels of chymotrypsin and β-glucosidase of digestive tissues of the Antarctic asteroids Odontaster validus and Odontaster meridionalis, and the Antarctic echinoid Sterechinus neumayeri, are comparable to levels of the asteroids Henricia downii and Astropecten articulatus from the Gulf of Mexico. α-Glucosidase activities are similar for S. neumayeri and A. articulatus. The pH of maximal activity is 7.5 for disaccharidases and 8.5–9.5 for chymotrypsin for both polar and subtropical species. Affinities for substrates were higher at 25°C than at 0°C for disaccharidases of all species. However, Km values for chymotrypsin increased from 0 to 25°C. Lack of sufficient adaptation by polar echinoderms to facilitate digestion at low temperature may reduce their capacity to digest food.

To evaluate levels of essential (zinc and copper) and non-essential (mercury and cadmium) heavy metals, 34 species of organisms from different areas close to the Antarctic Peninsula were analysed. These included algae, filter-feeders, omnivorous invertebrates and vertebrates. Mercury was not detected, while cadmium was found in the majority of organisms analysed (detection limit was 0.05 ppm for both metals). The highest cadmium concentration was observed in the starfish Odontaster validus. Anthozoans, sipunculids and nudibranchs showed maximum levels of zinc, while the highest copper level was found in the gastropod Trophon brevispira. Mercury and cadmium levels in fishes were below the detection limit. Concentrations of essential and non-essential metals in birds were highest in liver followed by muscle and eggs. Cadmium and mercury levels in muscle of southern elephant seals were above the detection limit, whereas in Antarctic fur seals they were below it. The objective of the study was to gather baseline information for metals in Antarctic Ocean biota that may be needed to detect, measure and monitor future environmental changes.

The Antarctic sea star Odontaster validus and the amphipod Cheirimedon femoratus are important predators in benthic communities. Some bryozoans are part of the diet of the asteroid and represent both potential host biosubstrata and prey for this omnivorous lysianassid amphipod. In response to such ecological pressure, bryozoans are expected to develop strategies to deter potential predators, ranging from physical to chemical mechanisms. However, the chemical ecology of Antarctic bryozoans has been scarcely studied. In this study we evaluated the presence of defenses against predation in selected species of Antarctic bryozoans. The sympatric omnivorous consumers O. validus and C. femoratus were selected to perform feeding assays with 16 ether extracts (EE) and 16 butanol extracts (BE) obtained from 16 samples that belonged to 13 different bryozoan species. Most species (9) were active (12 EE and 1 BE) in sea star bioassays. Only 1 BE displayed repellence, indicating that repellents against the sea star are mainly lipophilic. Repellence toward C. femoratus was found in all species in different extracts (10 EE and 12 BE), suggesting that defenses against the amphipod might be both lipophilic and hydrophilic. Interspecific and intraspecific variability of bioactivity was occasionally detected, suggesting possible environmental inductive responses, symbiotic associations, and/or genetic variability. Multivariate analysis revealed similarities among species in relation to bioactivities of EE and/or BE. These findings support the hypothesis that, while in some cases alternative chemical or physical mechanisms may also provide protection, repellent compounds play an important role in Antarctic bryozoans as defenses against predators.

Juvenile stages are often thought to be less resistant to thermal challenges than adults, yet few studies make direct comparisons using the same methods between different life history stages. We tested the resilience of juvenile stages compared to adults in 4 species of Antarctic marine invertebrate over 3 different rates of experimental warming. The species used represent 3 phyla and 4 classes, and were the soft-shelled clam Laternula elliptica, the sea cucumber Cucumaria georgiana, the sea urchin Sterechinus neumayeri, and the seastar Odontaster validus. All four species are widely distributed, locally abundant to very abundant and are amongst the most important in the ecosystem for their roles. At the slowest rate of warming used (1°C 3 days(-1)) juveniles survived to higher temperatures than adults in all species studied. At the intermediate rate (1°C day(-1)) juveniles performed better in 3 of the 4 species, with no difference in the 4(th), and at the fastest rate of warming (1°C h(-1)) L. elliptica adults survived to higher temperatures than juveniles, but in C. georgiana juveniles survived to higher temperatures than adults and there were no differences in the other species. Oxygen limitation may explain the better performance of juveniles at the slower rates of warming, whereas the loss of difference between juveniles and adults at the fastest rate of warming suggests another mechanism sets the temperature limit here.

