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Journal ArticleDOI

Marine chemical ecology: chemical signals and cues structure marine populations, communities, and ecosystems.

25 Mar 2009-Annual Review of Marine Science (Annual Reviews)-Vol. 1, Iss: 1, pp 193-212
TL;DR: How chemical cues regulate critical aspects of the behavior of marine organisms from bacteria to phytoplankton to benthic invertebrates and water column fishes is reviewed.
Abstract: Chemical cues constitute much of the language of life in the sea. Our understanding of biotic interactions and their effects on marine ecosystems will advance more rapidly if this language is studied and understood. Here, I review how chemical cues regulate critical aspects of the behavior of marine organisms from bacteria to phytoplankton to benthic invertebrates and water column fishes. These chemically mediated interactions strongly affect population structure, community organization, and ecosystem function. Chemical cues determine foraging strategies, feeding choices, commensal associations, selection of mates and habitats, competitive interactions, and transfer of energy and nutrients within and among ecosystems. In numerous cases, the indirect effects of chemical signals on behavior have as much or more effect on community structure and function as the direct effects of consumers and pathogens. Chemical cues are critical for understanding marine systems, but their omnipresence and impact are inadequ...

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Journal ArticleDOI
TL;DR: The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research as mentioned in this paper, and both are only imperfect analogs to current conditions.
Abstract: Rising atmospheric carbon dioxide (CO2), primarily from human fossil fuel combustion, reduces ocean pH and causes wholesale shifts in seawater carbonate chemistry. The process of ocean acidification is well documented in field data, and the rate will accelerate over this century unless future CO2 emissions are curbed dramatically. Acidification alters seawater chemical speciation and biogeochemical cycles of many elements and compounds. One well-known effect is the lowering of calcium carbonate saturation states, which impacts shell-forming marine organisms from plankton to benthic molluscs, echinoderms, and corals. Many calcifying species exhibit reduced calcification and growth rates in laboratory experiments under high-CO2 conditions. Ocean acidification also causes an increase in carbon fixation rates in some photosynthetic organisms (both calcifying and noncalcifying). The potential for marine organisms to adapt to increasing CO2 and broader implications for ocean ecosystems are not well known; both are high priorities for future research. Although ocean pH has varied in the geological past, paleo-events may be only imperfect analogs to current conditions.

2,995 citations

Journal ArticleDOI
TL;DR: Evidence from direct and indirect approaches using geochemical and genetic techniques suggests that populations range from fully open to fully closed and a full understanding of population connectivity has important applications for management and conservation.
Abstract: Connectivity, or the exchange of individuals among marine populations, is a central topic in marine ecology. For most benthic marine species with complex life cycles, this exchange occurs primarily during the pelagic larval stage. The small size of larvae coupled with the vast and complex fluid environment they occupy hamper our ability to quantify dispersal and connectivity. Evidence from direct and indirect approaches using geochemical and genetic techniques suggests that populations range from fully open to fully closed. Understanding the biophysical processes that contribute to observed dispersal patterns requires integrated interdisciplinary approaches that incorporate high-resolution biophysical modeling and empirical data. Further, differential postsettlement survival of larvae may add complexity to measurements of connectivity. The degree to which populations self recruit or receive subsidy from other populations has consequences for a number of fundamental ecological processes that affect population regulation and persistence. Finally, a full understanding of population connectivity has important applications for management and conservation.

1,640 citations

Journal ArticleDOI
TL;DR: It is concluded that the best way to protect salt marshes and the services they provide is through the integrated approach of ecosystem-based management.
Abstract: Salt marshes are among the most abundant, fertile, and accessible coastal habitats on earth, and they provide more ecosystem services to coastal populations than any other environment. Since the Middle Ages, humans have manipulated salt marshes at a grand scale, altering species composition, distribution, and ecosystem function. Here, we review historic and contemporary human activities in marsh ecosystems—exploitation of plant products; conversion to farmland, salt works, and urban land; introduction of non-native species; alteration of coastal hydrology; and metal and nutrient pollution. Unexpectedly, diverse types of impacts can have a similar consequence, turning salt marsh food webs upside down, dramatically increasing top down control. Of the various impacts, invasive species, runaway consumer effects, and sea level rise represent the greatest threats to salt marsh ecosystems. We conclude that the best way to protect salt marshes and the services they provide is through the integrated approach of ecosystem-based management.

