165 The Chloroplast Genome of the Toxic Stramenopile Heterosigma Akashiwo (Raphidophyceae)

Many temperate green macroalgae contain secondary meatbolites that provide protection from grazing by some herbivores. These include the production of dopamine hydrochloride by the ulvoid green alga Ulvaria obscura and the production of dimethylsulfoniopropionate (DMSP) by many species of Ulvales and Caulerpales. The dopamine hydrochloride defense was isolated using bioassay-guided fractionation and is effective against sea urchins ( Strongylocentrotus droebachiensis ) and littorinid snails ( Littorina sitkana ). The DMSP activated defense system involves enzymatic cleavage of DMSP into dimethyl sulfide (DMS) and acrylic acid. It is found in many of the Ulvales and several species of Codium in the northeastern Pacific and Australasian regions. Many green algae such as Ulva fenestrata and Enteromorpha linza are avoided by urchins, which are deterred by DMS and acrylic acid in laboratory assays. However, these algae are often preferred foods of snails, which are deterred by DMS and acrylic acid. Snails may preferentially consume ulvoid green algae, despite being deterred by DMS and acrylic acid, because these algae contain relatively high nitrogen concentrations.     

164 Chemical Antiherbivore Defenses of Temperate Green Macroalgae

Many temperate green macroalgae contain secondary meatbolites that provide protection from grazing by some herbivores. These include the production of dopamine hydrochloride by the ulvoid green alga Ulvaria obscura and the production of dimethylsulfoniopropionate (DMSP) by many species of Ulvales and Caulerpales. The dopamine hydrochloride defense was isolated using bioassay‐guided fractionation and is effective against sea urchins (Strongylocentrotus droebachiensis) and littorinid snails (Littorina sitkana). The DMSP activated defense system involves enzymatic cleavage of DMSP into dimethyl sulfide (DMS) and acrylic acid. It is found in many of the Ulvales and several species of Codium in the northeastern Pacific and Australasian regions. Many green algae such as Ulva fenestrata and Enteromorpha linza are avoided by urchins, which are deterred by DMS and acrylic acid in laboratory assays. However, these algae are often preferred foods of snails, which are deterred by DMS and acrylic acid. Snails may preferentially consume ulvoid green algae, despite being deterred by DMS and acrylic acid, because these algae contain relatively high nitrogen concentrations.     

CONFUSING CONVERGENT MORPHOLOGIES: DIVERSITY AND DIFFICULTIES IN NEW ZEALAND ERYTHROPELTIDALES AND BANGIALES

Blooms of green macroalgae can devastate important finfish and shellfish habitats. Ulvaria obscura , a relatively unstudied green alga, is a major contributor to these blooms in the San Juan Islands, Washington State, USA. The biomass and productivity of this and other ulvoid algae were measured seasonally for two years. Experiments comparing the growth rates, responses to desiccation, photoacclimation, and grazer preference of U. obscura and Ulva fenestrata were conducted. Ulvaria blooms tended to occur in the subtidal while Ulva blooms were often intertidal. Both genera bloomed between late June and September. Despite their superficial similarity, Ulvaria and Ulva display markedly different physiological and ecological responses. Ulva was capable of faster growth, had higher rates of photosynthesis, and was more desiccation tolerant than Ulvaria. Ulvaria , however, appears to be more resistant to grazing than Ulva.     

Novel morphology in Enteromorpha (Ulvophyceae) forming green tides

"Green tides" are vast accumulations of unattached green macroalgae associated with eutrophicated marine environments. They have major ecological and economic impacts globally, so an understanding of their origin and persistence is required in order to make management decisions. Blooms predominantly consist of two common fouling genera of the Ulvales, Ulva (distromatic sheets) and Enteromorpha (monostromatic tubes). In the Baltic Sea and elsewhere green tides have increased over the last few decades. On the west coast of Finland, summer blooms consist of monostromatic sheets resembling Monostroma (Codiolales). We identified these as Enteromorpha intestinalis by comparative analyses of rDNA internal transcribed spacer 1 (ITS1), 5.8S, and ITS2 sequences, the first time monostromatic sheets have been found in the genus Enteromorpha. Ordinary attached E. intestinalis sporulated freely in culture, but the sheets reproduced only by cell regeneration into typical tubular thalli. The ITS sequences were identical to those of attached E. intestinalis populations in southwestern Finland, but differed by two substitutions from other Baltic sequences. We infer that this bloom originated from local attached populations and now reproduces clonally by fragmentation. This study provides further evidence of radical changes in gross morphology of green algae under eutrophicated conditions and the need for molecular identification.     