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.

Although echinoderms constitute some of the most conspicuous taxa of the Antarctic benthic communities, the echinoderm fauna of Terra Nova has not been described yet. The present study provides the first species list of echinoids, ophiuroids and asteroids from Terra Nova Bay (30–500 m depth) and describes the depth distribution of these species. Preliminary observations of the summer reproductive condition of some of the species are also included.

Hexactinellids (glass sponges) are an understudied class with syncytial organization and poor procariotic associations, thought to lack defensive secondary metabolites. Poriferans, though, are outstanding sources of bioactive compounds; nonetheless, a growing suspicion suggests that many of these chemicals could be symbiont-derived. In Polar latitudes, sponges are readily invaded by diatoms, which could provide natural products. Hexactinellids are typical of deep waters; but in Antarctica, they dominate the upper shelf providing shelter and food supply to many opportunistic mesograzers and macroinvertebrates, which exert strong ecological pressures on them. Aiming to examine the incidence of defensive activities of hexactinellids against consumption, feeding experiments were conducted using their lipophilic fractions. Antarctic hexactinellid and demosponge extracts were tested against the asteroid Odontaster validus and the amphipod Cheirimedon femoratus as putative sympatric, omnivorous consumers. Hexactinellids yielded greater unpalatable activities towards the amphipod, while no apparent allocation of lipophilic defenses was noted. After chemical analyses on the lipophilic fractions from these Antarctic glass sponges, quite similar profiles were revealed, and no peculiar secondary metabolites, comparable to those characterizing other poriferans, were found. Instead, the lipidic compounds 5α(H)-cholestan-3-one and two glycoceramides were isolated for their particular outspread presence in our samples. The isolated compounds were further assessed in asteroid feeding assays, and their occurrence was evaluated for chemotaxonomical purposes in all the Antarctic samples as well as in glass sponges from other latitudes by NMR and MS. Characteristic sphingolipids are proposed as chemical markers in Hexactinellida, with possible contributions to the classification of this unsettled class.

The rates of oxygen consumption by embryos of antarctic echinoderms (Acodontaster hodgsoni, Odontaster validus, Psilaster charcoti, and Sterechinus neumayeri) were compared to the biomas (ash-free dry organic weight) of the egg of each species. These species could survive for months to years (range: 10 months to 5 years) by relying solely on the reserves present in the egg. However, certain species did not use any of the egg's reserves during early development. Embryonic stages of O. validus (a species with planktotrophic larvae) did not decrease in lipid, protein, or total biomass during the first 35 days of development. During the first 42 days of development, embryos of A. hodgsoni (a species with lecithotrophic development) used protein as an energy source. For both species lipid composed 40 to 50% of egg biomass, but was not used as an energy reserve. Larvae of O. validus have a high-affinity transport system for amino acids dissolved in seawater (K1 = 1.3 {mu}M for alanine). The rate of alanine transport from a low concentration (50 nM) could supply 32% of the larva's metabolic needs. This is a 10-fold higher input to metabolism than was determined (3% at 50 nM) for larvae of a temperate asteroid, Asterina miniata. Larvae of antarctic echinoderms live in an environment where the food supply is low for most of the year. Relative to their metabolic rates, antarctic larvae have larger energy stores and planktotrophic larvae have higher nutrient transport capacities when compared to larvae from temperate regions. These physiological differences allow antarctic larvae to survive for long periods without particulate food.

