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This article was downloaded by:[California Digital Library -CDL (CRC journals only) Consortium] [California Digital Library -CDL (CRC journals only) Consortium] On: 21 February 2007 Access Details: [subscription number 758075943] Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Reviews in Fisheries Science Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713610918 The Effects of Harmful Algal Blooms on Aquatic Organisms Jan H. Landsberg a a Florida Marine Research Institute, Florida and Wildlife Conservation Commission, 100 Eighth Avenue Southeast, St. Petersburg, FL 33701.

To link to this article: DOI: 10.1080/20026491051695 URL: http://dx.doi.org/10.1080/20026491051695

Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article maybe used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 113 Reviews in Fisheries Science, 10(2): 113–390 (2002) 10(2): 113–390 Fisheries Science, Reviews in TABLE OF CONTENTS Petersburg, FL 33701 . a.b. Intracellular ...... 171 Extracellular c. Toxins (Exotoxins) or Exudates .... 173 Cell Surface Contact ...... 173 Biomagnification ...... 174 1. Intact Cells...... 171 2. Lysed Cells ...... 174 1. Trophic Transfer, , 1. ...... 2. 226 Prymnesiophytes ...... 232 Abstract ...... 114 A. Direct Exposure ...... 171 B. Indirect Exposure ...... 174 A. AcuteB...... 175 ChronicC...... 176 Organisms and Habitats Affected ...... 176 A. SaxitoxinsB...... 191 TetrodotoxinC...... 204 Spirolides...... D. 205 BrevetoxinsE...... 205 GymnodimineF...... 213 Okadaic G.acid and Derivatives (Dinophysistoxins) ...... Pectenotoxins...... 213 H. 216 CiguatoxinsI...... 217 J. Yessotoxins ...... 223 K. Azaspiracid ...... 224 Hemolysins...... 225 L.M. ...... 237 Anatoxins ...... 241 Aquatic Organisms 1064-1262 /02/$.50 ©2002 by CRC Press LLC The Effects of Harmful AlgalJan H. Blooms LandsbergFlorida Marine Research Institute, Florida Fish and Wildlife Conservation Commission, 100 Eighth Avenue Southeast, St on I.II. Introduction ...... III. 115 Harmful Algal Blooms ...... 117 Routes of Exposure...... 117 IV. Impacts ...... 175 V. Toxins and Harmful Mechanisms ...... 189 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ...... 256 ...... 257 114 spp...... 256 Olisthodiscus luteus This review provides an in-depth survey of the recorded incidences in aquatic 1. Anatoxin-a2...... 241 Anatoxin-a(s)3...... 242 Homoanatoxin-a ...... 243 1. 2. 3. 1. Dinoflagellates...... 2. 258 ...... 260 Compounds ...... 261 1. Dinoflagellates...... 2. 261 Diatoms3...... 271 Raphidophytes4...... 273 Prymnesiophytes5...... 273 Cyanobacteria...... 275 1. Diatoms2...... 278 Silicoflagellates3...... 280 Pelagophytes ...... 280 1. Dinoflagellates...... 2. 282 ...... 285 1. Dinoflagellates...... 2. 285 Ciliates ...... 287 N. MueggeloneO...... 243 CylindrospermopsinP...... 244 MicrocystinsQ...... 245 ...... R. 253 Reactive Oxygen ...... 255 S. Species with Multiple Toxins ...... 257 T. Species with Uncharacterized Toxins or Bioactive U. Harmful Species that Cause Mechanical Damage ...... 278 V. Harmful Species and Water Quality ...... 282 W. Suspected Species with Unidentified Mechanisms...... 285 Disease ...... 288 A. HABs as B.Potential Vectors ...... 288 HABs as Stressors in Disease ...... 289 Acknowledgments ...... 295 References...... 295 ABSTRACT: organisms of mortality and disease events suspected or known dinoflagellates, to of be caused species by microalgal 200 or Some mechanisms. harmful their or biotoxins, their blooms, ciliate diatoms, , prymnesiophytes, silicoflagellates, cilliates, currently known to be, suspected to be, or have the potential to be toxic or harmful to a wide and cyanobacteria are spectrum of organisms. This review summarizes the current information on toxic terrestrial affect that those or relevant, when (and, organisms aquatic harmful affect that species microalgal organisms, including humans), provides an updated list of such species, cites pertinent case VI. HABs as Potential Vectors for Pathogens and Stressors in VII. Parasites and PathogensVIII...... 292 Conclusions ...... 294 LANDSBERG REVIEWS IN FISHERIES SCIENCE

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blooms in the Nile Delta, Alexandria (Halim, minutum

harmful algal blooms, effects, aquatic organisms, toxins. Alexandrium Often referenced as one of the first toxic red reported to kill fish was that Often referenced as one of the first toxic red tides reported to kill fish was benthic and nonphotosynthetic harmful species that do not typically

1960). In addition, if the river was used incident for occurred drinking this in would freshwater imply that farther the upstream than in the Nile Delta; such varies in many summary papers (Sournia, 1995; Hallegraeff, 1995), in some cases because certain groups of aquatic microalgae are ignored. Some of increase the in the number of harmful species can apparent be attributed to the recent inclusion of being as described previously were that species of inclusion the to also and “bloom” benign, but that have now been found to be harmful. In freshwater systems, more cases of being poisoned by drinking water containing toxic cyanobacteria increase The 1996). (Carmichael, reported being also are blooms ) (blue-green in the number of species known improvements in our ability to accurately to identify HAB species and their toxins, be as harmful or toxic well as enhancements reflects in global monitoring and technological surveillance. Anthropogenic influ- ences interacting with natural processes have helped to increase the frequency of blooms, and the frequency (Hallegraeff, with 1995). These interactions have also which brought about an increase toxic in the species number are of formerly transferred benign globally species that environmental have or genetic become adaptations, or both. toxic because of altered described in the Old Testament of the Bible (Exodus 7:20-21): “all the water of the river was changed into blood. The fish in the river died and the river itself became so polluted that the Egyptians could not drink the water” (Moore 1977; Hallegraeff, 1995). Although this may be a reference to red in the river Nile, there is little historical or fossil (?) evidence that red tides occurred often in this region, except possibly for histories, and includes relevant information brackish, on and marine ecosystems. It is hoped that this review harmful will provide documentation and or toxic species reference from understanding our freshwater, material to critical questions research suitable future stimulate will for it that and alike researchers, professionals systems. aquatic upon students, have they effects managers, significant the and species and (HAB) bloom algal harmful resource of and health KEY WORDS: I. INTRODUCTION The recent increase in harmful algal blooms (HABs) in aquatic systems has begun to demonstrate the farreaching effects of aquatic these health and population growth, ecology, human blooms health, and ecosys- on species 1995; 1993, (Hallegraeff, economies and interactions, industries major on as well as integrity, tem most which about HABs, The 2000). Dolah, Van 1998; Burkholder, 1993; Steidinger, is known, tend to be planktonic, often visibly obvious, and quickly lead to acute of aquatic organisms. Some 5000 species of poisonings or mass mortalities marine microalgae have been described worldwide, and although less than 2% are now known to be harmful (Smayda, or 1990; toxic, Hallegraeff, that 1993, percentage 1995; appears Sournia, species to 1995). (including freshwater species) be The number increasing of aquatic Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 Gymnodinium (= brevis ., 2000). Cabeza de Vaca (1542) et al 116 h after consuming toxic (Vancouver, 1798, cited in Kao, 1993). 2 / 1 ) Hansen and Moestrup (Daugjberg In , before Europeans reached the Pacific coast, Native Americans Native coast, Pacific the reached Europeans before America, North In One of the earliest reports linking red tides to a may have been referring been have may kill fish a to tides red linking reports earliest the of One Before the 1980s, most HAB research concerned the species that pose a risk to algal harmful particular about information provided have reviews recent Several Columbia, Canada, in 1793. Several of the crew became sick and one crew member crew one and sick became crew the of Several 1793. in Canada, Columbia, died 5 breve reported that the Native Americans in the used to refer to the seasons the to refer to used Mexico of Gulf the in Americans Native the that reported as follows: “none of these peoples reckoned time by the sun or the moon, nor did they keep track of the month or the year. But they do understand and know about the different seasons when fruits ripen or fish die...” In Florida, red tides are most common in the fall and kill thousands of fish, many of which wash up along the beaches. were reputed to have watched the sea for the luminescence streaks of or red streaks water red during the If day dinoflagellates). by (caused night at luminescence and those warn to guards posted and mussels of taking the forbade chiefs the seen, were 1969). Sapeika, and Grindley in cited 1951, (Carson, dangers the with acquainted not consumed they if poisoned be could humans that knew Americans Native Although toxic shellfish (Kao, 1993, and references therein), early visitors to North America obviously did not. One of the shellfish poisoning incident involving some of first Captain Vancouver’s crew in British documented cases concerned the paralytic aquatic mortality event. to the seasonality of Florida red tides caused by Since that early report and because of threats to public health, a concerted effort has effort concerted a health, public to threats of because and report early that Since events. poisoning shellfish regarding knowledge public’s the improve to made been Shellfish, which can accumulate a variety of biotoxins, continue to pose a public health risk in coastal areas where toxigenic microalgae occur. Today, a wealth of information exists about effects longer-term potential the the of many but short-term fish, and shellfish human toxic of consumption health risks associated with the are not well understood, nor are the effects of exposure to routes, toxins for through example, via other aerosols. were organisms aquatic to posed risks the but mammals, other to and health human largely unknown. More research was also needed on two other important subjects: they but bioassays, in mammals small or humans to nontoxic are species HAB many produce that species microalgal the of and organisms, aquatic effect significantly can multiple toxic compounds only a few toxicity. of the compounds have been tested for affected organisms of groups specific the detailed have others and groups, or species Hallegraeff, 1988; Carmichael, (e.g., environments particular in HABs by or HABs by the overall 1993; Van Dolah, 2000). However, none of these reviews has addressed current state of knowledge regarding HABs and their effects on aquatic organisms. Because so much progress has been made in this area recently, I felt that it was a good opportunity to provide readers with a summary of the information available. LANDSBERG upstream areas would not be a common habitat for red tides per se. Others have theorized that the coloration and associated fish kills were perhaps caused by the Mediterranean the from ichthyotoxic, is which oxide, iron red of discharge volcanic into the Nile River (Wilson, 1985). inferences Even are this sometimes made historical about example the illustrates co-occurrence how of a “” and an REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 is not a microalga, blooms of this of blooms microalga, a not is 117 Mesodinium rubrum Mesodinium ., 1993; Steidinger, 1996; Hasle and Syvertsen, 1996; Throndsen, 1996; Syvertsen, and Hasle 1996; Steidinger, 1993; ., et al et ., 1995) and, except where relevant to this review, are not discussed not are review, this to relevant where except and, 1995) ., et al et The groups of harmful microalgae considered here include the dinoflagellates, III. ROUTES OF EXPOSURETypically, aquatic animals are exposed to toxic or harmful concentrations of there algae but web, food the dominate and bloom species epibenthic or planktonic when exposed being organism an of chances The exposure. of means obvious less also are species, the to a toxic or harmful species depends on the basic ecology of the HAB likelihood the and formation, bloom to conducive are that conditions environmental For the purposes of this review, harmful algal blooms include all aquatic species that species aquatic all include blooms algal harmful review, this of purposes the For are known to produce toxins or to cause harm, directly organisms or indirectly, or to aquatic to terrestrial organisms products. Toxic or harmful species can affect the full spectrum of living systems — associated with aquatic habitats from the biochemical to the or ecosystem level. For example, at the their ecosystem level, high concentrations of cells may interfere subsurface communities such as submerged aquatic vegetation. with At the biochemical light penetration and influence level, secondary metabolites produced by microalgae may interfere with particular cellular processes in the organism, but may not adversely affect the organism as a whole. In this review, species that have demonstrated effects only at the cellular or many interactions, species of terms In briefly. only mentioned be will level microbial microalgae may adversely affect other organisms through, for example, predator- prey relationships, but this aspect is also outside the scope of this review. diatoms, cyanobacteria, raphidophytes, prymnesiophytes, silicoflagellates pelagophytes, (Table aquatic and to 1). harmful or toxic be These to potential the have microalgae or be, to suspected include be, to known about 200 species that are organisms (Tables 2 to 7). Harmful effects may also extend to terrestrial organisms (including man) that are exposed to ciliate the aquatic Although shellfish). systems or to their products (e.g., This review summarizes the current information on species that affect aquatic organisms (and when relevant those that affect terrestrial toxic or harmful microalgal organisms, including humans), provides an pertinent updated case list histories, of and such includes relevant species, species from freshwater, brackish, information and cites marine ecosystems. I on hope that this review harmful or toxic provides documentation and reference material suitable for researchers, students, managers, and resource and health professionals alike, and that future it research will questions stimulate critical to our understanding significant effects of they have on HAB aquatic systems. species and the II. HARMFUL ALGAL BLOOMS species are often associated with harmful effects and pertinent thus information will on the be of included. these For groups, numerous references are (Skulberg available Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS Taylor 1993; further. The species names used here are those currently accepted in the literature; when it is appropriate, synonyms are provided.

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metabolites that act at the cellular level are produced by many of these species but are not included. not are but species these of many by produced are level cellular the at act that metabolites

is not a classification scheme and there are other species that may fit into these categories. Other secondary Other categories. these into fit may that species other are there and scheme classification a not is

are listed in the column text reference; NDIT = not discussed in text. Note this is just for the organization of the text, it text, the of organization the for just is this Note text. in discussed not = NDIT reference; text column the in listed are

(as *). Species discussed in the text under toxins/harmful mechanisms (section V) are listed under bolded headings that headings bolded under listed are V) (section mechanisms toxins/harmful under text the in discussed Species *). (as

= marine, B = brackish, F = freshwater. Action listed alphabetically, first by known action in the field, then experimental then field, the in action known by first alphabetically, listed Action freshwater. = F brackish, = B marine, =

experimentally, ** = human toxicity or harmful event, ?** = potential for event but not yet documented, *** = parasites. M parasites. = *** documented, yet not but event for potential = ?** event, harmful or toxicity human = ** experimentally,

LANDSBERG REVIEWS IN FISHERIES SCIENCE demonstrated only = * suspected, or unknown = ? experimentally, or field the in either action and reference, TABLE 1. Harmful and toxic aquatic microalgae and ciliates, habitat, toxins or bioactive compound produced, text produced, compound bioactive or toxins habitat, ciliates, and microalgae aquatic toxic and Harmful 1. TABLE

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LANDSBERG REVIEWS IN FISHERIES SCIENCE TABLE 1. (continued) 1. TABLE

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LANDSBERG REVIEWS IN FISHERIES SCIENCE TABLE 1. (continued) 1. TABLE

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124 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 1. TABLE

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126 TABLE 1. (continued) 1. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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128 TABLE 1. (continued) 1. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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130 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 1. TABLE

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132 TABLE 1. (continued) 1. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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134 TABLE 1. (continued) 1. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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organisms LANDSBERGaquatic on impacts REVIEWS IN FISHERIES SCIENCE health or ) or (wild mortalities documented and species 2. TABLE

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138 TABLE 2. (continued) 2. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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140 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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142 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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144 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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146 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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148 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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150 TABLE 2. (continued) 2. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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152 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 2. TABLE

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organisms TABLE 3. species and documented mortalities (wild or aquaculture) or health impacts on aquatic on impacts health or aquaculture) or (wild mortalities documented and species Diatom 3. TABLE

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aquatic organisms aquatic TABLE 4. Prymnesiophyte species and documented mortalities (wild or aquaculture) or health impacts on impacts health or aquaculture) or (wild mortalities documented and species Prymnesiophyte 4. TABLE Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

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158 TABLE 4. (continued) 4. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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160 TABLE 4. (continued) 4. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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aquatic organisms aquatic TABLE 5. species and documented mortalities (wild or aquaculture) or health impacts on impacts health or aquaculture) or (wild mortalities documented and species Raphidophyte 5. TABLE

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or health impacts on aquatic organisms aquatic on impacts health or TABLE 6. Silicoflagellates, pelagophytes, and ciliates and documented mortalities (wild or aquaculture) or (wild mortalities documented and ciliates and pelagophytes, Silicoflagellates, 6. TABLE Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

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on aquatic organisms aquatic on TABLE 7. Cyanobacteria species and documented mortalities (wild or aquaculture) or health impacts health or aquaculture) or (wild mortalities documented and species Cyanobacteria 7. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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166 TABLE 7. (continued) 7. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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168 TABLE 7. (continued) 7. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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170 TABLE 7. (continued) 7. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 et 171 ) or may accumulate at the surface if aerotopes if surface the at accumulate may or ) Planktothrix ) are present (Christoffersen, 1996). Many organisms associated with associated organisms Many 1996). (Christoffersen, present are ) Microcystis Most of the known toxic cyanobacteria are waterborne and may form massive In other cases, there may be no obvious mechanism by which some organisms may be no obvious mechanism by In other cases, there Numerous toxic microalgae have sedimentary cyst or resting stages as part of ., 1989). Depending on bloom density and toxin content, animals ingesting 1. Intact Cells A. DIRECT EXPOSURE a. Intracellular Toxins drinking by either toxins their and cells microalgal to exposed directly are Organisms them or ingesting them via various feeding modes (e.g., filter feeding, predation). , sponges, and shellfish that filter feed can take up toxic cells directly from the water column; many of Planktivorous fish these that actively organisms prey retain on toxic toxins microalgae can in Because filter feeders and also predators are not necessarily discriminatory, they can be the absorb toxins. viscera. exposed to most of the known major microalgal toxins (Tables 1, 7, 9, 10, 11). (e.g., blooms subsurface that a susceptible organism will come into contact with the species are benthic HAB. or planktonic, predatory Whether or photosynthetic, or HAB in a resting cyst phase will influence which communities of marine organisms might be affected by exposure to that HAB species. The degree of harm incurred may, in turn, depend on whether the organisms can detect blooms or toxins and then avoid them. For example, terrestrial animals such as cattle, sheep, preferentially they goats, — scums cyanobacterial surface avoid , necessarily not do wildlife wasps, and other drink water from these areas and become intoxicated (Lopez 1999). Rodas and Costas, could detect toxins, and so they may inadvertently consume toxic HAB species or prey. In these cases, chronic or sublethal effects major may indirect; or occur. direct either Exposure is toxins to V) Section harmful (see characterized and microalgae below. outlined are and 8 Table in listed are affected organisms of groups and routes (e.g., include organisms These cyanobacteria. with contact into come therefore may water , macrophytes, pelagic and benthic fish and , littoral inverte- brates, and waterfowl (Christoffersen, 1996), as well as amphibians, reptiles, and mammals. Toxic cyanobacterial blooms can also terrestrial organisms have that water the drink of a water surface the (Table along 7). devastating scum dense Poisoning a usually forms impact that does bloom not heavy on a occur is there unless that scum surface thin a form can cyanobacteria weather, calm In 1996). (Carmichael, frequently action wave and Wind lake. a of surface entire the over dispersed be may can concentrate cells along the shoreline (Carmichael, 1988). Wildlife or domestic animals therefore may have limited subsurface water from which to drink (Codd al between a few milliliters and several liters can experience acute or lethal toxicity (Carmichael, 1996). their life cycle. In some cases these may sink swimming planktonic forms. In other cases, toxic cells or adsorbed toxins stages may be more toxic than their free- Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