770 citations

Journal ArticleDOI
TL;DR: It is shown how the use of global variables of environmental forcing that have recently become available and gas exchange relationships that incorporate the main forcing factors will lead to improved estimates of global and regional air-sea gas fluxes based on better fundamental physical, chemical, and biological foundations.
Abstract: The past decade has seen a substantial amount of research on air-sea gas exchange and its environmental controls. These studies have significantly advanced the understanding of processes that control gas transfer, led to higher quality field measurements, and improved estimates of the flux of climate-relevant gases between the ocean and atmosphere. This review discusses the fundamental principles of air-sea gas transfer and recent developments in gas transfer theory, parameterizations, and measurement techniques in the context of the exchange of carbon dioxide. However, much of this discussion is applicable to any sparingly soluble, non-reactive gas. We show how the use of global variables of environmental forcing that have recently become available and gas exchange relationships that incorporate the main forcing factors will lead to improved estimates of global and regional air-sea gas fluxes based on better fundamental physical, chemical, and biological foundations.

626 citations

Journal ArticleDOI
TL;DR: The results imply that humans could be substantially impacting iron and bioavailable iron deposition to ocean regions, but there are large uncertainties in the authors' understanding.

562 citations

References
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Journal ArticleDOI
TL;DR: A selectively review of the literature on chemical cues for colonization in the sea, focussing on contrasts between positive (inducers) and negative (inhibitors, deterrents) cues and on prokaryote/eukaryote interactions.
Abstract: Colonization of surfaces in marine benthic environments is often one of the most significant moments in the life history of benthic organisms, representing, for example, a change from a planktonic to a benthic existence, a shift from a mobile to a sessile life form, or the initiation of pathogenesis. Many of the surfaces that are colonized are, in fact, other marine organisms, and in a general sense there is widespread evidence that specific chemical cues derived from marine organisms affect colonization by both marine prokaryotes and eukaryotes. However, detailed information for any one system on the nature of such cues, their distribution in situ, and their effects on the demography of colonizers is rare. Here, we selectively review the literature on chemical cues for colonization in the sea, focussing on contrasts between positive (inducers) and negative (inhibitors, deterrents) cues and on prokaryote/eukaryote interactions. We also consider whether generalized life history or natural history characteristics of colonizers (i.e., the mobility of a propagule, the extent to which a species is a habitat generalist or specialist, etc.) affect their response to chemical cues, and we touch briefly on some recent highlights relevant to the critical interplay between hydrodynamics and chemistry. A number of important methodological concerns are now being addressed through the introduction of field assays and analyses for chemical cues, and through molecular techniques for the characterization of microbial biofilms. These developments are encouraging, as is the increasingly multidisciplinary and cross-taxonomic approach to research in this area.

195 citations


"Marine chemical ecology: chemical s..." refers background in this paper

  • ...…pulchra produces halogenated furanones in surfaceassociated gland cells, moves these compounds onto its surface, and inhibits specific colonization phenotypes of marine bacteria (Maximilien et al. 1998, Steinberg et al. 2002), while having little or no effect on growth or survival of the bacteria....