Two new asterosaponins, validosides A and B, were isolated from the butanolic extract of the starfish Odontaster validus and their structures established as (24S)‐24‐O‐[2,4‐di‐O‐methyl‐β‐xylopyranosyl‐(1→2)‐α‐arabinofuranosyl]‐5α‐cholestane‐3β,4β,6α,8,15β,24‐hexaol 6‐O‐sodium sulfate (1) and (24S)‐24‐O‐(3‐O‐methyl‐β‐xylopyranosyl)‐5α‐cholestane‐3β,6α,8,15β,24‐pentaol 2′‐O‐sodium sulfate (2) on the basis of extensive spectroscopic investigations and chemical correlations. Validoside A and B are extremely rarely occurring saponins due to the presence of a sulfated group at C‐6 of the aglycone in 1 and at C‐2′ of a pyranose in 2. The absolute configuration at C‐24 was found to be S by the application of the Trost‐Mosher methodology. Copyright © 1993 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim

Energy levels (x ± 1 SD) of whole soft body tissues and shells of the antarctic brachiopod Liothyrella uva (Jackson, 1912) were 9.0 ± 1.0 and 1.1 ± 0.4·kJ·g−1 dry wt, respectively. The majority of the energy in the soft body tissues was attributable to NaOH-soluble protein (17.4% dry wt) and in the shell to lipid (2.6% dry wt). An intact individual with a shell length of 3.4 cm and a wet and dry wt of 5.8 and 2.9 g, respectively, contained a total of 9.6 kJ. As dense populations of L. uva occur in the Southern Ocean, this species represents a considerable potential energy resource for predators, including sea stars and fish (ca 103kJ·m−2 in some areas). Total crude extracts of whole brachiopod soft tissues caused significant retraction of sensory tube-feet in 6 species of sympatric sea stars. Control assays employing whole crude extracts of the fresh soft tissues of the antarctic limpet Nacella concinna (Strebel, 1908) or seawater alone did not elicit significant sustained tube-foot retraction. Bioactivity was diminished in frozen extracts of brachiopod soft tissues exposed to the sensory tube-feet of the sea star Odontaster validus (Koehler, 1906), but was not diminished in the tube-foot retraction response of the sea star Neosmilaster georgianus (Studer, 1885). Lyophilized brachiopod soft tissues ground into a fine powder and embedded at a concentration of 2% (wt:vol) in agar pellets containing 5% krill caused significant feeding deterrence in an allopatric fish (the sheepshead minnow Cyprinidon variegatus Lacepede 1803). These findings suggest that, similar to temperate zone and tropical brachiopods, body tissues of this antarctic brachiopod are unpalatable to potential predators.

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.

Secondary metabolites are widespread among lower phyla and understanding their functional role(s) in the producing organism has been under study in recent decades. Considerable progress has been made in understanding chemical ecological interactions among terrestrial organisms, and similar research in the marine realm has been initiated in recent years. Polar regions are more difficult to access and thus progress has been slower. Nevertheless, the extreme and often unique marine environments surrounding Antarctica as well as the many unusual trophic interactions in antarctic marine communities might well be expected to select for novel secondary metabolites and/or novel functional roles for secondary metabolites. Indeed, recent studies have documented novel, chemically-mediated interactions between molluscs and amphipods, between algae, urchins and anemones, and between sponges and their predators. The Porifera are the dominant phylum on the McMurdo Sound benthos, and representatives of this phylum have been shown to elaborate sea star feeding deterrents, inhibitors of fouling or infectious organisms, and metabolites which mediate predation via molt inhibition. As a result of studies on Antarctic sponges, new insights into functional roles of pigments and the ability of sponges to sequester metabolites have been gained, and a new mechanism of chemical defense has been described. Herein we describe recent results of our studies of trophic interactions between sponges and their predators that are mediated by specific sponge secondary metabolites. Moreover, we highlight unusual chemically-mediated interactions in antarctic marine invertebrates other than sponges.

The mode of development was ascertained for 14 of the 16 species of sea stars known to occur in shallow waters of McMurdo Sound, Antarctica (77°51′S; 166°40′E). The species were collected between September 1984 and December 1985. Females of three species,Odontaster validus, O. meridionalis andPorania antarctica, spawn small to moderate eggs (0.17 to 0.55 mm), have a high fecundity, and produce feeding larvae. Females of an undescribedPorania species spawn a few eggs (150 to 310) that are 0.55 mm in diameter and develop into demersal non-feeding larvae. Females ofDiplasterias brucei andNotasterias armata produce a few (