Direct and indirect effects of high pCO2 on algal grazing by coral reef herbivores from the Gulf of Aqaba (Red Sea)

Grazing on marine macroalgae is a key structuring process for coral reef communities. However, ocean acidification from rising atmospheric CO₂ concentrations is predicted to adversely affect many marine animals, while seaweed communities may benefit and prosper. We tested how exposure to different pCO₂ (400, 1,800 and 4,000 μatm) may affect grazing on the green alga Ulva lactuca by herbivorous fish and sea urchins from the coral reefs in the northern Gulf of Aqaba (Red Sea), either directly, by changing herbivore behaviour, or indirectly via changes in algal palatability. We also determined the effects of pCO₂ on algal tissue concentrations of protein and the grazing-deterrent secondary metabolite dimethylsulfoniopropionate (DMSP). Grazing preferences and overall consumption were tested in a series of multiple-choice feeding experiments in the laboratory and in situ following exposure for 14 d (algae) and 28 d (herbivores). 4,000 μatm had a significant effect on the biochemical composition and palatability of U. lactuca. No effects were observed at 1,800 relative to 400 μatm (control). Exposure of U. lactuca to 4,000 μatm resulted in a significant decrease in protein and increase in DMSP concentration. This coincided with a reduced preference for these algae by the sea urchin Tripneustes gratilla and different herbivorous fish species in situ (Acanthuridae, Siganidae and Pomacanthidae). No feeding preferences were observed for the rabbitfish Siganus rivulatus under laboratory conditions. Exposure to elevated pCO₂ had no direct effect on the overall algal consumption by T. gratilla and S. rivulatus. Our results show that CO₂ has the potential to alter algal palatability to different herbivores which could have important implications for algal abundance and coral community structure. The fact that pCO₂ effects were observed only at a pCO₂ of 4,000 μatm, however, indicates that algal-grazer interactions may be resistant to predicted pCO₂ concentrations in the near future.     

181 Two Lineages Within Griffithsia (Ceramiaceae, Rhodophyta) Based on Plastid Protein-Coding psaA, psbA and rbcL Genes

"Green tides" or blooms of ulvoid green algae are frequent in Yaquina Bay estuary on the central Oregon coast, USA. Measurements of their biomass were made from late spring to early winter in 1999 at six intertidal sites in the estuary, and were continued through the winter of 2002 at two sites that showed the greatest accumulation. The dominant blooming species were the Enteromorpha linza complex, Ulva fenestrata, E. flexuosa , and E. intestinalis . Red and brown algal abundance was negligable. Nutrients in the central channel were monitored, along with incident light and water column absorbence, and sediment temperature. The abundance of benthic amphipods was negatively correlated with the concentration of dissolved sulfides in porewater. However, dissolved sulfide levels were not correlated with green macroalgae biomass. Additional correlations were sought between the variables measured in an effort to suggest or negate major cause and effect relationships. This research is relevant to discerning the effects of eutrophication as a stressor on estuarine processes and could contribute toward an understanding of anthropogenic impacts on biological communities in coastal ecosystems.     

Propagule banks, herbivory and nutrient supply control population development and dominance patterns in macroalgal blooms

Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis . Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella . Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella . Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha , thereby negatively affecting Pilayella . Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.     