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Journal ArticleDOI
TL;DR: Results of this broad survey cast doubt on the widely accepted notion that virtually all marine larvae are suitable prey for most generalized planktivores, and suggest that unpalatable larvae often may be aposematically colored.
Abstract: Risk of larval mortality is a critical component of models and debates concerning the ecology and evolution of the differing reproductive characteristics exhibited by marine invertebrates. In these discussions, predation often is assumed to be a major source of larval mortality. Despite limited empirical support, most marine larvae are thought to be palatable and broadly susceptible to generalist predators. Previous studies of larval—planktivore interactions have focused primarily on larvae that typically feed, grow, and develop for weeks to months in the plankton. Such planktotrophic species commonly produce large numbers of small larvae that disperse over vast distances. In contrast, the nonfeeding lecithotrophic larvae from sessile invertebrates that brood are often large and conspicuous, lack morphological defenses, and have limited dispersal distances because they typically are competent to settle minutes to hours after spawning. Interactions between lecithotrophic larvae and consumers are not well studied. This has limited the ability of previous authors to test broad generalities about marine larvae. We show that brooded larvae of Caribbean sponges (11 species) and gorgonians (three species) as well as brooded larvae of temperate hydroids (two species) and a bryozoan are unpalatable to co—occurring fishes. In contrast, brooded larvae of temperate ascidians (three species), a temperate sponge, and Caribbean hard corals (three species) are readily consumed by fishes, as are larvae from four of six species of synchronous broadcast—spawning gorgonians from the Florida Keys. Frequencies of survivorship for larvae attacked and rejected by fishes were high and statistically indistinguishable from frequencies for unattacked control larvae. Frequency of metamorphosis (when it occurred) of rejected larvae never differed significantly from that of unattacked control larvae. Assays testing for larval vs. adult chemical defenses for five species with distasteful larvae showed that larvae of all five species were chemically distasteful to fishes, whereas only three of five adult extracts deterred fish feeding. A comparison of larval palatability among chemically rich taxa showed that brooded larvae were significantly more likely to be unpalatable (86% of the species tested) than larvae of broadcasters (33%), and that palatable larvae were rarely released during the day (23%) while unpalatable larvae usually were (89%). Additionally, the frequency of bright coloration was high (60%) for unpalatable larvae and low (0%) for palatable larvae, suggesting that unpalatable larvae often may be aposematically colored. Results of this broad survey cast doubt on the widely accepted notion that virtually all marine larvae are suitable prey for most generalized planktivores. Among species that do not chemically or physically protect larvae against fishes, selection appears to favor the release of larvae at night, or the production of smaller more numerous offspring that grow and develop at sea as a way of escaping consumer—rich benthic habitats. Because distasteful larvae are not similarly constrained, distasteful species should exhibit reproductive and larval characteristics selected more by the fitness—related consequences of larval development mode and dispersal distance than by the necessity of avoiding benthic predators. Production of large larvae and retention of offspring in parental habitats that have proved to be suitable for growth and reproduction have both been proposed as advantageous, but these advantages often were assumed to be offset by losses due to increased larval apparency to fishes. This assumed trade—off is not mandatory because larvae can be defended chemically. Distasteful larvae tend to be conspicuous, localized dispersers that can co—occur with benthic fishes, and yet not be consumed.

189 citations


"Marine chemical ecology: chemical s..." refers background in this paper

  • ...Once larvae or embryos are released from brooding adults, they can be at considerable risk of predation in the plankton, but even more so as they recruit to the benthos where both fish and invertebrate predators are commonly concentrated (Lindquist & Hay 1996)....

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  • ...In other instances the larvae are chemically defended, but the adults are not and appear instead to rely more on physical/structural defenses (Lindquist & Hay 1996)....

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  • ...In a survey of numerous invertebrate larvae from the Caribbean and Western South Atlantic, Lindquist & Hay (1996) found that many were unpalatable and chemically defended from consumers....

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Journal ArticleDOI
TL;DR: It is demonstrated that taxonomically diverse ascidians from habitats characterized by intense predation pressure produce secondary metabolites that significantly reduce predation on both adults and larvae, and suggested that this defensive chemistry may be crucial in allowing the release of large, well—provisioned larvae during daylight periods when larvae have the greatest probability of using photic cues to select physically appropriate settlement sites.
Abstract: Previous investigations, focused primarily on vertebrates, have noted substantial losses of eggs and embryos to predators and questioned why selection has not more commonly resulted in the evolution of chemically defended eggs or embryos Hypotheses regarding the apparent rarity of such defenses have emphasized the potential incompatibility of actively developing tissues and toxic metabolites Alternatively, this apparent pattern could be an artifact of our greater knowledge of vertebrates, which in general show few tendencies for synthesizing defensive metabolites in either juvenile or adult stages In this study, we investigated adult and larval chemical defenses of a group of benthic marine invertebrates, the ascidians, in which the adults are often chemically rich, and we contrast our findings with what is known about chemical defenses of eggs and embryos from terrestrial and aquatic organisms Our findings suggest that there is no fundamental incompatibility of rapidly developing juvenile tissues and bioactive metabolites, and that chemically defended eggs and larval stages may be common among some taxonomic groups Ascidians are benthic invertebrates that often lack apparent physical defenses against predation, yet are common on coral reefs where predation by fishes is intense In contrast to most co—occurring invertebrates, many ascidians also release large, conspicuous larvae during daylight hours when exposure to fish predation would be highest Thus selection by predators might favor the evolution of distasteful larvae In situ observations indicate that many conspicuous ascidian larvae are distasteful to potential consumers We investigated the ability of secondary metabolites produced by taxonomically diverse ascidians from geographically distant locales to deter predation on both adults and larvae Larvae from the Caribbean ascidian Trididemnum solidum were distasteful to reef fishes, and when organic extracts of individual larvae were transferred onto eyes of freeze—dried krill (a good larval mimic in terms of size and color), these eyes were rejected by fishes while control eyes (solvent only) were readily eaten Larvae of the Indo—Pacific ascidian Sigillina cf signifera were also distasteful to coral—reef fishes and contained the unpalatable bipyrrole alkaloid tambjamine C When added to artificial foods at or below their natural mean concentrations and offered to consumers in field and laboratory feeding assays, the secondary metabolites produced by Trididemnum solidum (Caribbean Sea), Sigillina cf signifera (Indo—Pacific), and Polyandrocarpa sp (Gulf of California) significantly deterred feeding by co—occurring fishes and invertebrates Secondary metabolites produced by Trididemnum cf cyanophorum from the Caribbean Sea, Lissoclinum patella from the Indo—Pacific, and Aplidium californicum from the temperate Pacific, and the small stellate spicules common to many tropical didemnid ascidians did not significantly affect fish feeding High—pressure liquid chromatography (HPLC) analyses of six didemnin cyclic peptides in individual colonies of Trididemnum solidum from one patch reef at Little San Salvador, Bahamas found large inter—colony differences in their concentrations The mean concentration of didemnin B was more than double the concentration needed to significantly deter fish feeding in our field assays, and feeding tests with nordidemnin B showed that it deterred fish feeding across the entire range of natural concentrations HPLC analysis of the extract from a combined collection of T solidum larvae found adequate concentrations of didemnin B and nordidemnin B to account for their rejection by foraging fishes We demonstrate that taxonomically diverse ascidians from habitats characterized by intense predation pressure produce secondary metabolites that significantly reduce predation on both adults and larvae, and suggest that this defensive chemistry may be crucial in allowing the release of large, well—provisioned larvae during daylight periods when larvae have the greatest probability of using photic cues to select physically appropriate settlement sites Production of defensive secondary metabolites appears widespread among certain groups of ascidians, some of which are also known to concentrate acid and heavy metals as additional defensive strategies