As genomics converges with ecology and evolution to identify the fundamental linkages between genome structure and function, genome and transcriptome complexity will need to be measured in organisms from more diverse habitats, most often in the absence of complete sequence data. Here, we describe the complexity of ten genomes measured by a novel, high-throughput fluorescence-based kinetic hybridization assay. We applied the Shannon information index, H, and a related, fluorescence-adjusted index, Hf, as unique metrics of the hybridization kinetics to complement the conventional rate constant, k. A strong, positive relationship was present between Hf, and the repetitive DNA content of five eukaryotic genomes previously determined by Cot kinetic analyses (Onchorynchus keta, Ilyanassa obsoleta, Bos taurus, Limulus polyphemus, Saccharyomyces cerevisiae). This relationship was used to characterize the complexity of previously unstudied genomic samples in five metazoan taxa from three marine environments, including deep-sea hydrothermal vents (Alvinella pompejana), the temperate subtidal (Streblospio benedicti), and Antarctic coastal bays (Sterechinus neumayeri, Odontaster validus, Tritonia antarctica). Contrary to the predictions of nucleotypic theory, Antarctic invertebrates consistently had the lowest quantities of repetitive DNA in conjunction with low metabolic rates and highly protracted rates of cell division and larval development. Conversely, hydrothermal vent species with rapid cell division and growth do not have significantly different genome sizes or particularly low amounts of repetitive DNA as compared to non-vent, deep-sea taxa. Furthermore, there appears to be a positive correlation between the temperature at which the most abundant repetitive sequence classes anneal and habitat thermal stability. Thus, our study reveals a potential shift in repetitive sequence representation between these extreme environments that may be related to genome function in species living at these different thermal regimes.

Hexane, chloroform, and methanol extracts of 18 species of antarctic sponges were tested for their ability to induce sustained tube-foot retraction in the antarctic spongivorous sea starPerknaster fuscus. Extracts were imbedded in silicone and used to coat the tip of a glass rod, which was allowed to contact an extended tube-foot. Retraction times were measured and compared with three controls: contact with a glass rod coated with a hexane extract of fish (feeding stimulant), contact with the glass rod alone (mechanical control), and contact with the glass rod coated with silicone alone (silicone control). Only extracts of the spongeMycale acerata did not elicit significantly longer tube-foot retraction times than controls for at least one of the three organic extracts. Hexane sponge extracts elicited the lowest levels of significant tube-foot responses, with only 39% of the sponge species tested showing activity in this fraction. In contrast, chloroform and methanol extracts elicited a significant tube-foot retraction response in 73% and 78% of the species tested, respectively. This indicates that in this assay repellent metabolites are generally more polar substances. It remains to be determined that secondary metabolites are responsible for all of the tube-foot retraction responses detected in sea stars exposed to sponge extracts; bioactive secondary metabolites have been isolated from a number of these antarctic sponges. It may be of ecological significance that the two rapidly growing sponges,Homaxinella balfourensis andMycale acerata, were either not repellent or had low repellency, and thatM. acerata is the primary dietary item ofPerknaster fuscus.

A variety of Antarctic marine invertebrates contains secondary metabolites that may provide defense against potential predators. However, only in a few cases have tissues, extracts or isolated compounds of these invertebrates been tested against sympatric predators. The Antarctic nudibranch Bathydoris hodgsoni Eliot, 1907 contains hodgsonal, a compound only present in the external body (mantle tissues), which may protect the slugs from predators. To test this defensive hypothesis for hodgsonal, we carried out a series of experiments using the sympatric omnivorous seastar Odontaster validus Koehler, 1906 as a potential predator. Our experiments revealed that natural concentrations of hodgsonal elicit significant feeding deterrent responses in O. validus. Furthermore, hodgsonal is probably biosynthesized de novo by the nudibranch, since it was not detected in the viscera (as it should be in the case of a dietary compound), its concentration in the mantle (0.05–0.15% dry mass) is quite constant in individuals from different localities and depths, and its sequestration from a particular dietary source is unlikely because B. hodgsoni is an omnivorous feeder.