ULTRAVIOLET ABSORBING SUBSTANCES FROM AGARUM FIMBRIATUM (PHAEOPHYCEAE) PROTECT GRIFFITHSIA PACIFICA FROM UV RADIATION

Patterns of recruitment and abundance of Ulva pertusa and Enteromorpha linza in the three locations on the south coast of Korea where green tide frequently occurred were investigated with respect to the substrate profile, temporal and spatial variations, and their persistence. Artificial substrates (10 to 20 cm) were made of a mixture of a piece of granite rock and cement on ceramic tiles. These setting plates, representing different surface texture, were placed horizontally and vertically along the rocky shoreline, and replaced every 1 to 2 months. Both populations showed highly seasonal blooms, with peaks in October, 1998 to February, 1999 and another slight peak was observed in the fall of 1999 only for Ulva. However, the winter peak of 1998 did not appeared in the following year, indicating that a significant year-to-year variation existed. Regarding their settlement onto artificial plates with different roughness, Ulva preferred to settle on the rough surface both in the upper and lower intertidal zone; however, this pattern only appeared in the upper zone for Enteromorpha. Vertical distribution of the two populations was well separated; Enteromorpha was more abundant in the upper zone and Ulva in the lower zone. Their seasonal blooms were concomitant with ammonia levels, but occurrence was not well correlated with nitrate and phosphate. The ability of the populations to persist, once blooms occurred, was also distinct; Ulva persisted ca. twice as long as Enteromorpha , having a greater impact in the presence of green tide in this geographical area.     

Recent insights into oceanic dimethylsulfoniopropionate biosynthesis and catabolism

Dimethylsulfoniopropionate (DMSP), a globally important organosulfur compound is produced in prodigious amounts (2.0 Pg sulfur) annually in the marine environment by phytoplankton, macroalgae, heterotrophic bacteria, some corals and certain higher plants. It is an important marine osmolyte and a major precursor molecule for the production of climate-active volatile gas dimethyl sulfide (DMS). DMSP synthesis take place via three pathways: a transamination ‘pathway-’ in some marine bacteria and algae, a Met-methylation ‘pathway-’ in angiosperms and bacteria and a decarboxylation ‘pathway-’ in the dinoflagellate, Crypthecodinium. The enzymes DSYB and TpMMT are involved in the DMSP biosynthesis in eukaryotes while marine heterotrophic bacteria engage key enzymes such as DsyB and MmtN. Several marine bacterial communities import DMSP and degrade it via cleavage or demethylation pathways or oxidation pathway, thereby generating DMS, methanethiol, and dimethylsulfoxonium propionate, respectively. DMSP is cleaved through diverse DMSP lyase enzymes in bacteria and via Alma1 enzyme in phytoplankton. The demethylation pathway involves four different enzymes, namely DmdA, DmdB, DmdC and DmdD/AcuH. However, enzymes involved in the oxidation pathway have not been yet identified. We reviewed the recent advances on the synthesis and catabolism of DMSP and enzymes that are involved in these processes.     

Herbivory and patch dynamics on rocky reefs in temperate Australasia: The roles of fish and sea urchins

Sea urchins are widely considered to be the major grazers in temperate subtidal systems, with herbivorous fish being browsers of minor importance. This paper reviews spatial and temporal patterns in these herbivores on rocky reels in temperate Australasia, with the aim of assessing their relative impacts on patch structure and dynamics. Herbivorous fishes are widespread and make up a significant numerical component the reel fish fauna. Sea urchins are also abundant, but not all geographic locations support actively grazing species. Both fish and sea urchins exhibit distinct patterns of distribution among depth strata. Within depth strata, all herbivores are restricted to (sea urchins) or forage preferentially in (fish) particular habitat patches, causing a mosaic of different feeding activities. These patches are either related to specific features of the habitat (e.g. Kelp patches, topography) or behavioural interactions. Foraging by sea urchins and demersal-nesting damselfishes is intense and persistent, whereas in the kelp-feeding fish Odax cyanomelas, foraging reaches greatest intensity at predictable locations during a few months of every year. Many fish and sea urchins consume some algae in preference to others. However, feeding preferences may determine the nature of the impact only in fishes. For sea urchins, preference may occasionally determine the order in which algae are consumed, but at high densities they consume all available macroalgae. Impacts of both types of herbivore on the abundance of algae have been recorded. Some sea urchins (e.g. Evechinus chloroticus, Centrostephanus rodgersii) appear to severely modify biogenic habitat structure by maintaining ‘barrens’ (areas devoid of macroalgae) over long periods. In contrast to this, the effects of fishes may be more transitory (e.g. seasonal impact of Odax cyanomelas on brown algae) or occur at smaller spatial scales (e.g. nest sites maintained by male Parma victoriae) Herbivorous and other fishes appear to respond to spatial patterns in algal distributions, rallier than having it major impact upon them. The relative effects of fish and sea urchins on the long-term dynamics of kelp forests are unknown, hut temporal patterns in herbivore abundance and behaviour, and algal demography arc urgent targets for research.     