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172

currently characterized) microalgal toxins microalgal characterized) currently TABLE 8. Principle routes by which representative groups of organisms are exposed to harmful (as harmful to exposed are organisms of groups representative which by routes Principle 8. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 3 ., 10 × et al and as Heterocapsa Gyrodinium Heterocapsa Heterocapsa Favella circularisquama . Karenia mikimotoi Cylindrospermopsis H Gentien and Arzul, 1990) Arzul, and Gentien were more than 6.4 ., 1986). The inhibitory effect of the of effect inhibitory The 1986). ., spp. that have been dormant in the have been dormant spp. that et al et on the growth of both diatoms and the and diatoms both of growth the on Heterocapsa 173 (Shilo and Aschner, 1953). Release of exotox- of Release 1953). Aschner, and (Shilo cells increased, so did the likelihood of contact of likelihood the did so increased, cells ., 1995). Exudates from the toxic cyanobacterium toxic the from Exudates 1995). ., were also due to cell-to-cell contact. cell-to-cell to due also were Gyrodinium aureolum Gyrodinium (Huntley Alexandrium ., 1997). Compounds that inhibited feeding in the et al et et al seem to be mainly due to an extracellular product were derived from the extracellular products of the (Rausch de Traubenberg and Morlaix, 1995) and by the . (2000) (as (as (2000) . circularisquama circularisquama . et al et ., 1989). Redclaw crayfish accumulated by . (1995) proposed that the toxic properties of are on the cell surface. Cells of the dinoflagellate H adhered to the adoral zone of membranelles of Chrysochromulina polylepis Chrysochromulina Prymnesium cells and eventually died. The toxic effects of cells swelled, become immobilized, and eventually lysed (Kamiyama lysed eventually and immobilized, become swelled, cells Favella taraikaensis Favella Gonyaulax grindleyi Gonyaulax ., 1990; Myklestad 1990; ., et al cells. When concentrations of et al were immobilized immediately after contact with and by ingesting it (Saker and Eaglesham, 1999). sp. can inhibit feeding by cladocerans and (Ostrofsky pacificus et al et Favella ehrenbergii Favella Favella Prorocentrum lima Uchida Zooplankton are able to detect toxins via chemoreception. Soluble tintinnid cells/ml, dinoflagellate prymnesiophyte (Carlsson Anabaena c. Cell Surface Contact either microalgae, with contact cell-to-cell direct by affected be can organisms Many through exposure to microalgal toxins present gills the on penetrate the structures cell anatomical microalgal surface when or caused through damage mechanical the or skin of the exposed organism. and Arima, 1997). In another example, after direct contact with the brown tide to the bottom, are consumed by benthic organisms, and are then recycled into the of For example, cysts food chain. cells. vegetative the than toxic more times 10 to up are months several for sediments When newly formed, the cysts can even be up to 1000 times more toxic than the vegetative cells (Dale and Yentsch, 1978). Shellfish therefore could be exposed to high levels of toxins at all times if sediments are filtered during feeding. b. Extracellular Toxins (Exotoxins) or Exudates toxins extracellular as such exudates release microalgae many growth, active During (exotoxins) into the surrounding water, and organisms extracel- or exotoxins example, For cells. of absence apparent can the in even products be affected by such lular bioactive compounds are produced by the dinoflagellates Hansen in Hansen and prymnesiophyte ins by microalgae and the long-term persistence of toxins in water are determined by basic physicochemical properties that influence their stability. Release of toxins into a watery milieu may not always pose a threat to organisms if large volumes of water dilute the toxins. can be detected by copepods, and it has been suggested that these toxins may act as feeding deterrents (Shaw 1983; Burns raciborskii circularisquama instriatum circularisquama of concentrations with Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS cyanobacterium the containing water the from toxin the absorbing on the tintinnid the on circularisquama

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., et al were ., 1989; (Tracey, convolutus brevis et al . . C K , cause a type of at the sea surface, Mytilus edulis Mytilus concavicornis

Karenia brevis ., 1994). For this reason, treating have setae, barbs, processes, or ., 1995; Tester and Mahoney, 1995). Mahoney, and Tester 1995; ., Lyngbya majuscula et al et al et chunii, , adult blue mussels, mussels, blue adult ,

Chaetoceros 174 ., 1985). et al ., 1993; Kent 1993; ., Rhizosolenia ., 1995) and by causing them to reject food. When et al et and were inhibited from feeding (Bricelj and Kuenstner, 1989). et al

, ., 1989), Aureococcus anophagefferens Aureococcus ., 1989; Albright 1989; ., et al et al et Several planktonic species can release toxins that become aerosolized after lysis after aerosolized become that toxins release can species planktonic Several Blooms of the marine cyanobacterium Some diatom species, for example, 1. Trophic Toxin Transfer, Bioaccumulation, Biomagnification Biotoxins are transferred trophically when organisms consume other organisms that organisms other consume organisms when trophically transferred are Biotoxins have been exposed directly bioconverted, to or biomagnified toxic the toxins. microalgae In many cases, and the consumed has organism modified have the that toxins from the was form bioaccumulated, that was originally produced by B. INDIRECT EXPOSURE blooms with chemicals or by manipulating the environment can cause toxins to be released into the environment and ultimately may cause damage to aquatic organ- isms. transport Bubble-mediated transport. bubble-mediated in up caught become that or has been shown to concentrate from where concentrated toxins are subsequently released as an aerosol ( implicated in the death of goldfish in ponds along the Florida coast during a 1974 red tide (Quick and Henderson, 1974). 2. Lysed Cells Many species that normally produce environment under intracellular normal conditions. toxins Under release stressful of collapse and senescence during or currents, or action environmental wind change, salinity little (e.g., conditions into the a bloom), HABs release toxins as the cells lyse. For example, during senescence, cyanobacterial blooms release toxins into the (Schwimmer water and Schwimmer, that 1968; Rodger may cause fish to die Speare 1990). Terrestrial organisms and air-breathing mammals and versely reptiles affected can be by ad- these aerosolized toxins. Airborne toxins of LANDSBERG pelagophyte REVIEWS IN FISHERIES SCIENCE the by rejected usually are they chamber branchial the enter filaments cyanobacteria also it but feeding, cladoceran’s the interrupts only not rejection This postabdomen. causes nutritionally required that chamber has to be expelled collected (Gilbert, 1990). in the branchial Filaments or colonies of cyanobacteria can reduce the feeding rates of zooplanktonic of rates feeding the reduce can cyanobacteria of colonies or Filaments of the filtering appendages (Gliwicz and cladocerans by interfering with movement Siedlar, 1980; Haney 1988; Ward and Targett, 1989), (Gallager and larval bay , Skeletonema costatum spines that can become trapped in the gills of fish or shellfish, cause mechanical damage that impairs respiration, and eventually cause death (Parry contact dermatitis (swimmer’s itch) in humans swimming waters. or Symptoms include bathing itching, rash, in burning, affected blisters, and deep skin erosions that can be very painful (Fujiki

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 et ., 1989; Montoya , [OA] et al , azaspiracid [AZ]) (Table ., 1992], or PST for fish, birds, fish, for PST or 1992], ., ., 1996b). et al et et al Prorocentrum , paralytic shellfish toxins [PST] — and can also be transferred up the food chain food the up transferred be also can and domoic acid [DA]), Diarrheic Shellfish Diarrheic [DA]), acid domoic and Protoperidinium 175 Pyrodinium Dinophysis aeruginosa ., 1978; White, 1980; Geraci

, and et al Pseudo-nitzschia Microcystis Gymnodinium , brevetoxins [PbTx] and analogs), and, more recently, Azaspiracid , Karenia People who consume toxic tropical fish and become sick with tropical fish ., 1996]). IV. IMPACTS A. ACUTE Toxic HABs are usually planktonic and have acute rapidly effects. exposed When to high organisms concentrations of are these organisms toxic aquatic blooms, of mortalities shellfish-poisoning mass and problems, public-health significant events, can result. Exposure to a high concentration of toxin usually invokes an immediate biochemical or cellular response that may HABs of byproducts metabolic to exposed leads directly Animals change. behavioral to toxin of a concentration A effects. physiological, toxic minimize or avoid pathological, to ways different or in respond that thresh- above a certain threshold may be lethal, whereas concentrations below old may cause only a mild physiological, pathological, or behavioral response. In until they some cases, over a longer time period, organisms may accumulate toxins eventually exceed tolerable concentrations chronic and effects. The extent to which organisms will then accumulate toxins depends on stability, [lipophilic]), the soluble lipid or [hydrophilic] organism soluble water (either solubility the may suffer the microalgae. Filter-feeding zooplankton consume and retain toxic microalgae that microalgae toxic retain and consume zooplankton Filter-feeding microalgae. the are then passed one step up shellfish also consume the toxic microalgae and food accumulate the toxins, chain which in turn to zooplanktivores. become available to both animal and human Filter-feeding consumers. This transfer of toxins up the food chain is one of the most common ways in which including higher humans, trophic are levels, affected Shellfish Amnesic by poisonings: microalgal shellfish human toxins. with associated Several are microalgae that toxins produce specific diatoms (ASP, Poisoning Poisoning (DSP, dinoflagellates and dinophysistoxins [DTX]), Paralytic Shellfish Alexandrium Poisoning (PSP, dinoflagellates saxitoxins [STX] and derivatives), Neurotoxic Shellfish Poisoning (NSP, dinoflagel- lates Shellfish Poisoning (AZP, dinoflagellate Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1). These same toxins can also affect higher-level predators by a similar transfer of [Work birds for DA (e.g., chain food the up toxins and mammals [Armstrong al poisoning or ciguatera are also victims of indirect exposure to microalgal Some ciguatera toxins. toxins are biomagnified and some are biotransformed during their transfer up the food chain from toxic epiphytic dinoflagellates ingested by herbi- vores, to piscivores that feed on the ciguateric herbivores, and finally to consuming humans ciguateric piscivorous fish (Swift and Swift, 1993). The toxins found in benthic dinoflagellates are transformed from fish. into piscivorous in concentrated are a they that time the by form humans available to and harmful to cyanobacterium the and introduced into many trophic levels (Kotak

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., 1993; ., 1993; et al et al ., 1993; Anders and et al ., 1987; Harshbarger et al 176 ., 1988; Harshbarger et al Only recently has attention been drawn to the fact that microalgal toxins and Rarely are aquatic biotoxins considered to be etiological agents of tumor C. ORGANISMS ANDThe majority of documented HAB events are associated with , fish, and HABITATSbird mortalities, but there have also been AFFECTEDseveral accounts of other marine animal mortalities, such as of whales, dolphins, and sea turtles, for which biotoxins have Yoshimizu, 1994), yet biotoxins are rarely considered to be potential tumorigenic agents. B. CHRONIC health have received the algal species known to affect human For obvious reasons, affect lower vertebrate those algal species whose toxins severely most attention. Of and invertebrate species (many of them commercially important), most is known about the acute, lethal effects. Little information exists about the chronic, sublethal or lethal effects disease of to susceptible more bioaccumulated organisms render effects or such whether or biomagnified species animal algal toxins on human and and what is the fate of these toxins in the ecosystem. at all biological levels and to be recognized their chronic effects need to be studied as major threats to animal health, ecosystems (Shumway and Cucci, sustained 1987; Landsberg, 1995, 1996; Burkholder, 1998). fisheries, endangered species, long-term potential the that likely is it toxicants, and or contaminants other for true is As include will health public on or animals aquatic of health the on biotoxins of effects increased susceptibility to disease, immunosuppression, abnormal development, or long the in exposed are who levels trophic all at Animals neoplasia. of induction the term to biotoxins through the diet may die or display impaired feeding, avoidance behavior, physiological dysfunction, impaired immune function, reduced growth (Shumway, 1990; Luckenbach and reproduction, or pathological effects correlations defined clearly found have researchers cases, some In 1996). Landsberg, between the incidence of certain types of tumor and certain chemical contaminants or oncogenic (Malins LANDSBERGthe to species individual of tolerances Physiological toxins. the of toxicokinetics and various toxins will also determine the level of toxin accumulation. REVIEWS IN FISHERIES SCIENCE Wikfors and Smolowitz, 1993; Burkholder, 1998). the and distribution geographical widespread their given surprising is This induction. resultant probability that many aquatic animals are chronically exposed to tumor- promoting biotoxins. Several groups of toxins cyanobacteria have produced been shown to by have a variety dinoflagellates of short-term effects, nodularins, and Microcystins, and term. long these the in promoters tumor be also can toxins same lyngbyatoxin- and , , dinophysistoxin-1, acid, okadaic a have all been demonstrated to properties have only be been demonstrated experimentally tumorigenic Fujiki, in and small Sueoka mammals (Table 1996; or Humpage, cell and 1). Falconer 1993; Suganuma, These and (Fujiki assays tumorigenic 1998). The prevalence of tumors in aquatic animals has been worldwide steadily increasing for the last 30 years (Geraci

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., 1999) ., ., 1989; et al et et al Gyrodinium , ., 1998). For the et al ., 1997; Naviner 1997; ., Gymnodinium , et al et ., 1989). Unexplained fish kills and ., 1991). It is possible that many of et al et al Karlodinium 177 , ., 1995; Windust 1995; ., Karenia et al et ., 1992; Vidal and Gallo-Reynoso, 1996) may have been caused by ., 1990; Anderson and White, 1992; Hernández (Table 2). The wide variety of life strategies adopted by many HAB et al et al The sublethal effects of HAB species on molluscs (Table 9), zoop- HABs are responsible for numerous fish kills and disease events around the Traditionally, only planktonic HABs have been recognized as having acute Pfiesteria been implicated (Rounsefell and Nelson, 1966; Geraci, 1989; Geraci Hokama Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS purposes of this discussion, the organisms that HABs have been reported to affect will include zooplankton, macroinvertebrates, vertebrates, and, briefly, microalgae. A general chronology of mortality events, organisms affected, and the HAB species associated with the event is provided in Tables 2 to 7. When possible, organisms are listed both by their common and scientific names. These tables do not include general descriptions of red tides that are available in include the minor literature, nor fish do kills they or incidents identified. In many cases the reports simply document the during coincidental occurrence toxins for tested which necessarily not were animals multiple dead event; mortality a HAB and bloom a species of were or were bioassays conducted. Pertinent case histories of many of the larger-scale events are provided below. Allelopathic outlined. are 11) (Table plankton noncrustacean and 10), (Table lankton microbes other on toxins their or species HAB of effects antimicrobial or interactions Rao Subba (e.g., microalgae or will not be discussed, except for the brief reference to pertinent species in Tables lethal always not are zooplankton upon microalgae harmful of effects The 11. and 1 (Tables 10 and 11). Documented deleterious effects on zooplankton include avoid- ance; lethargic swimming or paralysis; grazing reduction or inhibition; starvation- induced mortality; and reduced rates of development, growth, and particularly on bivalves, have been covered short-term effects of HABs on shellfish, survival. The in extensive reviews by Shumway (1990, 1995). A summary of sublethal effects and by toxins specific of Effects 9. Table in outlined is molluscs on species HAB various harmful species are given below. world. At least 60 species are ichthyotoxic, and more than 30 species are harmful to fish (Tables 1 to 7). For many years, fish kills may have been attributed to the low dissolved-oxygen levels generated by by toxicity. For example, from 1980 to 1989, at necessarily considered to be caused high-biomass blooms were USA, Atlantic, South the in and 69% and Mexico of Gulf were the in kills fish of not 50% least attributed to low dissolved oxygen (Lowe or and marine mammal mortalities (e.g., Williams and Bunkley-Williams, Williams 1990; biotoxin transfer through the diet (Landsberg, 1995). species in aquatic systems suggests that fish in numerous affected. trophic niches can be effects, but benthic and food predatory the through HAB transferred be species can may biotoxins Other also diet. kill the through animals, exposure toxin usually by web and cause fish mortalities (e.g., White these kills were associated with harmful algal blooms caused by small, kills ephemeral fish Such toxic. being as recognized recently, until not, were that dinoflagellates bloom the with associated oxygen dissolved low the to attributed been have would blooms by caused be to known now are kills Many ichthyotoxicity. direct than rather as such dinoflagellates small of

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178 TABLE 9. Sublethal or chronic lethal effects of harmful microalgae on molluscs on microalgae harmful of effects lethal chronic or Sublethal 9. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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180 TABLE 9. (continued) 9. TABLE LANDSBERG REVIEWS IN FISHERIES SCIENCE

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181 TABLE 10. Sublethal or chronic lethal effects of harmful microalgae on crustacean zooplankton crustacean on microalgae harmful of effects lethal chronic or Sublethal 10. TABLE Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

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LANDSBERG REVIEWS IN FISHERIES SCIENCE TABLE 10. (continued) 10. TABLE

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184 LANDSBERG REVIEWS IN FISHERIES SCIENCE (continued) 10. TABLE