Parborlasia corrugatus McIntosh is a large, abundant, epibenthic antarctic nemertean which represents a considerable source of nutrition for predators. The energetic composition of adult body tissues (kJ·g −1 dry wt) is comprised primarily of energy derived from protein (11.4 kJ NaOH-insoluble, 6 kJ NaOH-soluble). Energy units associated with lipid (4.3 kJ) and carbohydrate (0.2 kJ) are much lower. Based on calculations of the energetic composition of whole body tissues, and a density of 0.3 ind·m −2 , mean population energetic density is estimated to be 65 kJ·m −2 . These values are often significantly higher as P. corrugatus aggregates when feeding. Despite their high abundance, nemerteans are not preyed upon and appear to be chemically defended. Sperm of the antarctic sea urchin Sterechinus neumeyeri (Meissner) were killed when exposed to a 3% aqueous extract of whole nemertean body tissues. In laboratory feeding experiments, two common species of antarctic fish showed significant rejection of nemerteans. Its toxic and feeding-deterrent characteristics are probably the result of the epithelial production of copious acidic mucus (pH = 3.5), although other toxic or noxious metabolites may be present. These results indicate that P. corrugatus , an important scavenger in antarctic benthic systems, is abundant, high in nutrients and energy content, and could be judged on this basis as a high-quality prey item. Nonetheless, due to its chemical defense, potential predators may avoid ingestion of this species.

The biochemical and energetic composition, population biology (size-weight relationship, abundance and size-frequency distribution) and chemical defense of the antarctic ascidian Cnemidocarpa verrucosa Lesson was investigated at McMurdo Sound, Antarctica, during the austral summer of 1989. The organic content (% organic material · g −1 dry tissue wt) of individual body components (tunic, body wall, endocarps, intestines, mature ovitestes and branchial basket) was generally high, with values ranging from 44.5% in the endocarps to 83.9% in the gonads. Most of this material was in the form of NaOH-soluble and insoluble protein. Tissue energy values ranged from 15.1 (tunic) to 22.4 (gonad) kJ · g −1 dry wt. Body height (cm) was positively related to body weight (g dry wt) by an exponential function. A representative individual (14 cm height, 550 g wet wt) contained a total of 493 kJ with most (75%) of this energy attributable to the body wall and tunic. The gonadal index [(kJ ovitestes · total kJ −1 ) x 100] for sexually mature individuals collected in November was 17.2 ± 4.7 ( n = 6). Population densities of C. verrucosa at depths of 20–30 m were 0.4 ind · m −2 at a site 3 km north of McMurdo Station. Energetic densities were estimated to be 197 kJ · m −2 . Size-frequency analysis revealed four modal peaks that probably represent distinct age cohorts, and may indicate predictable, annual recruitment events. Bioassays revealed that the tunic was noxious to sympatric pelagic and benthic fish, as well as an allopatric model fish. However, aqueous tunic extracts did not cause mortality in sea urchin sperm indicating the noxious compound(s) is not cytotoxic. pH values for body components were weakly acidic or neutral (5.86–6.93). Mature ovitestes were rejected by sympatric pelagic fish suggesting that gametes may be chemically defended. Although this common antarctic ascidian represents a significant resource of materials and energy, its tunic is tough and noxious, and probably provides an effective means of defense against potential predators.

Work on the life histories of common antarctic benthic marine invertebrates over the past several decades demands a revision of several widely held paradigms. First, contrary to expectations derived from work on temperate species, there is little or no evidence for temperature adaptation with respect to reproduction (gametogenesis), development, and growth. It remains to be determined whether the slow rates of these processes reflect some inherent inability to adapt to low temperatures, or are a response to features of the antarctic marine environment not directly related to low temperature, such as low food resources. Secondly, contrary to the widely accepted opinion designated as “Thorson's rule,” pelagic development is common in many groups of shallow-water marine invertebrates. In fact in some groups, such as asteroids,pelagic development is as prevalent in McMurdo Sound, the southern-most open-water marine environment in the world, as in central California. In other taxonomic groups, especially gastropods, there does seem to be a genuine trend toward non-pelagic development from tropical to antarctic latitudes. Although this trend has been predicted by theoretical models, its underlying causes appear to be group specific rather than general. Thirdly, pelagic lecithotrophic development, often considered to be of negligible importance, occurs in many shallow-water antarctic marine macroinvertebrates. Pelagic lecithotrophy may be an adaptation to a combination of poor food conditions in antarctic waters most of the year and slow rates of development. Nevertheless, some of the most abundant and widespread antarctic marine invertebrates have pelagic planktotrophic larvae that take very long times to complete development to metamorphosis. These species areparticularly prevalent in productive regions of shallow water (