In situ tests on inducible defenses in Dictyota kunthii and Macrocystis integrifolia (Phaeophyceae) from the Chilean coast

Numerous experimental studies have reported inducible defenses in macroalgae, but most of them have been conducted in laboratory environments where algae were maintained detached from the substratum and in artificial flow regimes. The results of those experiments might not reflect the natural situation, which can only be studied in situ. We examined whether the brown macroalgae Dictyota kunthii (C. Agardh) Greville and Macrocystis integrifolia (Bory) show inducible defenses following exposure to different grazing levels (direct, water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing) in field experiments, striving to maintain natural conditions as much as possible. We measured palatability of algae after exposure to different grazing levels by using live algae and agar-based food containing non-polar extracts. M. integrifolia showed no induction of defenses (at least not of non-polar compounds), suggesting constitutive defenses, absence of defenses (tolerance) or use of another strategy to avoid herbivory. These results are similar to those from previous laboratory experiments. In D. kunthii, defense was induced after two weeks of direct grazing by amphipods under field conditions. Water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing did not induce defenses. Induction of defense in response to direct grazing agrees with results from a previous laboratory study, but while indirect cues induced defenses in the laboratory, there was no measurable induced defense in the field. Probably chemical cues from grazers are diluted quickly in the field, not reaching concentrations that cause induction of defenses. This might be the reason why in some algae induction by direct grazing is a more important defensive strategy than induction by water-borne cues. The results from our study also suggest that laboratory experiments showing induced defenses in response to grazed neighbours or mere grazer presence need to be interpreted with caution.     

Ontogenetic changes in the diet of the herbivorous Scartichthys viridis in a rocky intertidal zone in central Chile

Scartichthys viridis maintains a herbivorous diet following recruitment to rocky intertidal areas, where it consumes almost exclusively macroalgae. The sheet–like green macroalgae Ulva and Enteromorpha were the main items consumed by individuals <130 mm L _T. The tough branching red macroalga Gelidium made the bulk of the gut contents of specimens >220 mm L _T, Ulva being consumed to a much lesser extent. Further, Gelidium increased in importance in the total gut contents during ontogeny. In contrast, both small (70–120 mm L _T) and medium–sized (140–210 mm L _T) S. viridis individuals preferred Ulva in the laboratory. It is suggested that the increasing consumption of Gelidium along the ontogeny of S. viridis results from the limited availability of Ulva in the field. Large S. viridis individuals possessed longer guts relative to their body length, in comparison with small individuals.     

180 Relationships Between Water and Sediment Characteristics and Benthic Green Macroalgal Abundance in Yaquina Bay,Oregon: 1999–2002

“Green tides” or blooms of ulvoid green algae are frequent in Yaquina Bay estuary on the central Oregon coast, USA. Measurements of their biomass were made from late spring to early winter in 1999 at six intertidal sites in the estuary, and were continued through the winter of 2002 at two sites that showed the greatest accumulation. The dominant blooming species were the Enteromorpha linza complex, Ulva fenestrata, E. flexuosa, and E. intestinalis. Red and brown algal abundance was negligable. Nutrients in the central channel were monitored, along with incident light and water column absorbence, and sediment temperature. The abundance of benthic amphipods was negatively correlated with the concentration of dissolved sulfides in porewater. However, dissolved sulfide levels were not correlated with green macroalgae biomass. Additional correlations were sought between the variables measured in an effort to suggest or negate major cause and effect relationships. This research is relevant to discerning the effects of eutrophication as a stressor on estuarine processes and could contribute toward an understanding of anthropogenic impacts on biological communities in coastal ecosystems.     