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Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . (2000). Some of the ., 1989). Such sublethal, et al et al 188 . (1995), and Duy et al . (1994), Yoo ., 1999) (see section on okadaic acid). Persistent algal blooms can et al et al The overall effects of HABs on food webs and ecosystems are probably the least the probably are ecosystems and webs food on HABs of effects overall The Surprisingly Surprisingly little information is available on the effects of HABs on reptiles, with associated are that birds) otters, raccoons, bears, (e.g., organisms Terrestrial chronic effects have farreaching impacts on animal and plant communities. cause severe shading, which ultimately can lead to the decimation of beds cause severe shading, which ultimately can lead to the decimation of seagrass (critical habitat for a multitude of species) (Dennison more unusual reports of animal mortalities associated with include cyanobacterial wildfowl, toxins honeybees, bats, and rhinoceros (Table 7). understood of all impacts. Not only are the pathways of complex algal-toxin transmission and incompletely known, long-term implications effects (e.g., recruitment failure and the subsequent loss of species from an ecosys- of sublethal, chronic tem, reduced filtration of water masses, and pelagic the coupling) subsequent are virtually effects unknown. on Long-term effects benthic- (e.g., (e.g., habitats many toxin In investigated. be accumula- to need also consumers higher-level on tion) coral reefs), there have been increased nutrient inputs and associated increases in macroalgal cover in recent years (Lapointe, 1989). An macrophyte increase in cover may macroalgal cover substrate or used increased The species. toxic these as of abundance the in substrate increases by toxic benthic microalgae also increase the distribution of toxic microalgal species, which would increase the can lead to likelihood they will be ingested by Potential herbivores effects browsing of through some the microalgae substrate. upon (Landsberg herbivores include tumor promotion LANDSBERG particularly sea turtles, alligators, and crocodiles, that are often exposed to toxins. Several cases of such effects have been documented (Tables 2 to 7), and it is likely that many more terres- and aquatic Both such analyses. biotoxin incorporate examinations mortality pathobiological or disease events will trial be birds are linked potential to victims of HABs reported HAB in as which events. the majority Numerous of deaths bird are associated kills with the have consumption toxic been of prey, for example, can succumb to toxins after toxins. In other cases, birds bioaccumulated microalgal fish or molluscs that drinking contaminated water, particularly after ingesting toxins from cyanobacterial have consumed or otherwise blooms in freshwater systems. Recently, biotoxins mammal mortalities have (Tables 2 been to linked 7), but to as was marine REVIEWS the biotoxins IN FISHERIES were case SCIENCE for with many years reptile overlooked mortalities, as possibly being the cause of marine mammal deaths. In those cases in which an infectious pathogen was immunosuppression implicated, and increased susceptibility to disease as a result of chronic toxin stress may compromised. be not may diagnoses final have investigations, mortality been in factor a considered. as included When microalgal biotoxins are not freshwater, marine, or estuarine food webs may also be at risk of toxin exposure from preying on cyanobacterial toxic which aquatic in freshwater prey. toxic However, drinking animals by those far are HABs the by largest affected terrestrial group blooms are occurring (Falconer, 1993). Animal mortalities caused by cyanobacteria have been reported since the late worldwide. 1800s Essentially, however, any animal can be at in risk because even a chance Australia (Francis, 1878) and encounter occur with contaminated systems examples can of terrestrial wildlife Carmichael result and in provided domestic are animal reviews in extensive intoxications and are exhaustive, not provided is illness in list This 7. Table or death. Selected (1992), Ressom

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al 189 ., 1998). At the cellular level (cytotoxic), toxins can be cytolytic, be can toxins (cytotoxic), level cellular the At 1998). ., et al et In general, the major producers of toxins are the dinoflagellate and diatom V. TOXINS AND HARMFULHarmful microalgae can produce a suite of toxic or noxious bioactive compounds or organisms other affect adversely can that ) or metabolites (secondary MECHANISMSprimary in involved those from distinct are metabolites secondary These ecosystems. functioning basic the in role key no have they because metabolism, intermediate and of the cell (Wright and Cembella, 1998). The role of secondary metabolites may be intrinsic (e.g., protection from UV light, intracellular nutrient storage, or a differen- tiation signal) or extrinsic (e.g., toxic to promoter predators, of an symbiotic allelopathic relationships, substance, a a [Plumley, metal 1997], scavenger or such as plankton as “pheromones” a [Wyatt and Jenkinson, 1997]). Extrinsic that compounds may have the potential to harm siderophore promote mating of gametes these of significance evolutionary or ecological the cases, many In organisms. other those are humans) (e.g., organisms “target” the when especially unclear, is products toxin-producing the as framework ecological same the of part normally not are that toxic in increase apparent the and byproducts these of role ecological The organism. 2000, Dolah, Van (e.g., discussed much been has world the around strains or species and references therein). Numerous environmental 1998; and genetic Cembella, factors and regulate Wright the 1991; Bomber, e.g., (see, toxins microalgal of production bacteria of role recognized The here. discussed not are and 1999) Jones, and Sivonen in the ability of some species to produce toxins (Doucette, 1995; Doucette Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1998) is important but is mentioned only pertain to when possible effects on aquatic such organisms (see section on bacterial-algal HABs as potential interactions vectors). groups in the marine environment and cyanobacteria in fresh and brackish water. Toxins are usually classified (1) by the level of other activity, or which humans can on range effects from their the by (2) and level, ecosystem the to level biochemical DNA affect and genotoxic are toxins microalgal level, biochemical the At mammals. (Huynh adducts hemolytic, antineoplastic, or tumor promoters. At the organ adversely can toxins level, phylum the level, At dermatotoxic. or hepatotoxic, neurotoxic, toxins can be affect certain groups of organisms, for example, they are , fungi), antimicrobial antialgal (algae), or ichthyotoxic (fish) (Table 1). In addition, (bacteria, some of these same toxins can be tumorigenic in the long term (Table 1). Microalgal toxins affect to able are biotoxins biotoxins; or cytotoxins as either distinguished usually are whole organisms (usually rodents) (Carmichael, in 1992). In others bioassays, whereas organisms, certain invertebrate and whereas vertebrate cases, of range depending a cytotoxins affect can on biotoxins their are mode microalgae not by of produced compounds action, bioactive Some groups. some certain only influence may affect seriously more may they but systems, mammalian on effects minimal have may aquatic organisms because of their mode of activity in an example, toxins aqueous from several dinoflagellate, prymnesiophyte, medium. or raphidophyte spe- For cies may affect fish (ichthyotoxic), but they apparently alone have cells of concentrations no high or toxins known level, ecosystem the At effect organisms. other on may have large-scale effects, but these effects are not usually so well defined as in narrower settings. In addition to the production of toxins, microalgae can harm

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al ., 1995b, 1996; Sogawa et al ., 1997), (3) those having minimal et al 190 ., 1995a; Mundt (Mahoney and Steimle, 1979; Adnan, 1989) (Table 2) (see 2) (Table 1989) Adnan, 1979; Steimle, and (Mahoney et al Noctiluca or ., 1998), (2) those having antimicrobial or antineoplastic properties (Patterson ., 1991, 1993; Hasui Harmful algal blooms are often considered to be directly responsible for In some cases, a toxic effect caused by a particular microalgal species may have may species microalgal particular a by caused effect toxic a cases, some In that are discussed are those that have In this review, the secondary metabolites Ceratium organismal effects that have been demonstrated only experimentally (Table 1), or (4) those acting as enzyme inhibitors (Namikoshi and Rinehart, 1996). toxic Terrestrial microalgae are not included. mortalities or In disease events that could many well be attributable to cases, toxins acting at the there may suborganismal level, but there is currently no field documentation of their possible be unexplained effects. However, possible correlations should certainly toxins be that are investigated. currently Certain classified as cytotoxic may organismal have chronic level, impacts but at again the this is unknown. Although determine lethality the is level often of used to toxicity, these tests bioassays and are therefore may often not necessarily made be correlated in with effects short-term on aquatic rodent organisms. indirectly observed aquatic mortality events, but in many cases such events may be due to the presence of the bloom rather than to direct toxicity. Changes in water chemistry as a result of a bloom may lead usually directly to associated mortalities, with but increased levels these of changes ammonia, are nitrite, or hydrogen sulphide toxicity, or anoxia caused by depleted oxygen levels (Mackenthun et al et al LANDSBERGaquatic organisms in other ways. Other metabolic byproducts, such as enzymes or not although and, systems organ particular or membranes cell affect can acids, fatty necessarily lethal, can affect animal health, perhaps ultimately causing disease or other sublethal effects. In other cases, anatomical structures such as setae, spines, or barbs can mechanically microalgae damage the to exposed organisms the of responses pathological or other physiological organisms. Often in these cases, may result in the pathology, disease, sublethal effects, or mortality. Organisms docu- mented to harm aquatic organisms through nontoxic mechanisms are included in Table 1. been only demonstrated experimentally experimen- by an using of whole presence the cells involving rather investigation than mortality a purified in Therefore, toxin. that by caused was mortality that infer often may scientists species, toxic proven REVIEWS tally IN FISHERIES SCIENCE mortality the that means which toxins, multiple produce species Many toxin. known combination a by toxin, individual an by caused be can species these with associated of toxins, or by other experimental uncharacterized impacts, a suite toxic of tests using whole compounds. cells, cell extracts, and In purified terms toxins of should be conducted studying whenever possible. or organisms aquatic to harm cause to suspected, strongly are or documented, been to terrestrial organisms that may be exposed to aquatic systems. Some secondary metabolites will not be discussed, but the species producing them are identified in the following categories: (1) those causing only Table 1. Those metabolites fall into cytotoxic effects (e.g., Nagle and Gerwick,1995; Hasui 1948; Prescott, 1948). Fish or invertebrate kills have been attributed to the low levels low the to attributed been have kills invertebrate or Fish 1948). Prescott, 1948; of dissolved oxygen associated with nontoxic, high-biomass plankton blooms such as section on water quality).

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., ., g/ µ , and et al et al Pyrodinium , and , Lyngbya wollei Anabaena circinalis , has been in flux, but , the species are discussed Alexandrium in mice (intraperitoneally [i.p]) of 10 of [i.p]) (intraperitoneally mice in Alexandrium 50 191 Gymnodinium catenatum Gymnodinium for sodium cyanide at 10 mg/kg (Oshima 50 species, Cylindrospermopsis raciborskii — and by the cyanobacteria , Alexandrium compressum sp. (Table 1). The taxonomy of var. Most humans who experience paralytic shellfish poisoning (PSP) have con- Principally because of their effects on human health, toxins rather than the HAB the than rather toxins health, human on effects their of because Principally Aphanizomenon flos-aquae Planktothrix recently both morphospecies and genospecies have been studied (Costas here using the currently accepted taxonomy (Balech, 1995); they are listed in Table here using the currently accepted taxonomy 1 with their synonyms. sumed toxic bivalves (Shumway, 1990), (Shumway, but 1995), and rarely occasionally toxic fish (Maclean, 1979; toxic Adnan, 1984), gastropods and can also cause this sickness. Numerous globally fatal (Kao, 1993), but the cases recent successful implementation of of programs moni- PSP have been toring the presence reported of PST-producing microalgae in many countries has helped to minimize the risks to humans. Thus far, all human cases of PSP have been caused by toxic dinoflagellates, and the geographical distribution of these cases is related to the global distribution of the various PST-producing species and their toxigenic LD an having lethal, highly are PST strains. 1995; Scholin, 1998). For the previous purposes mortality reports associated with of this discussion and in documenting kg (when compared with an LD A. SAXITOXINS Neurotoxic paralytic shellfish toxins 11 (PST) approximately are produced by dinoflagellates — species are usually considered according to their involvement in particular poisoning particular in involvement their to according considered usually are species events (e.g., saxitoxins in paralytic shellfish poisoning, brevetoxins in neurotoxic shellfish poisoning). Because microalgal species because may the same effects produce the of toxins descriptions event, one multiple in can implicated be can toxins, species be harmful numerous produced by species microalgal numerous lists 1 Table similar. species, very or identical are species and these of many of because prymnesiophytes, raphidophytes, diatoms, (dinoflagellates, microalgal by alphabetically the describes and group ciliate or cyanobacteria) and pelagophytes, silicoflagellates, or human on effects potential toxins’ the and species individual by produced toxins animal health. This table provides a comprehensive harmful list or of toxic the species; currently it known also includes those species demonstrated whose toxins experimentally have been to have on effects documented have they unless provided not is species activity these of discussion at the cellular level, aquatic organisms in the wild. In the following section, species are grouped by the but further toxins or bioactive compounds that they produce. Species that produce toxins are discussed under each appropriate multiple section, as outlined in Table 1. Where follows the order of the groups outlined in possible, the description of these toxins Table 1, except when multiple species are involved. bahamense 1989c). PST are potent that bind to site 1 sodium on channel, block the the influx of voltage-dependent sodium into excitable cells, first and numbness, restrict signal and paresthesia are PSP of Symptoms neurons. between transmission around the lips and mouth and then involving the face and neck; muscular Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., and et al et sp. was sp. ., 1988; et al tamarense ., 1988b, 1990a, 1988b, .,

., 1995; Teegarden 1995; ., et al et ., 1977), but corrobo- but 1977), ., Cochlodinium et al et et al et ., 1995), yet many species many yet 1995), ., type ’78) has been shown Alexandrium et al et ). In 1977, In ). ., 1996), produce sublethal effects ., 1996). Some of the PST derivatives ., 1998). A bacterial origin for PST has PST for origin bacterial A 1998). ., -sulfocarbamoyl toxins B1 (GTX5), B2 et al Cochlodinium N et al et et al ., 1998) (see section on HABs as vectors). 192 (as Cochlodinium et al ., 2000). Exposing euphausiid, lobster, and crab ., 1990, 1994). Because the toxin profiles of PST- et al et al ., 1993; Onodera et al ., 1990; Doucette et al (Table 2). The fact that PST are produced by several species of Cochlodinium polykrikoides The transport of PST through the food chain and the accumulation of toxins in PST are present in a wide range of aquatic organisms, and they have been PST comprise the saxitoxins (STX) and at least 21 derivatives (Oshima, 1995) that 1995) (Oshima, derivatives 21 least at and (STX) saxitoxins the comprise PST Even though numerous microalgal species have been documented to produce catenella . (Tables 10 and 11), or cause active avoidance (Turriff avoidance active cause or 11), and 10 (Tables and Cembella, 1996a; Turner larvae to PST did not produce reported. were any effects apparent adverse short-term effects studies in other in two yet studies 1991a), Robineau (Yazdandoust, 1985; Exposure to PST may be lethal (Bagøien Anderson, 1990; Anderson producing dinoflagellate species differ the exposure dose and highly the toxic proportion PST derivatives of to which animals are exposed will differ (Landsberg, 1996). become toxins which by mechanisms important as identified been have zooplankton Turriff 1981a; 1979, (White, levels trophic higher to available A cyanobacteria in freshwater systems suggests that aquatic and terrestrial animals also animals terrestrial and aquatic that suggests systems freshwater in cyanobacteria have a good chance of being exposed to these toxins. Toxin composition can vary environmen- with location, geographical with strains; and species microalgal among tal factors, and under different experimental conditions (Cembella LANDSBERG drowsi- ataxia; motor incoordination; and floating; sensation of lightness weakness; doses, high in and efficiency; ventilatory decreasing progressively incoherence; ness; respiratory paralysis and death (Catterall, 1985; Kao, 1993). (Sakamoto absent apparently were dinoflagellates when occur to documented REVIEWS IN FISHERIES SCIENCE in various combinations and concentrations have been natural associated with toxigenic PSP. dinoflagellate No or cyanobacteria population contain all has naturally occurring PST been derivatives, so found the toxin profile to (i.e., the toxin to be characteristic of the microalgal strain or components produced) is considered species (Cembella 1992). Knowledge of the widespread distribution of PST and results of a series of experimental studies have led to the conclusion that in many cases dinoflagellates (Doucette PST of source only the not are been demonstrated, and bacteria also play a significant (but not exclusive?) role in (Kodama species dinoflagellate certain in PST of production the 1990b; Ogata are highly toxic (as -blocking agents in mammals) and include the carbamate toxins, (STX), neosaxitoxins (GTX1-4). (NEO), The and decarbamoyl the gonyautoxins analogues deoxydecarbamoyl analogues (dcSTX, (doSTX, doGTX2, doGTX3) are dcNEO, of intermediate tox- dcGTX1-4) and icity. the The least toxic derivatives are the (GTX6), and C1-C4 (Sullivan, 1993; Oshima, 1995). Although not usually associated with PSP, to produce two unique, zinc-bound, toxins/ multiple (see neosaxitoxin-like 1989b) Nozawa, compounds (Onoue and (Reyes-Vasquez Venezuela in outbreaks PSP in implicated rative evidence is lacking. mortality of majority the organisms, aquatic to dangerous potentially are all and PST reports have been associated with the dinoflagellates

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., to et al et when ., 1995; ., 1995), ., ., 2000). Tigriopus ., 2000). et al et al et et al Alexandrium (Schantz et al tonsa tissues after 12 h. ., 1985b). In experi- and et al ., 1986; Turriff 1986; ., typicus ., 1985b; Turriff herdmani . et al et E et al and Saxidomus gigantea Saxidomus g PST/g dry weight. The toxin profile toxin The weight. dry PST/g g , they accumulated toxin readily over µ toxins is species specific. Copepods have Copepods specific. species is toxins ., 1995; Teegarden and Cembella, 1996a; 193 herdmani et al ., 1996) (see section on uncharacterized toxins and toxins uncharacterized on section (see 1996) ., Calanus finmarchius Alexandrium et al et Alexandrium catenella ., 1995; Teegarden and Cembella, 1996a; Turner gSTXeq/100 g were measured in et al µ were rejected by copepods (Huntley copepods by rejected were ., 1990; Shumway and Cembella, 1993; Bricelj and Shumway, 1998a, , a range of responses was noted. Physiological reactions of some ). , profiles of the toxins accumulated by these copepods were dramati- ., 1996; Teegarden and Cembella, 1996a). When the copepods the When 1996a). Cembella, and Teegarden 1996; ., , indicating some toxin transformation (Boyer ., 1995; Bagøien 1995; ., were fed ., 2000). When copepods were exposed to toxic and nontoxic strains of were exposed to toxic and nontoxic ., 2000). When copepods et al et al et et al et et al Because most outbreaks of PSP result from eating shellfish, most studies on PST on studies most shellfish, eating from result PSP of outbreaks most Because Regardless of the level at which zooplankton feed on or are exposed to toxic tamarense tamarense catenella tamarense Alexandrium . . . . mental exposures of the copepods A cally different from the toxin profiles of the dinoflagellates and had patterns more consistent with metabolic transformation than differential retention. Concentrations of up to 2332.3 compared with other taxa in the zooplankton community (Turner of the copepods showed higher levels of GTX2 and GTX3 than originally found in A californicus 12 of saturation a reached and period 3-day a That such high levels of PST zooplankton act as toxin vectors were (Teegarden and Cembella, 1996a). Recently, such accumulated reinforces the hypothesis a finding was confirmed in the field where PST concentrations were disproportion- that ately high in the copepods the Globally, resources. economic consumable, are that species vector those concern presence of PST has been documented in numerous and species biotransformation, exposure, of occurrence, the on molluscs. available are Several reviews extensive effects of PST in molluscs, especially in bivalves and gastropods (Shumway, 1990; Shumway was STX here. provided be will effects PST of survey brief a only therefore, 1998b); butterclams, Washington toxic from isolated first Bagøien Bagøien 1957; Schantz, 1960). Because STX and its derivatives block only voltage-gated A algae, the accumulation of saxitoxins in zooplankton and the subsequent transfer- ence of the toxins by zooplankton into the are well documented (White, 1981a, b; Turriff of zooplankton accumulate PST and act as vectors in transferring toxin up the food in transferring PST and act as vectors accumulate of zooplankton chain (White, 1981a, 1981b; Turriff Turner A copepods included paralysis (Ives, 1985, 1987), whereas other copepods enced experi- no obvious adverse effects (Teegarden and Cembella, 1996a). Toxic strains of ingested at lower rates in response to ingested at increasing a toxicity high rate (Ives, similar 1985, to 1987), the rate or at (Teegarden and which Cembella, 1996a). nontoxic This range strains of responses are suggests that ingested suscepti- to species copepod of bilities capable are they abilities; selective and chemoreceptive specific and sensitive highly of selecting preferred prey from mixed absence algal of populations, PST and is not the necessarily presence a or factor Cembella, in 1996a, grazing 1996b). The selectivity differential (Teegarden response and of copepods to species may also be due in part to (Turriff the presence of other bioactive compounds Although the rates at which copepods feed on PST-producing dinoflagellates may PST (Boyer vary, copepods can still bioaccumulate Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , , ., ., and et al et al ., 1990, et al , may retract nerve sensitivity nerve gigas ., 1993; Cembella Mercenaria mercenaria Mercenaria in vitro in Mya arenaria et al Alexandrium catenella ., 1971). Only in recent years have ., 1971). Only in recent ., 1994; Bricelj and Cembella, 1995; ., 1996; Bricelj and Shumway 1998a, et al ., 1985, 1987; Shumway and Cucci, 1987; Cucci, and Shumway 1987; 1985, ., et al ., 1993; White et al , northern quahogs, northern , 194 et al et et al ., 2001). In more tolerant species, if PST are not are PST if species, tolerant more In 2001). ., ., 1983; Beitler and Liston, 1990; Bricelj et al et ., 1999). et al Alexandrium et al These molluscan mortalities also usually occur in conjunc- spp. and PST can respond by a variety of mechanisms. Such mechanisms. of variety a by respond can PST and spp.