Palatability of macroalgae can be affected by many different factors including thallus toughness, nutritional quality (carbohydrates, lipids, proteins, ash), and the presence of deterrent secondary metabolites. Palatability bioassays were performed using two different antarctic algal grazers (the antarctic rockfish Notothenia coriiceps and the sea star Odontaster validus). These data were then compared to toughness and chemical composition data. Toughness was determined using a penetrometer and a wide range of thallus toughness was observed in individuals of differing species as well as of the same species. Some of the tougher individuals required more than 30 N of force to cut through the thallus while others required less than 1 N to break through. Each algal species is also being tested for percent composition of carbohydrates, lipids, proteins, as well as ash. It is hypothesized that the algal species with greater nutritional quality and/or less thallus toughness are more likely to be acceptable foods to consumers (NSF OPP9814538, OPP9901076).

Aqueous extracts of the mantle tissues of the opisthobranchs Austrodoris kerguelensis and Tritoniella belli and the lamellarid gastropod Marseniopsis mollis were cytotoxic to the sperm of the antarctic sea urchin Sterechinus neumayeri, and caused behavioral responses (sustained terminal sensory tube-feet retractions) in five species of antarctic sea stars. Pieces of mantle tissue of all three species were noxious to two species of antarctic fish. Primary body components (mantle, foot, viscera) of the three gastropod species contained high levels (% dry wt) of NaOH-insoluble protein (40–59%), moderate levels of NaOH-soluble protein (7–25%) and lipid (6–18%), and low levels of carbohydrate (< 1%). The energetic composition of the tissues reflected their organic make-up, with most energy (11–17 kJg−1 dry wt) associated with protein. Total energy contents of representative adult individuals were 192,26 and 69 kJ for A. kerguelensis (93 g wet wt), T. belli (22 g wet wt) and M. mollis (114 g wet wt), respectively. Although these sluggish gastropods lack an external shell and are nutrient-and energy-rich and therefore high quality prey, like many of their temperate and tropical counterparts, they appear to be protected by chemicals.

Studies of the benthos between 30 and 60 m at Cape Armitage, McMurdo Sound. Antarctica, reveal an epifaunal community in which sponges and their asteroid and nudibranch predators predominate. Field experiments demonstrated that, with the exception of Mycale accrata, the growth rates of the sponges are too slow to measure in one year. Mycale, however, was observed to increase its mass as much as 67%. Because of its more rapid growth rate, Mycale appears to be the potential dominant in competition for substratum space, the resource potentially limiting to the sessile species. This conclusion is supported by observations of Mycale growing over and, in some cases, apparently having smothered many other sessile species representing at least three phyla. The densities and size frequency distributions of all the predators were measured; numerous feeding observations allowed an accurate appraisal of dietary compositions. Because of the predators' very slow consumption rates, however, direct measures of ingestion and its impact on prey populations were not possible. Estimates of the ingestion rates were derived from measurements of predator respiration rates, growth rates, and gonad growth. Data from the field surveys and the energetics studies suggest that Mycale is prevented from dominating the space resource by the predation of two asteroids. Perknaster fuscus antarcticus and Acodontaster conspicuus. Adult Perknaster specialize on Mycale, and the sponge provides a small proportion of the diet of A. conspicuus. Acodontaster conspicuus and the dorid nudibranch Austrodoris memurdensis are the most important predators on three species of rossellid sponges (Rossella racovitzae, R. nuda, and Scolymastra joubini). Despite this relatively heavy consumption and despite the fact that none of these sponges has a refuge in growth from potential mortality from A. conspicuus, very large standing crops of the rossellid sponges have accumulated. This accumulation appears to result from predation on larval and young A. conspicuus and Austrodoris by Odontaster validus, which is primarily a detrital feeder and apparently acts as a filter against the settlement and survival of the A. conspicuus and Austrodoris larvae. In addition, predation upon adult A. conspicuus by O. validus and the actinian urticinopsis antarcticus annually kills approximately 3.5% of the A. conspicuus population. This mortality exceeds the apparent rate at which A. conspicuus escape the larval filter.