188 citations


"Marine chemical ecology: chemical s..." refers background in this paper

  • ...In some species, both larvae and adults are strongly defended by similar metabolites (Lindquist et al. 1992)....

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Journal ArticleDOI
TL;DR: It is shown that waterborne cues from the copepod Acartia tonsa induce paralytic shellfish toxin (PST) production in the harmful algal bloom-forming dinoflagellate Alexandrium minutum.
Abstract: Among the thousands of unicellular phytoplankton species described in the sea, some frequently occurring and bloom-forming marine dinoflagellates are known to produce the potent neurotoxins causing paralytic shellfish poisoning. The natural function of these toxins is not clear, although they have been hypothesized to act as a chemical defence towards grazers. Here, we show that waterborne cues from the copepod Acartia tonsa induce paralytic shellfish toxin (PST) production in the harmful algal bloom-forming dinoflagellate Alexandrium minutum. Induced A. minutum contained up to 2.5 times more toxins than controls and was more resistant to further copepod grazing. Ingestion of non-toxic alternative prey was not affected by the presence of induced A. minutum. The ability of A. minutum to sense and respond to the presence of grazers by increased PST production and increased resistance to grazing may facilitate the formation of harmful algal blooms in the sea.

187 citations


"Marine chemical ecology: chemical s..." refers background in this paper

  • ...In a similar way, waterborne cues from the copepod Acartia tonsa induce paralytic shellfish toxin (PST) production in the bloom-forming dinoflagellate Alexandrium minutum (Selander et al. 2006)....

    [...]

  • ...Given the increasing instances of subtle chemical signaling that are being discovered in marine systems (e.g., Toth & Pavia 2000, Stachowicz 2001, Selander et al. 2006, Long et al. 2007), multitrophic level signaling could be common and important, but underinvestigated, in marine systems (Byrnes et…...

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Journal ArticleDOI
TL;DR: These compounds, including okadaic acid, brevetoxins, lyngbyatoxin A, caulerpenyne, bryostatins, and isocyano terpenes, are selected because their impacts on marine organisms and communities are some of the best-studied among marine natural products.
Abstract: Marine natural products play critical roles in the chemical defense of many marine organisms and in some cases can influence the community structure of entire ecosystems. Although many marine natural products have been studied for biomedical activity, yielding important information about their biochemical effects and mechanisms of action, much less is known about ecological functions. The way in which marine consumers perceive chemical defenses can influence their health and survival and determine whether some natural products persist through a food chain. This article focuses on selected marine natural products, including okadaic acid, brevetoxins, lyngbyatoxin A, caulerpenyne, bryostatins, and isocyano terpenes, and examines their biosynthesis (sometimes by symbiotic microorganisms), mechanisms of action, and biological and ecological activity. We selected these compounds because their impacts on marine organisms and communities are some of the best-studied among marine natural products. We discuss the ...

185 citations