., 1993; Shumway ., 1992; Chebib ., 1985; Shumway 1985; ., et al ., 2000; Smith 2000; ., et al et al et (Table 2). et al et ., 1996; Lassus Alexandrium et al The fate and distribution of PST in bivalves varies according to HAB bloom Of the known PSP-producing microalgae, documented mass mortalities of tamarense . mechanisms include increasing mucus and pseudofeces production, modifying valve modifying production, pseudofeces and mucus increasing include mechanisms altering behavior, burrowing or feeding reducing rate, filtration changing and activity consumption oxygen or activity cardiac changing and production, byssus of rate the (Cucci 9) (Table reduce pumping rates (Dupuy and Sparks, 1968). Bivalves that are exposed for short for exposed are that Bivalves 1968). Sparks, and (Dupuy rates pumping reduce to periods 1992; Shumway, and Marsden 1990; Shumway, 1988b; 1988a, Shumway, and Gainey Bricelj intrapo- species, exposure; of history prior conditions; environmental characteristics; mechanisms; detoxification and dynamics uptake variability; individual and pulation, biotransformation or breakdown physiological retention; and localization anatomical mechanisms; and differences in initial Yoshioka, 1981; Maruyama toxicity of dinoflagellates (Shimizu 1991, 1996; Kitts and 1993, 1994; White to STX and ability to continue actively feeding during toxic blooms (Twarog LANDSBERGsodium channels, which function in mammalian nerves as well as in skeletal and cardiac muscle fibers (Kao, 1993), researchers formerly thought that bivalves were not affected by PST to considered typically because not were STX of levels High channels. calcium voltage-gated their neuromuscular functions operate mainly to bivalves (Prakash be lethal or pathogenic by REVIEWS IN FISHERIES SCIENCE researchers focused on the fact that these harmful and mortalities many for responsible been have they that and ways toxins various can and do affect molluscs in effects. molluscs have thus far been associated only with Cembella and Shumway, 1995; Lassus tion with other marine animal mortalities and sometimes with PSP outbreaks (Table outbreaks PSP with sometimes and mortalities animal marine other with tion 2) (see below). The potential effects of different responses. HAB individual species or species-specific and of their series toxins a on on depending vary, may bivalves Some species may initially avoid exposure by behavioral or physiological mecha- nisms or by their individual response to ingested toxins. Because microalgae can produce toxic benthic cysts and/or vegetative planktonic stages, bivalves may be differentially exposed to toxins because of their feeding modes. Some species feed on resuspended particulate matter from sediments, whereas others filter plankton from the water. Bivalve feeding behavior may be one of the may bivalves Some principal ingested. are toxins controlling which to extent the limit initially that factors dinoflagellates toxic of consumption the avoid to behaviorally respond immediately (Shumway and Cucci, 1987; Gainey and of Shumway, presence the 1988a; in Shumway, example, 1990). For A close the shell valves (Shumway, 1990); softshell , the siphon (Shumway and Cucci, 1987); and Pacific , Curtis 1998b; lethal or usually cause only short-term responses, then suboptimal conditions may allow sublethal effects to occur. Differences between bivalve species in the ability species’ each with correlated been have PST accumulate to

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , , ., ., ), et var. et al et et al accu- gigas Pecten from a .

C ., 1993). weeks had weeks 2 g STXeq/100 / 1 µ Mytilus edulis et al spp. — PST are PST — spp. Mytilus trossulus Mytilus minutum . A for 4 for . The digestive gland digestive The . Patinopecten yessoensis (Bricelj ., 1992). Toxins are not , and Placopecten Crassostrea and et al Pyrodinium bahamense . In other bivalves — such as such bivalves — other In . fundyense fundyense tamarense .

A ) bloom conditions (Chebib conditions bloom ) catenella . sea scallops ( scallops sea

, and oysters, and , ), are known to retain high levels of toxins of levels high retain to known are ), 195 Alexandrium excavatum . A Thalassiosira weissflogii ., 1994). Populations of bivalves and species of accumulate very low levels of GTX (Landsberg, (as Placopecten magellanicus Placopecten magellanicus et al ., 1993; Shumway and Cembella, 1993; Cembella and Spisula solidissima Spisula et al tamarense Alexandrium tamarense, A catenatum ., 1994; Bricelj and Cembella, 1995). The location and deposition and location The 1995). Cembella, and Bricelj 1994; ., . . Mercenaria mercenaria Mercenaria A G ., 1996). For example, STX-resistant blue mussels, et al et , in softshell clams and Washington butterclams, and in the scallops and et al , these three tissues contained almost no GTX1 and GTX4 15 days after the after days 15 GTX4 and GTX1 no almost contained tissues three these , edulis . M ., 1990; Cembella Atlantic surfclams ( surfclams Atlantic The toxin profiles of bivalve populations vary depending on the toxigenicity of The toxin profiles of bivalve populations Bivalves store PST for different lengths of time, and the toxic components

., 1990), whereas Washington butterclams, Washington whereas 1990), ., readily accumulated in the adductor muscle of scallops, and because this is the only the is this because and scallops, of muscle adductor the in accumulated readily part of the shellfish usually consumed, they are normally safe for public consump- tion (Shumway and Cembella, 1993). 1996). Bivalve toxin profiles also vary by geographic region, by season, and by the Martin 1990; Liston, and (Beitler tissues different in components toxic of distribution et al are able to depurate toxins from their tissues in less than nine weeks (Shumway al compressum Shumway 1994; weight of toxin components in the various bivalve organs varies between species. For example, in the scallops levels toxicity and gland, digestive the in concentrated are toxins the of majority the in the gills, gonads, and adductor muscles are typically less than 80 g (Shumway and Cembella, 1993). transformed in the gonad, kidney, and adductor muscle of the great , Because toxins are accumulated maximus and then to exposed experimentally was scallop still contained GTX1-4 and NEO after 35 days (Lassus northern quahogs, northern for long periods of time (from Cembella, 1993; months Shumway to more than 5 years) (Shumway and at low levels or are absent (Shumway, 1990; Landsberg, 1996; Bricelj and Shumway, and Bricelj 1996; Landsberg, 1990; (Shumway, absent are or levels low at 1998 and references therein). Depuration times also vary between species can eliminate PST within weeks (Shumway, species. 1990). Pacific oysters, Most and bivalves may also vary in populations with a history of exposure to PSP their toxins appear to have a lower toxin susceptibility to effects sensitivity than do mussels from unexposed populations. Wild blue mussels of PST. Mussels from region chronically contaminated with PST accumulated twice as much toxin on a weight-normalized basis as did cultured mussels from a pristine zone when identical to jected sub- 1993). Such species-specific and intraspecific differences at least partly explain the variations in toxicity profiles reported in molluscs. the dinoflagellate to which each population is exposed. For example, in general, bivalves exposed to mulate high GTX levels, whereas bivalves exposed to that had been experimentally fed a toxic isolate of isolate toxic a fed experimentally been had that were that mussels in those than lower, index condition a but to, similar rates growth fed a nontoxic diet of the diatom rapidly, whereas others are species are able to depurate toxins retained vary. Some slow to detoxify. A mussels the in range found been have concentrations PST of high Extremely PST toxicity levels is found and in different bivalves. Patinopecten yessoensis 1972; Bricelj Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ; ., et al ., 1995, ., 1985; ., 1998). ., spp. STX et al et al et et al ., 1969, 1985; (Sullivan et al Anabaena circinalis Protothaca staminea ., 1982), the crab ., 1983). PST have also have PST 1983). ., ., 1996). However, it is Alexandrium et al et al et et al ., 1983; Daigo g/g dry weight, with the toxin the with weight, dry g/g ., 1979; Shumway 1995); the ., 1988; Nagashima 1988; ., µ et al Mactra chinensis et al et al et (Fusetani and ., 1994, 1995a, 1995b; Tsai 196 et al ., 1981; Raj therefore may suggest a potential for sublethal spp. (Foxall et al Cancer circinalis Peronidia venulosa . A sp., consumed by the crabs (Kotaki crabs the by consumed sp., accumulated high levels of PST when fed Carcinoscorpius rotundicauda Jania ., 1981; Yasumoto et al Although the acute effects of PST exposure on behavioral and physiological Limited information is available on the short-term responses of shellfish to exposure PST with associated effects documented no are there term, short the In Because they naturally ingest a variety of dinoflagellate species and strains, cells/ml). Total PST concentration reached 1580 reached concentration PST Total cells/ml). 6 1996; Negri and Llewellyn, 1998), alga calcareous and in some cases were derived from the horseshoe crab (10 Llewellyn and Endean, 1989; Arakawa been found in the crabs and the Japanese clams 1983; Bricelj and Cembella, 1995; Oshima, 1995; Bricelj forms milder into toxins convert not do that species those of health the how unclear is affected by long-term exposure to toxic components. responses have been demonstrated, other effects generally of unknown. Because PST of on public bivalve health concerns health and are safety the development protocols, of it has been critical distribution that in different species, we particularly in understand edible tissues. In the general, although might dynamics there that thought usually not is it of shellfish, among distributed widely are PST toxin be negative health effects associated general, shellfish are thought with to be immune from such effects. Yet long-term some consider- toxin deposition, ation and, of possible in effects HABs, is between association appropriate, possible a especially example, in For unknown. cases still is where diseases the certain etiology of toxin deposition, and the incidence of recently (Landsberg, neoplasia 1996). in shellfish was hypothesized cyanobacterial exposure. Under experimental conditions, the Alathyria condola freshwater profile comprising mostly C toxins and high significant very levels to exposed of Mussels GTX2 viscera. and the in GTX3. contained The were (96%) toxins of majority Feeding feeding. cease then and hours few a for feed filter to tended concentrations resumed from 1 to 3 days later. The mussels’ erratic feeding concentrations behavior were when high cell was not observed when animals were exposed concentrations to of cells. low The reduction in feeding rate associated with exposure to high concentrations of deleterious effects in mussel populations (Negri and Jones, 1995). accumu- in other invertebrates. Crustaceans, echinoderms, and (rarely) polychaetes (Daigo effects ill any without apparently PST late PST have been found most commonly in xanthid crabs (Noguchi Koyama toxins are typically stored in the tissues of these species, whereas other potentially carcinogenic substances such as the carbamate-derivative gonyautoxins are verted to con- less toxic compounds. The ability to convert corresponding carbamate nontoxic decarbamoyl toxins derivatives to has been their demonstrated in a few bivalves, such as Atlantic surfclams; Pacific littleneck clams, LANDSBERGbivalves are exposed to a variety of toxic components. Knowledge of which toxins are deposited in which tissues may be critical for determining the effects of each toxin on shellfish health. For example, naturally exposed in Atlantic New England to surfclams PST associated and with sea scallops are REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , , , Clupea ., 1971). ., Portunus Homarus et al et sanguineus sp. (Jonas- ., 1998). Alexandrium . Daira perlata Metopograspus gSTX eq/100 g H , µ et al gSTX eq/100 g (1MU Astropecten scoparius Eudistylia nudus µ . spp. (Sang and Ming, , H ) were confirmed to be , ., 1982; Jonas-Davies and et al in Australia, even though the Panulirus ., 1988), and presumably neural ., 1993); penaeid shrimp (Sang and (Sang shrimp penaeid 1993); ., et al ., 1998). Some predatory invertebrates et al et g PST/100 g (Negri and Llewellyn, 1998). Llewellyn, and (Negri g PST/100 g gSTX eq/100 g (Desbiens and Cembella, (Noguchi µ µ Asterias amurensis ., 1993); the blue manna crab, et al 197 contained nearly 16,500 Mouse Units (MU) (Sato et al Atergatis floridus Hemigraspus oregonensis Hemigraspus spp. (Jonas-Davies and Liston, 1985); and other have a very low affinity for toxins (Llewellyn, 1997). have a very low affinity for toxins (Llewellyn, ., 1957; Negri and Llewellyn, 1998). The nerves of (Sato ., 1997; Lin et al Balanus Zosimus aeneus et al Pisaster ochraceus floridus , apparently did not cause any significant health problems. . ., 1983), which is equivalent to 300,000 A and algal remnants. In this case, the pteropods acted as vectors as acted pteropods the case, this In remnants. algal and , were observed dead and dying. A bloom of Procambrus clarkii Procambrus gSTXeq/100g in the of the American lobster, ; the shore crabs crabs shore the ; µ , and et al ., 1985). Maximum toxin levels of more than 16,000 (Desbiens and Cembella, 1995) and are 1000-fold less sensitive to PST than are the nerves of was coincident with the mortality, and shellfish (softshell clams; blue et al (Sang and Ming, 1984; Negri and Llewellyn, 1998); the lobsters gSTX; Schantz (Jonas-Davies and Liston, 1985; Negri and Llewellyn, 1998), Llewellyn, and Negri 1985; Liston, and (Jonas-Davies µ Pachygrapsus crassipes As they did historically with shellfish, researchers believed that cold-blooded It has been suggested that lobsters exposed to high PST concentrations do not floridus polyacanthus . . sodium channels in a hemolymph protein, saxiphilin, that can Several species of xanthid crabs produce bind with STX and thus confer some resistance to possible toxic effects (Llewellyn, exception- tolerate to appear species some why explain may mechanism This 1997). ally high levels of toxin (Llewellyn, xanthid crabs, the rate of 1997). dissipation within the crab was fairly high and suggested When saxitoxins were injected into that high concentrations of toxin are not accumulated (Arakawa contaminated with PST. Stomach contents from dead fish contained the pteropods retroversa Limacina a xanthid not known to carry PST (Daigo (Prakash PST by affected negatively not were finfish as such vertebrates In recent years, however, several studies have demonstrated the lethality of PST to finfish, and researchers have also begun investigating the finfish tissues. distribution In July of 1976 in PST the Bay in of Fundy, Canada, Atlantic herring, harengus harengus tamarense mussels; and northern horsemussels, Davies and Liston, 1985) and the Davies and Liston, A 80 than less contained samples of majority In Japan, an individual per g (Koyama = 0.18 A Liston, 1985; Asakawa pelagicus americanus crayfish the 1984); Ming, 1984); barnacles crustacean species (in Shumway, 1995, and invertebrates that Negri accumulate and PST Llewellyn, include 1998). the Other tubeworms accumulate toxins through the consumption of toxic bivalves (Shumway, 1995). survive (Yentsch and Balch, 1975), but it including lobsters, are able to tolerate extremely high levels of PST. Concentrations appears that many macrocrustaceans, of up to 1512 Homarus americanus can which others, among mussels, blue on preying by PST accumulate can Lobsters have maximum toxicities of up to 23,000 1995). Xanthid crabs can harbor toxins in their tissues at concentrations that would be fatal to other animals (Llewellyn, 1997). When some xanthid crabs are handled, mechanism defense a as used are toxins these that suggests which PST, release they (Noguchi were found in the xanthid crab frontalis and Pugettia producta Pugettia Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., (as et al Mallotus ) by first- Lythrypnus tamarense . cultures, gobies A Engraulis japonica Engraulis excavata . A catenella ., 1991a, 1991b). The level . A , and within several minutes spp.). et al -toxic crab larvae, mortalities were mortalities larvae, crab -toxic , resulted in high mortalities and, strains, mode of exposure to toxin, to exposure of mode strains, ., 1989). Fish kills associated with -fed zooplankton, some (22 to 38%) (reported as ., 1991a, 1991b). With increased strain larvae exposed to concentrations of

tamarense , . et al A et al catenella 198 Alexandrium . A ., 1991b). When bluebanded gobies, bluebanded When 1991b). ., Alexandrium Pagrus major ., 1989; Robineau tamarense . Alexandrium et al et A cells/ml died sooner than those exposed to lower et al 2 10 × ., 1989; Robineau et al had died, and moribund larvae lay paralysed on the tank bottom. in the estuary and Gulf of St. Lawrence showed that capelin, that showed Lawrence St. of Gulf and estuary the in above 1

species, either directly or via toxic zooplankton vectors, results in ) spp. have been reported in numerous geographical locations (Table ., 1986; Gosselin ., 1989). After 4 days, 95% of the red sea bream larvae exposed directly Pseudopleuronectes americanus et al et al , could not detect the presence of a toxic bloom and confirmed laboratory , were experimentally exposed to exposed experimentally were , ., 1989; White Preliminary results on the codistribution of fish larvae and Experimental ingestion of Alexandrium excavatum tamarense . . observations that these fish cannot discriminate between toxic and nontoxic strains. nontoxic and toxic between discriminate cannot fish these that observations In addition, local survival of fish larvae appeared to be determined by the density villosus toxicity, mortality from vectoral intoxication tended to increase, but not as rapidly as mortality from direct intoxication. These results are consistent with the observa- tions that zooplankton eliminate the toxins and that larvae first-feeding nature, in the Thus, toxicity. final the in variability saturation reduced in results of the tissues that readily feed on dinoflagellates will (Robineau be postlarvae carnivorous at a higher risk of intoxication than concentrations or than larvae on control diets did (Mills and Klein-MacPhee, 1985). of mortality caused (Gosselin by the directly toxins toxins was the invariably to less exposed when were the fish fish the were when exposed than vectors via toxins to et al dalli A Within 1 to 2 h after feeding on their on swam and equilibrium their lost species both of larvae weeks) 6 to (4 older larvae died within a few hours (White sides, upside down, or in circles. Most and the susceptibility of the fish to Huntley the toxins (Mills and Klein-MacPhee, 1985; observed within 24 to 60 h. When exposed directly to all treated fish displayed similar symptoms. Fish swam in an irregular, jerky manner; jerky irregular, an in swam Fish symptoms. similar displayed fish treated all tight in swimming some paralyzed, became sides; their on swam and equilibrium lost after treatment 79% of them breathing slowly. Within 1 to 2 days circles; and began which field, the in had they as widely, gaping mouths their with died fish The died. is suggestive of asphyxiation (White, 1977). White (1981b) suggested that fish may be as sensitive as homeotherms are to PST and has provided further corroborative evidence for the role of zooplankton as vectors of PST up the food chain (White 1979, 1980, 1981a, 1981b, 1984; White feeding larvae of red sea anchovy, bream, Japanese of larvae first-feeding of debilitation eventually, (White to 1989). Several studies have demonstrated Alexandrium further that feeding and age the on depends exposing toxins to Vulnerability growth. reduced or fish mortality larvae to of toxigenicity fish, the of behavior exhibited erratic swimming, loss of balance and buoyancy, rapid darting prior to death, and death within 13 flounder, to 14 h (Yazdandoust, 1985). First-feeding winter LANDSBERGfor transmitting the PST to the fish. In exposed by gavage laboratory to crude experiments, extracts of 28 herring were REVIEWS IN FISHERIES SCIENCE Alexandrium 2). In some cases, these kills may have been caused by toxins other than PST (see section on uncharacterized toxins and A