Activated chemical defenses suppress herbivory on freshwater red algae

The rapid life cycles of freshwater algae are hypothesized to suppress selection for chemical defenses against herbivores, but this notion remains untested. Investigations of chemical defenses are rare for freshwater macrophytes and absent for freshwater red algae. We used crayfish to assess the palatability of five freshwater red algae relative to a palatable green alga and a chemically defended aquatic moss. We then assessed the roles of structural, nutritional, and chemical traits in reducing palatability. Both native and non-native crayfish preferred the green alga Cladophora glomerata to four of the five red algae. Batrachospermum helminthosum, Kumanoa holtonii, and Tuomeya americana employed activated chemical defenses that suppressed feeding by 30–60 % following damage to algal tissues. Paralemanea annulata was defended by its cartilaginous structure, while Boldia erythrosiphon was palatable. Activated defenses are thought to reduce ecological costs by expressing potent defenses only when actually needed; thus, activation might be favored in freshwater red algae whose short-lived gametophytes must grow and reproduce rapidly over a brief growing season. The frequency of activated chemical defenses found here (three of five species) is 3–20× higher than for surveys of marine algae or aquatic vascular plants. If typical for freshwater red algae, this suggests that (1) their chemical defenses may go undetected if chemical activation is not considered and (2) herbivory has been an important selective force in the evolution of freshwater Rhodophyta. Investigations of defenses in freshwater rhodophytes contribute to among-system comparisons and provide insights into the generality of plant–herbivore interactions and their evolution.     

Influences of green macroalgae blooms on nutrients and sulfide dynamics in hypereutrophic intertidal ecosystems

Limnology - Blooms of green macroalgae (Ulva spp.) occur frequently in the hypereutrophic Yatsu tidal flat of eastern Japan. In this study, to elucidate the effects of such blooms on nutrients and...     

PRELIMINARY STUDIES OF SEASONALITY,ECOLOGY, AND SPECIES COMPOSITION OF ULVOID ALGAL BLOOMS IN WASHINGTON STATE

Blooms of green macroalgae can devastate important finfish and shellfish habitats. Ulvaria obscura, a relatively unstudied green alga, is a major contributor to these blooms in the San Juan Islands, Washington State, USA. The biomass and productivity of this and other ulvoid algae were measured seasonally for two years. Experiments comparing the growth rates, responses to desiccation, photoacclimation, and grazer preference of U. obscura and Ulva fenestrata were conducted. Ulvaria blooms tended to occur in the subtidal while Ulva blooms were often intertidal. Both genera bloomed between late June and September. Despite their superficial similarity, Ulvaria and Ulva display markedly different physiological and ecological responses. Ulva was capable of faster growth, had higher rates of photosynthesis, and was more desiccation tolerant than Ulvaria. Ulvaria, however, appears to be more resistant to grazing than Ulva.     

INTRASPECIFIC VARIATION IN STRESS‐INDUCED HYDROGEN PEROXIDE SCAVENGING BY THE ULVOID MACROALGA ULVA LACTUCA1

We investigated the presence and kinetics of the oxidative stress response in intertidal and subtidal individuals of the ulvoid macroalga Ulva lactuca L. Stress responses, as measured with both enzymatic and fluorescent‐based antioxidant assays, differed between individuals collected from a subtidal and an intertidal habitat. Subtidal individuals secreted significantly more hydrogen peroxide (H₂O₂) than intertidal individuals when subjected to osmotic stress or desiccation. The activity of reactive‐oxygen‐scavenging enzymes and the ability to scavenge exogenous H₂O₂ were lower in subtidal than in intertidal individuals, suggesting that subtidal individuals are less stress tolerant. In vitro experimentation demonstrated that millimolar concentrations of dimethylsulfoniopropionate (DMSP) and its breakdown products could efficiently scavenge H₂O₂, with DMSP being a less‐effective scavenger than dimethyl sulfide (DMS), acrylic acid, and acrylate. The addition of H₂O₂ at concentrations of 2.5 mM or greater induced the cleavage of DMSP into DMS and acrylic acid in subtidal individuals. Intertidal individuals were affected in the same manner with the addition of 5 mM H₂O₂. There were no differences in the amounts of DMSP cleavage in subtidal and intertidal algae when the algae were subjected to hyposaline conditions. Our data suggest that the oxidative‐stress‐induced cleavage of DMSP affords products with efficient H₂O₂‐scavenging abilities. In addition, U. lactuca individuals growing in intertidal habitats are better acclimatized to changing environments and thus have a higher threshold for oxidative stress than conspecifics in subtidal habitats.     