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . , 1 ., 10 × et al et (Sato ., 1984) ., fundyense fangi et al et , mortalities , . . tamarense ., 1993). . A T A were reported (GTX); , et al cells directly but (GTX, NEO); and , liver extracts were Takifugu poecilonotus (STX, GTX); , ., 1997) and 1997) ., ., 1993, 1997). It is unclear is It 1997). 1993, ., g STXeq/100 g wet weight. wet g STXeq/100 g µ tamarense . in Bangladesh (Zaman Bangladesh in et al et ., 1984; Nakamura 1984; ., et al et Scomber japonicus Scomber A cells that were found both free et al et ., 1998). Katsuwonus pelamis Scomber scombrus Alexandrium tamarense Lamna ditropis et al (Kungsuwan Gadus macrocephalus 199 tamarense . A Tetraodon cutcutia Tetraodon ., 2000). leiurus . in Japan (Kodama Japan in T and (GTX, NEO, STX) (Sato STX) NEO, (GTX, et al ., 1996). The PST profile of fish differed from that of the ., 1996). The PST profile of fish differed and strain (Montoya occurred in southwestern Bay of Fundy, New Brunswick, et al pardalis . (NEO, STX); skipjack, (GTX, STX); cod, T cells/ml when mortalities were reported. Dying fish were gasping , and no larvae were found when cell densities exceeded 1.3 2 tamarense , and , . 10 A × Chelonodon patoca Chelonodon Ypsiscarus ovifrons Ypsiscarus g STXeq/100 g were detected in the stomach, but no PST was detected in detected in the stomach, but no PST g STXeq/100 g were Tetraodon suvatii Tetraodon µ ., 1997) in Thailand, and in the marine puffer fish, ., 1990). Recently, PST were implicated in chub mackerel, chub in implicated were PST Recently, Although not usually targeted, PST have been incidentally found in numerous As they are in bivalves, the toxic profiles of PST that accumulate in fish are likely are fish in accumulate that PST of profiles toxic the bivalves, in are they As Even though PST can be toxic to fish, it appears that fish are not usually vectors usually not are fish that appears it fish, to toxic be can PST though Even Alexandrium vermicularis . of survival the compromise can therefore dinoflagellates toxic of presence The cells/ml. of larval cohorts and recruitment to local populations (Robineau to be 3.45 at the surface, and swimming on their sides or upside down (loss of equilibrium). Mackerel stomach contents included Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS in Argentina. In August 1993, cell counts of in Argentina. In August or within mucus strands from ingested presence salps. of PST in stomach A contents, liver, intestine, and mouse gills. The highest bioassay values confirmed the of 2800 the muscle of the fish. The most abundant toxins in the stomach were GTX1 and GTX4. The toxin profile of the liver included GTX2 and GTX3 and minor amounts of STX and NEO. Mackerel did not appear to ingest Although toxins were principally exposure transmitted other to the suggest mackerel also through the may diet, gills the the in toxin of amount significant a of presence mechanisms (Montoya Argentinian was attributed to the variety of (Haya chain food the up PST of toxic transfer the demonstrates PST accumulate mackerel prey consumed by the fish. The fact et al that concentrations low with identified been have species marine Several fish. of species of PST in the liver or intestine: shark, rather were exposed by preying on toxic salps that had fed on to be partially determined by process species-specific or to be differences dependent upon the in variety of toxic the prey species — bioconversion and toxins of they the contain — consumed by the fish. During July 1988, a small bloom of Alexandrium fundyense Canada. The highest concentration in a Concentrations of surface-water PST in Atlantic mackerel, sample was 7.5 cells/ml. Oncorhynchus keta Navodon modestus parrotfish, By far the dominant component in mackerel liver was STX except where in a few NEO fish, was also detectable. dominant. The difference between the toxic profiles GTX2 of the fish and and GTX3, and rarely B2, were also if or to what extent these toxins may contribute to chronic health effects. As part of a study for TTX (see below), STX and GTX were also found in fish the puffer freshwater measured by mouse bioassay and ranged from 40 to 209 to 40 from ranged and bioassay mouse by measured T 1997), in 1997), et al and in the Philippines (Sato for PST transfer if humans only eat fish muscle. The accumulation of toxins is usually is toxins of accumulation The muscle. fish eat only humans if transfer PST for

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . sp. Selaroides g/kg body g per 100 g µ Pyrodinium compressum µ Alexandrium ., 1990) and in Rastrelliger have been con- var. bloom (Table 2). (Table bloom et al spp. and spp. (Maclean, 1979). One bloom in southwest Mexico Sardinella compressum ., 1989). During these var. compressum et al var. Alexandrium catenella Alexandrium compressum Pyrodinium bahamense Pyrodinium 200 var. sp. from Brunei (Oshima , toxic shellfish, and fish that were reported from reported were that fish and shellfish, toxic , ., 1989). These incidents likely occurred because it is it because occurred likely incidents These 1989). ., red tides, fish kills, and PSP outbreaks (Maclean, 1973). (Maclean, outbreaks PSP and kills, fish tides, red et al et Sardinella bahamense . Pyrodinium bahamense P Pyrodinium g/kg body weight) or i.p. (intraperitoneal) (4 to 12 g/kg body weight) Pyrodinium bahamense µ blooms, but there have been a few reports of turtles in both Asia blooms, but there have been a few Pyrodinium . In a second incident in November 1983, 45 people became ill after = 400 to 750 Because PST does not appear to accumulate in fish muscle, humans who of areas in PST by affected be to reported typically not are Turtles are not If PST are ingested by fish or other secondary producers and the toxins 50 Gymnodinium From the mid 1950s until with concurrent the early 1970s, several dead During a winter (1995) turtles were reported weight) doses of PST showed reduced locomotor similar activity; short, irregular, hyperactive symptoms: periods; and death within loss of equilibrium; 1 h. Heavy accumulation of PST gasping; was confined to the gut (340 to 840 tissue); PST occurred in the muscle tissues at a level an order of magnitude lower than in the gut (White, 1984). consume only the muscle are unlikely to become intoxicated, whereas those who eat the viscera are likely to become sick. In 1976 in Brunei, 14 nonfatal PSP cases were associated with the consumption of the during a bloom planktivorous of fish PSP incident in 1983 in Indonesia involved 191 cases and four human fatalities due fish clupeoid planktivorous the of consumption the to leptolepis firmed in gut contents of involving incidents PSP blooms, mortalities of invertebrates, fish, turtles, and dolphins were also reported (Table 2) (see below). or — more than 145 turtles (species not specified) stranded on the beaches — along with fish and hundreds of lobsters (Orellana-Cepeda, the 1998). with Whether associated the conditions fish hypoxic and or anoxic to due were strandings invertebrate bloom is unknown. In neither the Asian nor Mexican incidents was made for examination the presence of PST or for possible vectors of toxin exposure. lethal to those organisms (unlike several of the previously cited examples in which be will ingested toxin the that possibility the is there then occurred), mortalities fish PST involving events mortality Mass chain. food the up passed and bioaccumulated piscivorous cases, most In 2). (Table occasionally documented been have birds and least at 1942, May In PST. by contaminated fish consuming after affected were birds 2000 dead sea birds were observed along the coastal beaches of Washington State, an with coincident was mortality The USA. At the same time, six human cases of fatal PSP had been reported, three from the LANDSBERGconfined to the fish’s gut, concentra- 1980, 1984) or negligible the muscle (White, of toxin appear in amounts and either certain species tions of perish toxins accumulate in the muscle. Several before fish species challenged with oral detectable (LD REVIEWS IN FISHERIES SCIENCE lips, the of sensations tingling and dizziness, numbness, suffered and fish consuming the with associated was dinoflagellate toxic known no Although throat. and tongue, with PST 1985). White, and (Maclean suspected highly was PSP 1984), (Adnan, event toxin profiles similar to of blooms by affected were that Mexico and the Philippines (Gonzalez Philippines the customary in southeast Asia to eat small fish whole, including any potentially toxic viscera (Maclean and White, 1985; Gonzalez

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 et ., 1968). et al ., 1968). The et al sp., were reported were sp., ., 1978). ; and Baltic macomas, Baltic and ; et al , had very low STX levels Ammodytes Cardium edule Cardium cells. 201 Crassostrea virginica catenella . ; edible cockles, edible ; A bloom in the same area (Adams , were found in a moribund state along the shore. Although blue Venus striatula Venus The first red tide bloom that led to a major PSP outbreak in the USA occurred In May 1968, dying sea birds and dead sand eels, sand dead and birds sea dying 1968, May In Alexandrium tamarense ., 1968.] Although no sick individuals were reported, a total of 636 birds (species From May to June 1975, a estimated similar 500 shags were incident affected (Table occurred 2) (Armstrong in the same area, and an in September 1972 from southern Maine to Cape Ann, Massachusetts. Blue mussels and softshell clams were the bivalves most did quahogs susceptible Northern reported. were clams to softshell Paralyzed toxic. most PST the were accumulation and not accumulate toxins, even in areas where blue mussels and softshell clams had high toxin levels. Eastern oysters, (Twarog and Yamaguchi, 1975). In a few isolated areas, softshell clams and blue mussels remained toxic until April 1973 (Hartwell, 1975), but this may have been attributable to a reocurrence of dinoflagellate cells (S. Shumway, personal commu- neopla- disseminated of reports several with coincided also outbreak This nication). sia in softshell clams, but whether there is a connection with PST remains uncon- firmed (Landsberg, 1996). Birds were affected during this red tide, with mortalities of at least 620 reported waterfowl, gulls, and shorebirds, representing 13 species, presence the showed contents gut of samples Representative ducks. black 1600 and Olympic Peninsula and three from Vancouver Island, British Washing- the along dying also Columbia. were viscera razor consumed Cats had that chickens and ton beaches. At least eight crustaceans and species clams in their stomachs of and inflamed intestines. The birds peak occur- (Table 2) rence of dead were birds was about found 2 weeks after dead the crest of with the red 2 weeks tide, after the and disappearance of the about bloom, the numbers of dead birds seen on the beaches began to decline. Based on the available information, McKernan and Scheffer (1942) suggested that the birds were killed by feeding on toxic crustaceans fish that and had ingested Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS around the possible Farne Islands, a northeast England. When continued, several humans were fish admitted dead of reports the and PSP of symptoms with hospital a to connection with the presence of a red tide bloom was investigated. Large numbers Venus, striped of Macoma balthica mussels were found to be highly toxic, there were no mortalities observed in this were recorded about a week after the peak of species. The first deaths of sand eels an 100 per (MU) Units Mouse 20,800 to 10,062 from ranged mussels in toxin of amount intraperito- injected when that toxin of amount the as defined was MU (One tissue. g neally, causes typical neurotoxic signs and kills a 20 g mouse in 15 min [Ingham al listed in Table 2) were found dead along 65 miles of the northeast coast. Many of they died. The majority of the dead birds the birds had been feeding shortly before breeding the of 82% days, few a Within fulmars. and terns, cormorants, shags, were found on the Farne Islands exhibited a loss shag population had died. Dying shags of equilibrium, lack of motor coordination, paralysis, restriction of pupils, excess intestinal hemorrhage, failure of the circu- vomiting, abnormal green-brown faeces, latory system, and congestion of organs, including the lungs. All these symptoms in humans and domestic chicks affected were remarkably similar to those observed by PSP. Seventy-eight cases of PSP occurred almost simultaneously during this May 1968 period in (Coulson humans caused by eating toxic mussels

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . ., ). et al et ., 1999). Scomber Megaptera et al ., 1975). ., 1998; Reyero 1998; ., et al g STX eq./100 g), of µ et al et Dicentrarchus punctatus Dicentrarchus Gymnodinium catenatum Gymnodinium and ., 1998). A morbillivirus (MSMV-WA) et al 202 , and spotted seabass, spotted and , g STX eq/100 g tissues in six seal samples and µ Alexandrium minutum Alexandrium , died along the coast of the western Sahara, Africa. The seals Diplodus sargus Diplodus , died in Cape Cod Bay after eating Atlantic mackerel, , containing STX. The absence of STX in New England waters and shellfish and waters England New in STX of absence The STX. containing , . Many other birds apparently perished after feeding on toxic shellfish, but ., 1989). ., 1998). The mortality affected mainly adults, which is not consistent with the ., 1998), so other epizootiological factors in this mortality may be relevant. The Recently, from May to July 1997, at least 117 Mediterranean monk seals, A classic case of transfer of PST up the food chain and subsequent mortality at Other examples have been documented of marine mammals having been Mouse bioassays yielded 30 to 40 (blacktail, fish two in were in a good nutritional state and had no obvious gross lesions, but their lungs were congested and filled with fluid. Terminally ill animals were lethargic, lacked motor coodination, and were paralyzed in the clinical water; of onset animals the between floated period The horizontally movement. effective of incapable were and signs and the death of the seals was short. Water samples indicated producers the PST presence known the of Monachus monachus directus these were not recovered (Bicknell and Walsh, 1975; Sasner a higher involved humpback whales in New England, USA. During a 5-week period beginning in late November 1987, 14 humpback whales, novaeangliae scombrus toxins farther north in suggested that the mackerel accumulated during the episode consuming after STX to exposed were whales the that and Lawrence, St. of Gulf the toxic fish that had migrated south. Animals were in good condition, there were no significant lesions, and at least six of the whales had partially digested mackerel in their stomachs, suggesting that they had been feeding just prior to death. STX was present in the viscera, especially the liver mackerel (mean caught where of the whales had 153 been feeding. Extracts from kidney, liver, and bioassay, standard a in mice to lethal were whales the of contents stomach and the mice showed the classic signs of STX toxicity. This was the first case demon- strating the transfer of PST through a commercially important fish and the first one documenting that PST were involved in a marine mammal mortality event (Geraci et al PST. to exposure after seen usually those with consistent were mice the in Symptoms (liver, samples seal all in found were GTX1) and NEO, (dcSTX, PST of levels Variable (Hernández fish all in brain), and muscle, skeletal kidney, 1999), and a few mussel samples. Results were liquid chromatography confirmed (HPLC) and by mass spectrometry high-performance (MS) (Reyero LANDSBERG jacknife, Atlantic the and mussels blue as such bivalves filter-feeding small, of REVIEWS IN FISHERIES SCIENCE The evidence suggested that PST were a likely cause for these mortalities (Hernández mortalities these for cause likely a were PST that suggested evidence The et al juveniles where mammals, aquatic in outbreaks morbillivirus recent of demographics are more severely affected (Hernández (Osterhaus found was morbillivirus dolphin related closely the of that from different et al potential immunosuppressive role of PST and associated increased susceptibility of marine mammals to virus infections should also be considered, along with potential interactive other causal factors. affected or suspected of having been affected by PST, but the evidence has often conducted. been not have tests appropriate the or inconclusive or unavailable been alone tissues in toxins microalgal of finding the contaminants, of kinds other with As

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ; g PST/ µ Anabaena ; and the livers the and ; , mortalities in the , mortalities Anabaena circinalis ., 1995). Two extensive et al ., 1990). Enhydra lutris Enhydra Kuhlia sandvicensis Kuhlia et al Acanthurus triostegus sandvicensis channel inhibitor could implicate PST, implicate could inhibitor channel + . Fish such as capelin (imported from blooms were recognized to be due to 203 blooms showed that approximately half the ; the aholehole, the ; circinalis . A circinalis g STX/100 g tissue is considered the regulatory limit for . µ A chaptalli ., 1998). Interestingly, although there is a worldwide distribution channel inhibition was shown after injections of extracts of et al ., 1990). Necropsy examination revealed no significant findings except findings significant no revealed examination Necropsy 1990). ., + Cheilinus rhodochrous Cheilinus extract injected i.p. into mice caused death within 4 to 7 min. Symptoms et al et Although dinoflagellates have been implicated in most of the documented cases documented the of most in implicated been have dinoflagellates Although One interesting, but unconfirmed, example from Hawaii indicated that a potent a that indicated Hawaii from example unconfirmed, but interesting, One channel inhibitor, characteristic of TTX () (see below) or STX, may channel inhibitor, characteristic of TTX + which would be unusual in this area (Hokama Kodiak Archipelago, Alaska, USA, during the summer of 1987. At least 60 animals such in were animals dead cases, many in but October, and June between died had suitable samples obtain to impossible was it that decomposition of state extreme an for analysis. Necropsies of four otters showed no obvious pathology, but they had were There drowning. to prior water of aspiration suggests which lungs, the in foam dive to only able was that animal moribund a of one and otters sick of reports three feebly. Coincident with the discovery of the dead otters, two cases of human PSP on Kodiak Island resulting from documented. One mussel sample collected the from the site contained > 5800 consumption of toxic blue mussels were from died dolphins the that suggested was It respectively. livers, dolphin and viscera toxins present in the gut contents of mullet, wrasse, and aholehole, but Na that a these was toxin the That CTX. not were toxins in which PST have affected animals, other organisms are now known to produce similar effects. Recently, and for the livestock first previous Although Australia. time, in mortalities PST sheep in from implicated were bloom an mortalities associated with have been involved in the death of two Atlantic dolphins living in an artificial lagoon artificial an in living dolphins Atlantic two of death the in involved been have (Hokama does not necessarily imply a direct cause and effect. A strongly circumstantial case otter, of PST in sea for the involvement was made 100 g mussel meat (80 human consumption). Consequently, De Gange and Vacca (1989) suggested that PSP could have been the cause of the sea otter mortalities. Na changes in the intestinal tract. Testing of fish for hypersensitivity-type inflammatory present in the same lagoon by stick enzyme immunoassay (S-EIA) confirmed the presence of (CTX) (see ), with viscera high of levels the present in mullet the usually were that California) from (imported mackerel and herring, Newfoundland), fed to the dolphins were negative for CTX by S-EIA as were extracts from dolphin livers and stomach contents. In from resulted a legs hind the in mouse paralysis and breathing, irregular bioassay, slow cyanosis, walk, symptoms, including wobbly injections of the viscera of mullet; the manini, wrasse the and gastric contents of two dolphins. In the guinea pig atrium moderate assay, Na strong and neurotoxic exposure, the neurotoxins involved were not identified. Affected sheep showed symptoms of trembling, recumbency, and crawling; necropsies and histo- pathology showed no significant lesions. Except at circinalis the lowest dose, included staggering at low doses, followed by gasping, leaping, respiratory and failure; salivation death was not observed from (Negri surveys of Australian concentrations detectable contained two-thirds and mice to neurotoxic were samples of PST (Negri Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ; ; ., ; the , the et al , and Atergatis Salvelinus Planocera Gadus macro- rivularis . (Kodama Tachypleus gigas T , Oncorhynchus keta Oncorhynchus and , or at least by bacteria produce only anatoxin- only produce ., 1993; Shumway, 1995; Shumway, 1993; ., ; the cod Katuwonus pelamis Procambarus clarkii et al et ; the xanthid crab , and char, ; and the flatworm ; the salmon the ; ., 1993, 1996). Although TTX is circinalis circinalis . torosa A ., 1988), which appear to be immune et al ., 1986; Tamplin, 1990). Recently, TTX Cololabis saira ; the skipjack ., 1988). Although TTX acts in a manner a in acts TTX Although 1988). ., Patinopecten yessoensis scoparius ; the crayfish et al . et al 204 et al et A Alexandrium tamarense Lamna ditropis Lamna and ; the saury Oncorhynchus mykiss ., 1982, 1983; Mosher and Fuhrmann, 1984; Miyazawa 1984; Fuhrmann, and Mosher 1983; 1982, ., (Noguchi Carcinoscorpius rotundicauda the salamanders

Hapalochlaena maculosa ., 1987; Tamplin, 1990; Sato 1990; Tamplin, 1987; ., ; et al et Ypsiscarus ovifrons ; the shark the ; et al et Lophozozymus pictor Aeromonas (Noguchi (Noguchi Conger myriaster , it appears that there is a geographical segregation of and ; the Gobius criniger Gobius ., 1995; Lin and Hwang, 2001). ; the parrotfish can detect extremely low levels of TTX via specialized gustatory receptors. Navodon modestus Astropecten polyacanthus

; the horseshoe crabs Vibrio , circinalis et al . TTX has been found in numerous other species of aquatic organisms, including organisms, aquatic of species other numerous in found been has TTX Chemically, TTX is very different from STX, but it produces similar symptoms ., 1986; Kungsuwan 1986; ., granulosa A . a (see later in text), whereas Australian isolates produce exclusively PST (Beltran and (Beltran PST exclusively produce isolates Australian whereas text), in later (see a Neilan, 2000). B. TETRODOTOXIN Another type of fatal human poisoning with characteristics similar to those of PSP the consumption of the fish to tetrodotoxin (TTX) through is attributed to exposure found been also has TTX 1984). Fuhrmann, and (Mosher fish) (puffer fugu’ delicacy in molluscs such as the Japanese scallop, LANDSBERGof of isolates European and American production. REVIEWS IN FISHERIES SCIENCE 1993), and at least eight species of gastropods (see Shumway, 1995; Lin and Hwang, and Lin 1995; Shumway, (see gastropods of species eight least at and 1993), 2001); these organisms represent another Although poisoning events associated with eating fugu usually occur potential in Asia, cases source for also have 1993) (Kao, Europe in and 1984), human Fuhrmann, and (Mosher USA Florida, in toxicity. been reported. TTX acquired through consumption of puffer fish, along with other with associated state “zombie” human the for responsible largely also is ingredients, the vodoun religion (Davis, 1985). goby the the conger eel cephalus filefish present in a wide range of aquatic organisms, it is not known to what extent the toxin originates from dinoflagellates. High concentrations of TTX, which are fatally toxic to other organisms, are known to (Mosher and Fuhrmann, 1984; Yamamori occur in puffer fish and xanthid to crabs its effects. Rainbow trout, T associated with this dinoflagellate (Kodama alpinus Some other fish species have been shown to reject toxic puffer livers and artificial (Yamamori TTX containing pellets food similar to PST, it is currently unclear what effects TTX has on aquatic organisms. was also shown to be produced by in mammals because they both act on channel. Like PST, TTX site exerts its toxic 1 action by specifically blocking of the voltage- the voltage-dependent sodium sensitive sodium channels in nerve biogenetic origin of TTX has been linked to bacteria such as the widely distributed and muscle membranes (Kao, genera 1993). The floridus the Taiwanese crab starfish multitentaculata al et Tsai

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., ., et al et et al et et al ., 1995; ., antiqua . Fibrocapsa C et al et ), and ), ., 2000; Hu 2000; ., strains originally that correspond to

et al et ., 1990). PbTx-1 is the ., 2002) carterae sp. and the raphidophytes the and sp. . (HaTx) japonica et al ., 1996b). Four toxins have toxins Four 1996b). ., (Table 1). -like . H ., 1986; Baden and Tomas, F et al (= et al et ., 1986). Dominant brevetoxin (Cembella et al ., 1984; Catterall, 1985; Poli 1985; Catterall, 1984; ., Karenia akashiwo et al et al et . verruculosa H (Bourdelais cf. 205 ) Hansen and Moestrup (Daugjberg Heterosigma akashiwo Heterosigma cultures (Shimizu (Baden, 1989; Schulman , (CmTx) and verruculosa Chattonella Alexandrium ostenfeldii Alexandrium brevis brevis . cf. . . C marina K K . marina . C C , Gymnodinium breve ., 1987). The effects and toxicokinetics of brevetoxins in mamma- in brevetoxins of toxicokinetics and effects The 1987). ., blooms (Table 2) that produce brevetoxins are far more devastat- blooms (Table 2) that produce brevetoxins (= et al et ., 1995, 1996a, 1996c). Further chemical characterization is required to Karenia (Table 1). Brevetoxins can cause serious health effects and significant et al ., 1990). PbTx-3 is the primary toxin responsible for respiratory irritation (Baden irritation respiratory for responsible toxin primary the is PbTx-3 1990). ., Five neurotoxic components (FjTx-I, FjTx-II, FjTx-IIIa, FjTx-IIIb, and FjTx-IV) By binding to specific sites on the voltage-sensitive channels, brevetoxins alter Brevetoxins are complex, polycyclic ethers, which are differentiated into two PbTx-2, oxidized PbTx-2, PbTx-3, and PbTx-9, (Ahmed PbTx-3 and and PbTx-2, three oxidized PbTx-2, toxins to correspond from that (CaTx) Khan definitively show that FjTx, CmTx, CaTx, and HaTx are the same as brevetoxins, as recently confirmed for most potent brevetoxin described (Shimizu components are PbTx-2 (4.93 to 12.6 pg/cell), PbTx-3 (0-2.31 pg/cell), and PBTx-1 (0.07-1.72 pg/cell) in Pierce 1986; (Pierce, aerosols bubble-generated in PbTx-3 and PbTx-2 are and 1989) al et and Mende, 1982). corresponding to brevetoxins PbTx-1, PbTx-2, PbTx-9, PbTx-3, and oxidized PbTx-2, respectively, were identified by HPLC from cultures of (Khan Sea North the of part Dutch the from isolated been isolated from membrane properties of excitable cells. Both brevetoxins and ciguatoxins bind to sodium channel site 5 to shift activation to more negative potentials, (Huang depolarisation membrane subsequent resulting in C. SPIROLIDES Spirolides are a group of macrocyclic amines that have recently been confirmed in shellfish from aquaculture sites in Nova Scotia, Canada. animal Although health the significance human of or these toxins is unknown, when spirolides caused injected rapid death due to into neurointoxication. The bioorigin of mice, spirolides be to determined been has D. BREVETOXINS Brevetoxins are potent neurotoxins and hemolysins produced by the dinoflagellate 2000) and the raphidophyte dinoflagellate the in found been have compounds japonica animal mortalities (Table 2). In terms of their frequency and the range of species affected, ing than blooms of raphidophytes (Table exception of 5). one case [Table 5], and an unconfirmed case [Section T.3]) Thus have been far raphidophytes (with the reported to be ichthyotoxic only (Table 5). type PbTx-2 rings; 10 type (PbTx-1 structure backbone their to according types main been have PbTx-10) through PbTx-5 PbTx-3, to (PbTx-1 brevetoxins Nine rings). 11 isolated thus far from Lombet 1986; Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 2001). antiqua Chattonella

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al ., 1995; ., 1991; et al et al ., 1996; Chang et al ., 1973; Baden, 1983, 1988; ., 1993; Chang ., 1991; Tester et al et al et al ., 1992). cells by breaking waves, surf, or et al aerosols can also cause conjunctival sp. (Haywood brevis ., 1990). In humans, NSP is characterized . 206 K brevis et al . K Karenia ., 1965; Steidinger, 1993). Neurotoxic Shellfish red tide that originated off the west coast of Florida of coast west the off originated that tide red ., 1989; Cattet and Geraci, 1993) and the chemistry, et al brevis skin penetration of PbTx-3 (< 3% of dose) in guinea pigs skin penetration of PbTx-3 (< 3% of . et al K ., 1975). Occasionally there are reports of skin irritation and is one of the oldest reported HABs. Fish kills have been reported been have kills Fish HABs. reported oldest the of one is in vitro et al is principally distributed throughout the Gulf of Mexico, with (Baden, 1989) and for brevetoxins in mice ranges from 0.05 mg/kg body weight for i.v. ., 1991). Exposure to ., 1996a). Toxic shellfish were detected in the Bay of Plenty, northeast 50 brevis

et al et al NSP cases have been associated principally with brevetoxins produced by Karenia brevis Karenia Steidinger, 1993). From December 1992 to January 1993, a large-scale NSP incident occurred for the first time in New Zealand (Bates 2 h after the initial exposure (Kemppainen irritation, rhinorrhea, and nonproductive cough; asthmatics may also wheezing (Music experience was there Experimentally, tides. red by affected waters in swim people when itching some evidence of Karenia were There Stream. Gulf the by waters coastal Carolina North to transported was and 48 documented cases of people contracting NSP by eating toxic shellfish; 35 and cases (Fowler officials state by implemented be could bans harvesting before occurred Tester, 1989; Tester and Fowler, 1990; Morris occasional red tides occurring along the mid and south Atlantic coast of the USA (Steidinger, 1993). Although shellfish poisonings identified from not was cause the eating 1884), (Walker, 1880s the since bivalves known been have USA, in Florida, until the 1960s (McFarren pharmacology, pharmacology, and toxicology of brevetoxins in experimental systems (see Baden, 1983, 1989) have been studied. Mice administered brevetoxins i.p. show an irrita- bility immediately after injection, paralysis. followed respiratory from death and defecation, urination, lachrymation, salivation, by hind-quarter paralysis, dyspnea, Whole LD (intravenously) administration to 0.5 mg/kg for oral and i.p. administration (Baden and Mende, 1982; Baden, 1989). since 1844 (Ingersoll, 1993). Steidinger, 1948; 1882) (Davis, outbreak 1947 (Table to 1946 the until 2), named and identified but the causative dinoflagellate was not Karenia brevis brevetoxins by contaminated shellfish of consumption the by caused is (NSP) Poisoning first The bivalves. with only associated been has far, thus and, analogs brevetoxin or 1917 in made was tide red Florida a by caused irritation respiratory of report written become brevetoxins when effects respiratory from suffer can People 1917a). (Taylor, aerosolized through the disruption of Morris 1998; A. Haywood, personal communication). Thus far, no human fatalities have been attributed to NSP, and there have been no documented cases of NSP caused by raphidophytes. New NSP cases may occur when people consume unregulated humans in incidents NSP 1987, Until harvested. illegally shellfish or species shellfish were limited to the Gulf of Mexico (Steidinger, hectares of shellfish-growing waters along the Atlantic coast 1993). were closed to harvest In 1987 to 1988, 145,280 entrained an of because Satake that symptoms having NSP, of definition case the met people 186 and Island, North LANDSBERGlian models (Templeton REVIEWS IN FISHERIES SCIENCE onshore winds (Pierce, 1986; Pierce by paresthesia, reversal of hot-cold temperature sensation, myalgia, vertigo, ataxia, abdominal pain, nausea, diarrhea, burning pain in the rectum, headache, bradycar- dia, and dilated pupils (Hemmert, 1975; Steidinger

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al ., 1973). Menippe ) (Chang, ) was also , the crab Crassostrea et al blooms have Brachidontes Oliva sayana ; variable co- ., 1998a). The breve and other filter

; Fasciolaria lilium Fasciolaria (as cf. ; Atlantic surfclams, Atlantic ; et al brevis exposure. When the When exposure. . were eastern oysters, eastern were ., 1979). Other species Other 1979). ., K et al et , was significantly affected significantly was , brevis ; and stone crabs, brevis . Balanus eburneus . K ., 1998). In coquinas that have that coquinas In 1998). ., K mercenaria . . The molluscs The . Gymnodinium M et al et ., 1986; Sykes and Huntley, 1987). Huntley, and Sykes 1986; ., (Baden, 1988). Analysis of shellfish Chione cancellata , there was a noticeable increase in (crown ), and (as and ., 1979). Branchiostoma caribbaeum et al et Macrocallista maculata brevis . Ischadium recurvum K brevis brevis et al ; the barnacle . . 207 brevis , were exposed to between 10 to 25 million K K ., 1973, 1993; Hemmert, 1975; Roberts ., 1993). The dinoflagellate implicated in this Argopecten irradians Argopecten . cf. . K et al et al sp. and several amphipod species, including . PbTx-3 predominates in coquinas, but it forms only forms it but coquinas, in predominates PbTx-3 . were fed ; sunray venus, Melongena corona brevis ; cross-barred venus, Travisia . ; blue crabs, Polydora websteri K Crassostrea gigas ; hooked mussels, ., 1996) but is now being classified as a new species (Haywood species new a as classified being now is but 1996) ., sp. The cephalochordate Mercenaria campechiensis Mercenaria ., 1982; Tester and Fowler, 1990; Steidinger and et al et ., 1965; Steidinger lost muscle control and could not right themselves; mortalities varied mortalities themselves; right not could and control muscle lost

et al (Ray and Aldrich, 1967; Sievers, 1969; Roberts 1969; Sievers, 1967; Aldrich, and (Ray et al (banded tulip), cells/ over a 24 h period, NSP levels were 25 to 100 MU per 100 g ; quahogs, ; ); the annelid Donax variabilis ., in preparation). There have been several documented cases in which The recruitment of bay scallops, bay of recruitment The brevis . 1995; Haywood 1995; et al killed invertebrates in the field (Simon and Dauer, 1972; Steidinger Except for five benthic species invertebrate species of normally polychaetes present in and Tampa a Bay, immediately Florida, brachiopod, after USA, at a were absent red least tide. 17 Dominant Onuphis species killed emerita included the polychaetes Acanthohaustorius Crassostrea virginica recurvus affected (Simon and Dauer, 1972). Several experimental studies have investigated to species invertebrate of susceptibility differential the copepods (Huntley control motor of loss and rate heart Eurypanopeus depressus event was tentatively identified as identified tentatively was event of concentrations varying to exposure by Unaffected included systemic and general CNS, gastrointestinal, neurosensory, neurocerebellar, neurosensory, gastrointestinal, CNS, general and systemic included and neuromuscular effects (Bates mercenaria Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS have been acutely affected by exposure to exposure by affected acutely been have hunteria olive) (lettered not did tissues internal of histopathology and pathology Gross 69.0%. to 55.5% from reveal any differences between exposed and control animals. Crabs did not retain toxins when fed toxic shellfish (Roberts Spisula solidissima raveneli quinas, persistence of toxicity is species dependent and varies according to the concentra- tion of the bloom and the bivalves’ rates of feeding and elimination (Baden, 1983). When Pacific oysters, of drained oyster meat. After 3 days of depuration, oysters brevetoxin were concentrations considerably in reduced and almost (Fletcher at g 100 acceptable per MU regulatory 20 of limits consumption human for the from different are PbTx-3 and PbTx-2 of proportions the toxin, the accumulated in present proportions K about 28% of the toxic component in by the North Carolina red tide of 1987 (Summerson and , 1990). Bay scallops Bay 1990). Peterson, and (Summerson 1987 of tide red Carolina North the by are a potential health risk, but they are usually safe if people eat only the adductor muscle, which does not accumulate brevetoxins. Cases of NSP in the southeastern USA have been associated with the consumption of virginica eastern oysters, feeders. Brevetoxins accumulate in the gut and hepatopancreas in (McFarren these species 1979; Baden

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., ., et et et al et al brevis et al

cells and (Ishida Chattonella ., 1999) or in Karenia brevis ., 1995). BTXB2 1995). ., . et al K et al et ., 1998; Kimm-Brinson 1998; ., et al et ., 2002). ., 1995, 1999; Satake et al et al Austrovenus collected at the time of the NSP event. collected at the time