Demethylation of Dimethylsulfoniopropionate and Production of Thiols in Anoxic Marine Sediments

Dimethylsulfoniopropionate (DMSP) is a natural product of algae and aquatic plants, particularly those from saline environments. We investigated whether DMSP could serve as a precursor of thiols in anoxic coastal marine sediments. The addition of 10 or 60 μM DMSP to anoxic sediment slurries caused the concentrations of 3-mercaptopropionate (3-MPA) and methanethiol (MSH) to increase. Antibiotics prevented the appearance of these thiols, indicating biological formation. Dimethyl sulfide (DMS) and acrylate also accumulated after the addition of DMSP, but these compounds were rapidly metabolized by microbes and did not reach high levels. Acrylate and DMS were probably generated by the enzymatic cleavage of DMSP. MSH arose from the microbial metabolism of DMS, since the direct addition of DMS greatly increased MSH production. Additions of 3-methiolpropionate gave rise to 3-MPA at rates similar to those with DMSP, suggesting that sequential demethylation of DMSP leads to 3-MPA formation. Only small amounts of MSH were liberated from 3-methiolpropionate, indicating that demethiolation was not a major transformation for 3-methiolpropionate. We conclude that DMSP was degraded in anoxic sediments by two different pathways. One involved the well-known enzymatic cleavage to acrylate and DMS, with DMS subsequently serving as a precursor of MSH. In the other pathway, successive demethylations of the sulfur atom proceeded via 3-methiolpropionate to 3-MPA.     

Dimethylsulfoniopropionate lyase from the marine macroalga Ulva curvata: purification and characterization of the enzyme

This is a report on the purification and characterization of an algal dimethylsulfoniopropionate (DMSP) lyase. This enzyme, also found in bacteria, is responsible for producing most of the dimethylsulfide (DMS) in marine environments. It was purified from the green macroalga, Ulva curvata (Kützing) De Toni. Initial in-vivo experiments showed that DMSP lyase activity from endogenous DMSP in Ulva increased for 24 h and then decreased as the culture aged and endogenous DMSP levels were depleted. When amended with exogenous DMSP, rates of DMSP lyase activity remained high even when the culture was 5 d old. Following disruption of the DMSP-depleted U. curvata cells by grinding, a soluble DMSP lyase was purified. This enzyme is a monomer of 78 kDa which has a K _m for DMSP of 0.52 mM. Soluble DMSP lyase had an optimum pH of 8 and an optimum osmotic strength of 75 mM NaCl. Following disruption of the algae by either grinding with sand or blending, and washing out the soluble enzyme, the green tissue, when treated with the non-ionic detergent, Triton X-100, solubilized additional DMSP lyase activity. Three hydrophobic variant forms of Ulva DMSP lyase were isolated and partially characterized from the detergent-solubilized activity. While the molecular and kinetic properties of the algal enzyme are different from the bacterial enzymes we purified earlier, both the soluble and membrane-bound forms did, nevertheless, cross-react with antibodies raised against the bacterial (Alcaligenes strain M3A) DMSP lyase.Abbreviations DMS dimethylsulfide - DMSP dimethylsulfoniopropionate This paper is dedicated to D.I. Arnon (1910–1995). We thank Dr. Richard Zingmark for helpful discussions on the speciation of the natural algal samples used in these experiments, and Robin Krest for collecting samples for us on numerous occasions. This work was supported, in part, by a grant from the University of South Carolina Venture Fund.