, ., 1975 in Baden, 1983). Evidence for in ., 1989; Warlen Warlen 1989; ., 208 et al et al et ., 2000). Although there is some information on et al , collected from Tiki Road in January 1993 and Rangaunu , in Delaware, USA (Bourdelais ., 1994, 1995), while PbTx-3 was detected in cockles (Ishida and (Ishida cockles in detected was PbTx-3 while 1995), 1994, ., ., 1998). Of the two major brevetoxins, PbTx-2 (= BTXB) and ., 1973; Quick and Henderson, 1975; Baden and Mende, 1982; et al et et al were implicated in a mortality of 1 to 2.5 million Atlantic menhaden, Atlantic million 2.5 to 1 of mortality a in implicated were ., 1975), although ingestion is another documented route (Tester et al Crassostrea gigas et al brain synaptosomes (Stuart and Baden, 1988). Signs of intoxication in fish Several brevetoxin analogs have been described from New Zealand shellfish. Brevetoxins are potent ichthyotoxins and have been responsible for the deaths Brevetoxin is thought to be absorbed directly across the gill membranes of fish verruculosa ., 1994, 1995) and was lethal to mice (i.p.) at a dose of 0.05 mg/kg. Mice showed was lethal to mice (i.p.) at a dose of ., 1994, 1995) and ., 1973; Quick and Henderson, 1975) (Table 2) and some life-history stages may the toxin deposition of brevetoxins in a few molluscs, there is no information on potential chronic health effect to these molluscs. cockles the in found was B1) (brevetoxin BTXB1 al irritability immediately after injection, followed (Ishida paralysis by respiratory by death hind-quarter and convulsions, dyspnea, paralysis, severe (brevetoxin B4) were isolated (brevetoxin B3), and BTXB4 (brevetoxin B2), BTXB3 from greenshell mussels, LANDSBERGextracts from an NSP outbreak in June metabolic conversion of the parent brevetoxin PbTx-2 into PbTx-3. 1996 No PbTx-2 was in Florida demonstrated the likely detected in the shellfish (Poli REVIEWS IN FISHERIES SCIENCE BTXB4 extracted from the hepatopancreas of toxic mussels was lethal to mice by i.p. injection at a dose of 0.1 mg/kg. Symptoms observed in mice were similar to those caused by BTXB2: paralysis of limbs, diarrhea, dyspnea, and convulsions. In accounted and metabolite brevetoxin toxic most the was BTXB4 mussels, greenshell for two-thirds of the mouse toxicity (Morohashi 1996a; Murata (Ishida cockles events are very common, widespread, and affect hundreds of species (Steidinger al (Riley others than susceptible more be PbTx-3, PbTx-2 was not detected in green mussels (Morohashi Harbour (northern North Island) in February 1994 and June 1995 (Ishida Tsuji, 1999) in New Zealand waters. PbTx-2 and PbTx-3 were also detected in Pacific in detected also were PbTx-3 and PbTx-2 waters. Zealand New in 1999) Tsuji, oysters, several although fragmentary, is fraction hemolytic the to related ichthyotoxicity situ of chronic species of fish collected during red tides in 1973 to 1974 displayed signs include violent twisting and corkscrew swimming, defecation pectoral fin paralysis, caudal fin curvature, loss and of equilibrium, quiescence, vasodi- regurgitation, lation, and convulsions, culminating in death due to respiratory failure. Chronically neurointoxicated fish show little pathology aside from slight precipitate hemolysis (Steidinger Baden, 1989). There is also some indication that there are hemolytic components, but these are not toxic to mice (Trieff 1996a, 1996b). of billions of fish over the years. Fish mortalities associated with 2000, brevetoxins produced by and Ramsdell, 2001). In July-September cf. Brevoortia tyrannus (Abbott 2000). Fish kills are postulated to originate with the lysis of toxic the subsequent tissues excitable liberation in sites receptor specific of to PbTx of toxin binding with begins and Intoxication passage across the gills (Baden, of fish 1988). (Baden and Mende, 1982); toxin binds with high affinity, Tilapia for example, to

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 M cells/ ; pin- , 1989) , Fibrocapsa et al. et [Morohashi brevis . cells/ml), fish K ., unpublished ., 1995], or 0.1

2 cells/ml, and 2.23 and cells/ml, et al 4 10 et al Chattonella marina × ., 1996a). 10 japonica × Fundulus majalis . et al F and various juvenile fish cells are required to yield

[Ishida

Type 2 brevetoxins (PbTx- brevetoxins 2 Type 3 ) during different seasons. 5 ) 10 10 × × brevis Tanichthys albonubes Tanichthys

strains involved, the component cells/ml) killed the fish in 2 to 4 h, 3 cells/ml, 1.67 cells/ml, 4 Danio rerio 10 brevis × cells/ml) (Landsberg ; striped killifish, 10 Karenia 3 . × , K 209 , 10 × neurotoxins, a significant decrease in heart rate Leiostomus xanthurus for Type 1 toxins (Baden, 1989; Baden 1989; (Baden, toxins 1 Type for M were traced through experimental food chains from marina ; spot, . brevis C . Gambusia affinis Gambusia ., 1989), probably due to depolarization of the vagus nerve. K ., 1992) instead of the significant pathology caused by reactive by caused pathology significant the of instead 1992) ., Labidocera aestiva produces 11 pg/cell, then 4.85 , et al Eucinostomus argenteus et al et were intensely toxic to these fish (Khan , were exposed to raphidophytes in different phases of the raphidophyte’s the of phases different in raphidophytes to exposed were , brevis . K was the most toxic during the mid-logarithmic phase and the least toxic antiqua . Lagodon rhomboides cells/ml) killed the fish within 66, 37, and 30 min, respectively. When red sea cells/ml) killed the fish within 66, 37, and 30 min, respectively. When red of toxin (Stuart and Baden, 1988), which is sufficient to kill fish. In ichthyotoxicity In fish. kill to sufficient is which 1988), Baden, and (Stuart toxin of C ., 1999]). 4 In order to examine differential brevetoxin-like production, juvenile red seabream, red juvenile production, brevetoxin-like differential examine to order In Brevetoxins from In some instances, mortalities caused by red tides are not immediate but may Unlike brevetoxins, thus far there is no evidence that brevetoxin analogs are M 10 turbinata showed abnormal movements for about half an hour, and then recovered gradually. recovered then and hour, an half about for movements abnormal showed Early-logarithmic-phase cultures (4.1 to 6.5 ml which is often considered to be a lethal concentration (Quick and Henderson, 1974). However, it is known that fish can die at lower cell concentrations (Morton and Burklew, 1969; Quick and Henderson, 1974) and can also apparently survive in much higher concentrations (at 3 et al major Pagrus growth cycle. From mid-logarithmic to early stationary phases, and japonica during the stationary phase. In 2-day cultures (1.68 (1.06 cultures 10-day and 8-, 6-, whereas × bream were exposed to was noted (Endo Exposure to neurotoxic fractions (Endo caused epithelium minor alterations in the gill lamellar oxygen species (ROS) (see section on ROS). (spotted mojarra, tissue damage and hemopathy. Chronic hemolysis was detected via anemia, cyanosis, evidence hyperviscous of blood, splenomegaly, hepatic hemosiderosis, and dehydration (Quick and Henderson, 1974, 1975; Baden, 1983). occur over a period of days or weeks of exposure to subacute concentrations toxins. Mortalities typically occur at a cell concentration of of 2.5 data). Some of these differences in toxicity level will depend on the susceptibility of each fish species to exposure the exposure; and toxins; extracellular of stability the the concentrations; cell and toxins, routes. If (see also Lewis, 1992, section on ciguatoxins). toxic not were shellfish Zealand New from isolated analogs Brevetoxin ichthyotoxic. to fish (100 ng/ml of BTXB1 for zebrafish, minnow, mountain cloud white for BTXB4 of ppm food this of generality The fish. juvenile to grazers, copepod through dinoflagellates, copepods of combinations different three using by demonstrated was transfer chain ( could be Juvenile spot were fed toxin-laden copepods so that vectorial intoxication examined. Toxins were shown to move from fish viscera to muscle tissue within Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS n 6 PbTx- PbTx-8, PbTx-3, (PbTx-2, brevetoxins 1 Type than potent more are PbTx-7) 1, the test population within 24 h were 3 to 5 n 9). Lethal concentrations for 50% of assays of mosquitofish, of assays n 37 to 10 and toxins 2 Type for fish,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ; , ., et al . Dur- cells/ ., 1992). ., brevis et al et

Opsanus beta Opsanus (assayed at 48 at (assayed red tide events tide red ., 1977). Anecdotal Aythya affinis brevis Karenia

. 4 Phalacrocorax auritus K et al 10 × , there is no evidence that brevis . K Mercenaria mercenaria Mercenaria ; and lesser scaup, red tide of October 1973 to May 1974, ., 2000). When gulf toadfish, gulf When 2000). ., 210 ., 1948; Rounsefell and Nelson, 1966). Cata- brevis et al et . ., unpublished data). K et al et al . Clinical signs included weakness, reluctance to fly, Mergus merganser Gemma gemma During the Florida west coast During 1946 to 1947 and 1953 to 1955, two of the largest the of two 1955, to 1953 and 1947 to 1946 During red-breasted mergansers, MU per 100 g tissue), or red tide water containing 2.2 slumping of the head, clear dysfunction, nasal excessive discharge, lacrimation, viscous chalky oral yellow diarrhea, discharge, tachycardia, dyspnea, oil decreased tachypnea, blood gland pressure, depressed body temperature, they toxins, tide red to ducklings Pekin white expose To dehydration. and reflexes, diminished quahogs, northern toxic with fed force were 1994). The use of different exposure protocols in experimental settings is critical to understanding brevetoxin toxicokinetics. Because experimental exposures do not necessarily mimic natural toxin that exposures, indicate misleadingly may care studies experimental some should example, For results. be taken in extrapolating the accumulates in the muscle, which would have important health. implications for However, public in natural exposures to ing an 8-week period, several thousand lesser scaup died. All scaup examined had substantial subcutaneous fat and normal breast muscles, which indicates that the birds died quickly. Most of contained the turritellid, ducks pyramidellid, had and fed opisthobranch recently, gastropods, gemclams, and and the amethyst proventriculi in addition to the usual reports of dead fish, large numbers moribund of or birds dead, particularly were double-crested cormorants, found When PbTx-3 was given orally to gulf toadfish in a fish-meal slurry, 40% of the total the of 40% slurry, fish-meal a in toadfish gulf to orally given was PbTx-3 When PbTx-3 body burden was concentrated in the hepatobiliary system, 27% in muscle, and 25% in the gastrointestinal tract. The hepatobiliary system therefore is the key of method the of regardless fish, in brevetoxin excreting and metabolizing for route toxin administration. Significant amounts of toxin were gastrointestinal also associated tract, with another the important site for detoxification (Washburn neither that determined been has it fish; of muscles the in deposited are brevetoxins PbTx-2 nor PbTx-3 is found in the muscle — both are principally concentrated in the intestinal tract (Landsberg were experimentally exposed to PbTx-3 by injection, the toxin was found principally found was toxin the injection, by PbTx-3 to exposed experimentally were concentration The tract. intestinal and stomach, gill, kidney, muscle, bile, liver, the in toxin of deposition the and time, with exponentially declined blood the in PbTx-3 of in tissues was were PbTx-3 of levels rapid high time, With h. 6 (1 after blood the in found h were metabolites following intravenous administration). No brevetoxin found in the gill and kidney, suggesting that both renal and branchial (Kennedy routes metabolites may its or brevetoxin of excretion the in important be reports of dead birds during red tide events are not documented unusual in the but scientific literature. are not always on record, in both geographical distribution and longevity, occurred in central and southwest Florida, USA (Gunter ml. The ducklings showed signs lethargic. On day of three, the ducklings toxicity showed signs of ataxia, 2 spastic movements days days few next the Over side. one to head the droop to tendency a and head, the of after exposure, appearing the ducks died and showed signs similar to those noted moribund in field birds observations (Quick of and Henderson, 1974; Forrester LANDSBERG (Tester h 25 to h 6 to 2 from periods REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 - /ml) 2 which brevis

, taken , 10 . × K ., 1991), and 1991), ., et al et cells (>1 Karenia brevis brevis . K Brevoortia tyrannus , virus) were found in numerous Vibrio ), sea turtles, and numerous fish species 211 , in 1963 (Layne, 1965), 1982 (O’Shea (O’Shea 1982 1965), (Layne, 1963 in , Tursiops truncatus ., 1998), when 7, 39, and 149 animals, respectively, died in was implicated in mortalities of the endangered Florida manatee, Florida endangered the of mortalities in implicated was et al ., 1948; Rounsefell and Nelson, 1966). Mass mortalities of dolphins were dolphins of mortalities Mass 1966). Nelson, and Rounsefell 1948; ., et al et Karenia brevis Karenia to be maintained. In the spring, as usually the disperse water downstream into temperature the warms inshore bays. up, If manatees red tide has come inshore along the Atlantic coast from June 1987 to May 1988. In the fall of 1987, in an unusual associated an dolphin mortality involved more in than 740 bottlenose dolphin 1987, strandings of fall the In 1988. May to 1987 June from coast Atlantic the along event, a Florida red tide was transported to North Carolina (see above), therefore exposing a wider range of dolphins and fish to toxins than are exposed when red tide is restricted to Florida. Dolphins migrating south in estimated the been has It fish. contaminated on there fed and Carolina North fall off bloom encountered the Florida, and Jersey New between stock migratory coastal the of 50% than more that cause proximate the be to considered was Brevetoxin period. this during died USA, agents ( of the mortality. Although infectious strophic mortalities of marine animals were recorded from Tarpon Springs to Key dead of reports were there events, these During coastline). of miles 150 (some West bottlenose dolphins ( (Gunter reported recently in north Florida (August to December 1999) when more than 100 animals stranded and brevetoxins were confirmed in dolphin tissues (B. Mase and T. Leighfield, NOAA, personal communications). In other years, there have numerous been reports of individual dolphin strandings. By far the largest temporal and spatial The brevetoxicosis. to secondary likely were these individuals, Contami- disease. infectious of patterns hallmark the lacked also mortality of patterns nant loadings (PCBs) were not considered to be abnormal during this event. Eight brevetoxins, for positive were mortality the during collected dolphins stranded 17 of and brevetoxin was found in the viscera of menhaden, from one dolphin stomach. No brevetoxin was detected in any of the control prey taken from dolphins that had died in captivity or prior concentrations to in this event. dolphin Brevetoxin livers ranged from 80 sublethally to exposed to brevetoxins 16,000 by ingesting toxic ng/g. fish and Dolphins then succumbed to were a range of chronic disorders, including fibrosis of the liver and lung, adhesions of the abdominal and thoracic viscera, and secondary microbial infections associated with immunosuppression (Geraci, 1989). latirostris manatus Trichechus 1996 (Bossart southwest Florida, USA, during the winter-spring. In 1963, in addition to manatee deaths, newspapers (Sarasota Herald Tribune, March 31, 1963) reported red tide in fish, and cormorants of mortalities turtles, of strandings and months, 6 for region the and neurotoxic symptoms in sick raccoons occurred coincidentally. The large-scale mortalities of manatees in 1982 environmental and conditions (Landsberg and 1996 Steidinger, inshore 1998). comes were usually concentrations, low at attributed offshore km 74 to to 18 develops typically a set of unusual do tides Red 1997). Steidinger, and (Tester dissipates then and fall-winter the during not usually appear inshore during the power winter-spring coastal the of months, waters warmer the in when areas, salinity zero or manatees low in congregated are Wilcox, plants, at warm-water spring refugia, or in residential canals (Reynolds and 1986). In the winter-spring periods appearances of inside 1982 the and barrier 1996, islands red of (above 24 ppt) allowed persistently high tide concentrations of southwest made Florida. unusual High-salinity areas Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 et ., 1998). The chronic mortalities of mortalities chronic The 1998). ., ., 1998). et al et et al 212 ., 1991). In 1982, it was suspected that filter- that suspected was it 1982, In 1991). ., et al et cells/ml (Steidinger, unpublished data) and the time the and data) unpublished (Steidinger, cells/ml 2 10 × ., 1998): brevetoxins were confirmed in manatee tissues, brevetoxin et al Several aspects of the manatee mortality events in 1982 and 1996 were different. were 1996 and 1982 in events mortality manatee the of aspects Several Although acute exposure to lethal doses of brevetoxin results in massive animal massive in results brevetoxin of doses lethal to exposure acute Although ., 1991). In 1996, the duration of the mortality event was coincident with the LANDSBERGduring this period (as occurred in 1982 and 1996), then the likelihood of manatees being exposed to tide red the red to proximity their and to move manatees the areas tide which on depending during their post-winter movements bloom. is fairly high, In 1982, there was a lag of 3 weeks between the time that the red tide cell counts 1 below to decreased had REVIEWS IN FISHERIES SCIENCE that the last manatee died (O’Shea died manatee last the that manatees to toxic still were they that and brevetoxins accumulated tunicates feeding some 3 weeks after seagrass. the to attached the were that tunicates toxic to dissipation exposed also were but seagrass of the bloom. Manatees normally consume Sick and dying manatees were observed throughout the red tide event (O’Shea al duration of the red tide manatee last the between time bloom lag significant no was There affected. that were manatees occurred in the same the At 1998). Steidinger, and inshore (Landsberg tide red the of areas dissipation the and mortality where gross level of examination, severe nasopharyngeal, pulmonary, hepatic, renal, and cerebral congestion were present in all manatees examined. Nasopharhyngeal and pulmonary edema and hemorrhage and hemosiderosis, were multiorgan edema, seen. and hemorrhage Microscopic pulmonary rhinitis, catarrhal lesions consisted of nonsuppurative leptomeningitis. Immunohistochemical staining with a polyclonal antibody to brevetoxin showed intense positive staining of lymphocytes and mac- rophages in the lung, liver, and secondary lymphoid tissues. Additionally, lympho- cytes and macrophages associated with inflammatory lesions of the nasal mucosa and meninges were also brevitoxicosis as positive a component of and the for likely primary etiology for the brevetoxin. epizootic These (Bossart findings implicated mortalities, effects from low-level exposure to brevetoxins are harder to interpret. at all Chronic dietary exposure to brevetoxins could exert lethal or sublethal effects physiological behavior, avoidance bloom feeding, impaired to leading levels, trophic dysfunction, impaired immune function, reduced growth and reproduction, patho- logical effects, or mortality. There is some evidence for immunosuppressive effects influence an suggests and brevetoxins to exposure chronic persistent with associated (Bossart term long the in health manatee on dolphins in 1987 together with numerous secondary health effects in dolphins are also suggestive of longer-term toxicity exposure (Geraci, 1989). shows high affinity for binding to manatee brain nerves (Trainer and Baden, 1999), shows high affinity for binding to manatee and manatee tissue extracts were toxic to mice in bioassays (Baden, unpublished data). The coincidence of the mortality with the red tide bloom would suggest that manatees were exposed to high levels of brevetoxins through inhalation. Residual toxin bound up in the food web, water, or substrate would persist after the bloom had dissipated, but aerosolized disperse brevetoxin relatively that quickly manatees (Landsberg could brevetoxins in nasal and and lung tissue implicate the aerosol, and lesions of inhale the upper Steidinger, would 1998). The respiratory presence tract were the only of severe and consistent inflammatory lesions seen in the manatees from the epizootic (Bossart

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 Liza ., 1978, ., 1996; Karenia Karenia et al et al Tanichthys ., 1995, 1996; 1995, ., ., 1996). Mice Gymnodinium Gymnodinium ., 1996). When 1996). ., , from Foveaux et al et et al (= et al et ., 2001) present at sp. (as sp. sp. (A. Haywood and sp. is the same species same the is sp. et al sp.) (Seki sp.) sp.), debilitating and lethal Karenia ., 1996a). The examination of Karenia ., 1996a; Seki Karenia ., 1996a). Thus far, there is no Tiostrea chilensis ., 1996a) but the potential public ., 2000]) (Haywood et al et al et al et al Karenia mikimotoi sp.) (Heil et al ) and wild largescale mullet ( sp.) cultures or oysters fed with with fed oysters or cultures sp.) sp. (A. Haywood and K. Steidinger, Gymnodinium Gymnodinium 213 ., 1996). Gymnodimine showed no hemolytic no showed Gymnodimine 1996). ., cells/ml. This cells/ml. 3 Karenia gymnodimine, the toxin targeted and damaged and targeted toxin the gymnodimine, 10 et al et M sp. (as × ., 1996). It is not yet clear what the mechanism of Gymnodinium g/kg (Mackenzie Gymnodinium µ et al ., 1996a). Gymnodimine was also a potent ichthyotoxin Karenia sp. (as sp. ., 1995, 1996; Mackenzie et al sp.) bloom (Mackenzie Acanthopagrus cuvieri sp.) were lethal to mice. The minimum lethal dose of gymnodimine of dose lethal minimum The mice. to lethal were sp.) et al ., 1996). These symptoms were identical to those shown by mice that mice by shown those to identical were symptoms These 1996). ., Karenia sp. known to produce gymnodimine in New Zealand (A. Haywood, ., 1996a) were associated with this toxicity. this with associated were 1996a) ., et al et were exposed to 2 m 2 to exposed were ), were reported in Kuwait Bay, Arabian Sea. The mortalities coincided et al et , at 250 to 500 ppb (pH 7) or 50 to 100 ppb (pH 8) (Seki 8) (pH ppb 100 to 50 or 7) (pH ppb 500 to 250 at , Hansen and Moestrup [Daugjberg Gymnodinium Karenia Gymnodinium From September to October 1999, wide-scale mortalities of cultured fish, When larvae were exposed to whole cells, culture filtrates, or albonubes . 1979b) results from the consumption of toxic shellfish containing okadaic acid (OA) acid okadaic containing shellfish toxic of consumption the from results 1979b) personal communications) (as F. OKADAIC ACID ANDDiarrheic (diarrhetic) shellfish poisoning (DSP) in humans (Yasumoto DERIVATIVES (DINOPHYSISTOXINS) Mackenzie mikimotoi crude using bioassay Mouse communications). personal Steidinger, K. and Haywood A. either of extracts (as sp. injected i.p. into mice was 450 macrolepis with a bloom of the dinoflagellate 6.0 > of concentrations maximum as the personal communications). Although the Kuwait strain has not yet been tested for toxicity it is quite likely that gymnodimine was associated with the fish kills. evidence that there are gymnodimine-contaminated shellfish human (Mackenzie health risks associated with the consumption of gymnodimine toxicity is nor what impact the toxin has under natural conditions. principally sobiaty ( E. GYMNODIMINE In 1994, screening of contaminated dredge oysters, Strait, New Zealand (South Island), led to the discovery of a novel toxic compound — gymnodimine (Seki phytoplankton samples showed that high numbers of (as communications) personal Steidinger, K. sp.) bears a close morphological relationship to and jumping, tail, curled symptoms, neurological with presented and min 15 to 5 in died (Seki paralysis a from gymnodimine with contaminated naturally shellfish to exposed been had sp. (as health threats still need to be considered. sonicated cell extracts of swimming primary the shed larvae the exposure, of min 30 Within occurred. effects after h 24 to 7 occurred death disintegrated; gradually margins mantle their and cilia exposure (Mackenzie to a freshwater fish, the white cloud albonubes mountain minnow (= red fin), T both the gill lamellar epithelial cells and the chloride cells. The severity of lesions (Terao dependent calcium not was activity or cytotoxicity (Seki Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et spp. et al ., 1985). ., spp. are et al et Dinophysis cells in the water, the in cells ., 1989; Yamashita Dinophysis ., 1995). When mice When 1995). ., ., 1990). DTX-1 causes et al lima . et al et P in tropical regions (Table

et al (Table 1). spp. or or spp. is the primary source of DTX-1, is cosmopolitan. DSP cases usually produce mainly OA. When bivalves When OA. mainly produce species found lima . ., 1988a; Lange P 214 Dinophysis ., 1989; Herschmann et al acuminata et al . Dinophysis fortii Prorocentrum lima D and Prorocentrum are suspected in incidences of ciguatera (Table 1) (see acuta . D ., 1989; Haystead ; and (4) induces various mitosis-specific events (Bialojan and Takai, et al Prorocentrum in vitro ., 1990; Sakai and Fujiki, 1991; Fujiki and Suganuma, 1993; Rossini, 2000). OA In the short term, when humans consume DST, they can experience gastrointes- experience can they DST, consume humans when term, short the In There are few reports implicating DST in animal mortalities or as having Okadaic acid inhibits protein phosphatases types 1 and 2A, thus increasing ., 1990, 1993). typically temperate in distribution, and occur in temperate climates (Aune and Yndestad, produced 1993). by a few benthic OA and OA- DTX benthic the of Some are DSP. in involved be to also known not are species these but 1), producing section on ciguatoxins) or as possibly promoting tumor development (see below). tinal distress, diarrhea, nausea, and vomiting (Aune and Yndestad, 1993; Quilliam protect to exposure DST minimize to strive programs Monitoring 1995). Wright, and public health and have developed of counts monitoring by either action shellfish plans to estimate exposure levels in and the epibenthic dinoflagellate and the epibenthic by determining DST levels in the meats, or both (Shumway both or meats, the in levels DST determining by et al control that proteins of phosphorylation the stimulating by diarrhea causes probably cytoskeletal of phosphorylation enhancing by or cells intestinal by secretion sodium or junctional elements that regulate permeability to solutes, passive loss of thereby fluids (Aune resulting and Yndestad, in 1993). sublethal effects. Although numerous studies have threats been of DSP to human health through made shellfish consumption, the potential effect on the potential of DST on molluscs has not numerous been species well of investigated. bivalves DST (Shumway, have Europe, 1990). Japan, and North America, the principal toxins have been either been OA, DTX- In found separate in DSP outbreaks species of 1, or DTX-3. Dominant toxins in the shellfish will vary depending on the in microalga to which the molluscs produces. For example, in Japan, are exposed and the toxins each microalga LANDSBERGand the OA and DTX derivatives toxins). shellfish (diarrheic DST dinophysistoxins as to referred be (DTX1-4) can toxins these Together (Aune and Yndestad, 1993). dinoflagellates planktonic the by produced toxins major the are OA REVIEWS IN FISHERIES SCIENCE whereas in Europe in whereas were experimentally exposed to OA, DTX-1, and DTX-3 administration, short-term either effects orally on the or digestive by tract were i.p. similar (Terao (Hamano mice suckling of intestines the in accumulation fluid excessive Within 15 effects min Potent occurred. diarrhea of severe by accompanied exposure villi ileum the to of epithelium the toxin, marked destruction of the of DTX-3 in the liver absorptive led to degeneration of the hepatocytes within 24 h (Terao al protein phosphorylation that (1) affects intracellular processes, including metabo- lism, membrane transport and secretion, contractility, gene transcription, mainte- nance of cytoskeletal structure, receptor-mediated signal transduction, and cellular division; (2) stimulates the expression of certain proto-oncogenes; (3) activates H1 kinase 1988; Fujiki 1993). OA induces rapid changes in the small and large intestine of the rat, resulting rat, the of intestine large and small the in changes rapid induces OA 1993). in hypersecretion, selective shedding of the enterocytes at the top of the villi, and accumulation of goblet cells (Edebo

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al Mytilus g OA/g g/g wet g/g Artemia Artemia . (1995) ., 2001). µ µ cells also Dinophysis Dinophysis Argopecten g/g on day µ et al et al Prorocentrum Prorocentrum lima g/g in the gills, the in g/g

µ accumulated OA lima . cells/ml. Adult P 4 cells/ml was reported in 3 10 cells/ml for 2 days) total DSP total days) 2 for cells/ml Prorocentrum × 10 1 bioassays using bioassays × 10 transformed DTX-1 into DTX-3 × to 1 ., 1990; Aune and Yndestad, 1993), and Yndestad, ., 1990; Aune 1 (8 10 et al × ., 1988b). When bay scallops, bay When 1988b). ., 215 et al et Artemia franciscana Artemia ., 1988b; Alvito ., 1988b; . (1995) found that more than 50% of 1-day-old of 50% than more that found (1995) . Prorocentrum lima lima Prorocentrum Patinopecten yessoensis et al nauplii and metanauplii. Therefore, Demaret et al et , there is a strong likelihood that fish are exposed to these , but this has not been confirmed since its original isolation by isolation original its since confirmed been not has this but , ., 1995). Blue mussels appear to have some protective mecha- , filtration rates decreased after 1 h, most likely because of the et al g/g in visceral and gonadal tissues and less than 0.1 than less and tissues gonadal and visceral in g/g Artemia . (1992). µ ., 1999). concavum ., 1990; Suzuki and Mitsuya, 2001), can be seasonally toxic (Sato . P with maximum cell concentrations of 1.5 et al , were exposed to exposed were ,

et al , et al , in Swedish waters remained toxic for up to 5 months after toxin accumulation toxin after months 5 to up for toxic remained waters Swedish in , , Japanese scallops and cultures. Demaret cultures. In the summer of 1993 and lasting into 1994, a large bloom of Toxins are also modified by bivalves. After experimental exposures to exposures experimental After bivalves. by modified also are Toxins reef tropical In fish. in OA of distribution the regarding information little is There As with other aquatic groups, there is little information on the potential effects As with PSP, different bivalve species accumulate varying concentrations of DST of concentrations varying accumulate species bivalve different PSP, with As concluded that DSP toxins were potentially highly toxic to zooplankton. caudata rapidly killed mantle, and adductor muscle. There were no observed detrimental physiological effects or mortalities during the exposure period of 13 days (Bauder were also rapidly affected, with 77% mortality in 5 h. there tintinnids, of species few a for Except India. of coast east the on Bay, Tuticorin toxin concentrations (including OA, DTX-1 and okadaic acid diolester) 1 were proximately ap- fortii (Suzuki systems, where OA is produced by benthic dinoflagellates such as lima toxins through the diet (see ciguatera). Few studies have investigated the presence of DST in fish, unless the studies were conducted in combination with ciguatoxin assays. There has been one report barracuda Sphyraena of OA in a carnivorous fish, Gamboa the barracuda of DST on zooplankton. A potentially lethal effect of OA on zooplankton has been experimental from extrapolated but only a few studies have investigated their potential effects on aquatic organisms. aquatic on effects potential their investigated have studies few a only but (Alvito feed on microalgae containing OA or DTX, the toxins are accumulated mainly in the in mainly accumulated are toxins the DTX, or OA containing microalgae on feed (Edebo hepatopancreas 1996), and may remain toxic for different periods of time. edulis Blue mussels, mussel Swedish in found are OA of concentrations high Although 1990). (Shumway, populations for several months every year, the toxins are not associated with any known health effects. When blue mussels were exposed to high concentrations of Prorocentrum lima toxicity associated with inhibitory or cytotoxic 1995). Blue mussels that were effects experimentally exposed to (Pillet and Houvenaghel, lima larvae died after they were exposed to 5.0 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS and DTX1 in the hepatopancreas. A slow buildup of toxin, from 1.2 to 2.0 to 1.2 from toxin, of buildup slow A hepatopancreas. the in DTX1 and weight occurred in the hepatopancreas by day 14, with a peak of 3.8 with associated was mortality no and documented, not was accumulation Further 16. exposure (Pillet nism whereby the activity of certain enzymes, such as glycogen synthase, are not affected by OA (Svensson and Förlin, 1998). Mussels can also have rapid clearance mechanisms; in one report, concentrations of OA declined from 7.2 to 1.8 (Edebo week 1 within hepatopancreas irradians

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ) et and . The nana . O ., 1999). lima PTX1-7 andPTX1-7 .

serrata et al Prorocentrum . P and F Oithona nana Oithona ., 2000). species (Table 1). et al in the Galician Rias and ., 1989; Suganuma et al longicornis . T ., 1999). Dinophysis spp., whereas other copepods acuminata et al . ., 1982, 1986). The bioorigin of ., 1988). An epidemiological study D et al Temora et al . (2000) recognized the need for more did not. During the bloom, populations of populations bloom, the During not. did . (2000) determined that several zooplank- several that determined (2000) .

Dinophysis ) 216 et al et al et fed on m showed a close correlation with µ ., 1985; Murata et al Euterpina acutifrons Euterpina Favella serrata and spp. are epiphytic on the macroalgae and that are normal spp. are epiphytic on the macroalgae , and high-risk FP areas are linked to areas where are both widespread and abundant. The presence of OA in the tissues the in OA of presence The abundant. and widespread both are In addition to the involvement of OA in acute human shellfish poisoning events, poisoning shellfish human acute in OA of involvement the to addition In concavum ., 1990; Sakai and Fujiki, 1991; Fujiki and Suganuma, 1993). DTX has also been . Acartia Because both PTX and DTX have often co-occurred in shellfish, the potential specific potential the shellfish, in co-occurred often have DTX and PTX both Because defined. clearly been not has organisms these of health the on PTX of impacts chemically been have PTXs 10 least at and shellfish from isolated been have PTX-10 pectenotoxins has just recently been confirmed to be G. PECTENOTOXINS For many years, researchers have been aware of (PTX) in the toxic shellfish presence poisoning events, and of PTX were implicated pectenotoxins in some cases of DSP (Yasumoto ( and the tintinnid tintinnids noticeably increased, whereas densities of LANDSBERGwas a total absence of zooplankton as well as a general decline in the number of diatoms and fish during the bloom (Santhanam and situation was Srinivasan, also recorded 1996). during a A bloom of similar REVIEWS IN FISHERIES SCIENCE ton species could ingest and transfer toxins through the pelagic food web. Grazing ( copepods some that indicated experiments Bajas in northwestern Spain. Maneiro Spain. northwestern in Bajas results suggest that tintinnids can play a significant role in the transfer of okadaic acid toxins to higher trophic levels in the food web (Maneiro declined dramatically. Okadaic acid content found in seston size fractions 100 to 200, to 100 fractions size seston in found content acid Okadaic dramatically. declined 200 to 300, and 300 to 1000 there is an increasing awareness of the potential role of OA and OA derivatives as tumor promoters. In two-stage carcinogenesis experiments, OA has been shown to induce skin papillomas and carcinomas in mice and adenomatous hyperplasia and adenocarcinomas in the glandular stomach of rats (Fujiki al to shown to induce tumors in mice (Fujiki of Hawaiian green turtles indicates exposure and that this tumor-promoter may have may tumor-promoter this that and exposure indicates turtles green Hawaiian of a potential role in the etiology of FP (Landsberg of digestive-tract cancer mortality in relation to the distribution of DSP was recently of digestive-tract cancer mortality in conducted in , and although there appeared to be a very tentative positive association between the two, Cordier is a debilitating neoplastic disease that has Fibropapillomatosis (FP) in green turtles been has FP but unknown, is FP of etiology The worldwide. levels epizootic reached ( dinoflagellates benthic Toxic 1994). (Herbst, viruses oncogenic to linked spp.) are not typically considered tumorigenic agents, but Benthic they 1993). have Suganuma, a and worldwide (Fujiki OA tumor-promoting produce and distribution Prorocentrum components of green turtle diets. In the Hawaiian Islands, green turtles consume Prorocentrum P extensive surveys and in-depth research before any link can be definitively proven. extensive surveys and in-depth research has animals marine in development tumor in toxins microalgal of role potential The also been discussed recently (Landsberg, 1995, 1996; Landsberg

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al Mytilus ., 1989; ., 1991). ., ., 1991), ., and the species are et al et et al et ., 1996) and et al ., 1990), and species have Prorocentrum (Table 1), and 1), (Table ., 1993a, 1993c; 1993a, ., et al toxicus et al . et al et G toxicus (Holmes . ), further supporting a G Prorocentrum Prorocentrum (Holmes ., 1988; Hu toxicus . G species, each of which produce ., 2001). et al (Table 1) pass up the food chain toxicus . et al Scarus gibbus G ., 1999); some of which have been species have been described recently and Mediterranean mussels,

toxicus toxins. When toxic herbivorous fish are , (as

. 217 et al G strains from the same areas can produce Prorocentrum ., 2000), maitotoxin (MTX1-3), gambierols, and cells or et al , and ., 2000; Suzuki ., 2000). PTX-2 is converted into PTX-1, PTX-3, PTX-6, Gambierdiscus ., 1997c). At least two types of maitotoxin (MTX-1 and (Table 1), which inhabits reefs and hard grounds in et al et al Coolia et al , is the only genus that has been definitely implicated as an Chlorurus gibbus Perna canaliculus Gambierdiscus (Draisci Gambierdiscus toxicus ., 1993b; Lewis Ostreopsis ., 1991). et al et al Gambierdiscus Gambierdiscus toxicus Although several other defined thus far (Draisci PTX-2 seco acid (PTX-2SA) and other derivatives greenshell by shellfish, mussels, such as scallops, galloprovincialis H. CIGUATOXINS tropical of consumption the by caused poisoning food human of type a is Ciguatera C-CTX) (Pacific or Caribbean with ciguatoxins (P-CTX or fish that are contaminated origin respectively) (Lewis and Holmes, 1993; Swift and Swift, 1993; Lewis Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 2000). There are a suite of ciguatera toxins that originate from the benthic dinoflagel- benthic the from originate that toxins ciguatera of suite a are There 2000). late 1987). Ciguatoxin precursors regions (Anderson and Lobel, subtropical and tropical and maitotoxin (MTX) originating from when herbivorous fish browse on substrates or on primary consumers been that exposed have to consumed by carnivorous fish such as barracuda or grouper and the ciguatoxins are ciguatoxins the and grouper or barracuda as such fish carnivorous by consumed bioaccumulated in their muscles, they pose a risk to human consumers (Lewis and not has (P-CTX-1) se per ciguatoxin Although 1993). Swift, and Swift 1993; Holmes, of cultures experimental or wild from isolated been (Satake P-CTX-4B) and P-CTX-4A, (P-CTX-3C, ciguatoxins other Yasumoto (Holmes strains certain in detected been have 1) (Table acids gambieric CTX-4A (scaritoxin) has been heavybeak confirmed parrotfish, recently in both food-chain link (Satake MTX-2) have been isolated from Australian Holmes (Faust, 1995; Holmes, 1998; Chinain demonstrated to produce cigua-like toxins (Table 1), it is unclear at this stage what role they may have in ciguatera. Also known dinoflagellate to be community part of (Tindall the tropical and benthic ciguatera are Morton, 1998) and often their own unique toxins and toxin derivatives (Table 1). CTX has not implicated been isolated in for except dinoflagellate benthic of species other any from additional water-soluble toxins have been isolated from Caribbean strains (Miller and (Miller strains Caribbean from isolated been have toxins water-soluble additional Tindall, 1988). precursors of varying toxicities in experimental bioassays (Bomber thus far origin of ciguatera (Lewis and Holmes, 1993). Benthic common in tropical communities; those producing okadaic previously acid (see okadaic acid). were In addition to OA discussed and its derivatives, which prorocentrolide, toxins fast-acting the produce to shown been also have spp. is lethal to mice in experimental assays (Torigoe hoffmanniolide (Table 1), but it is currently unclear how these toxins animal health may in the affect wild. Although additional benthic

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 Ostreopsis species have species and , in Madagascar , are also suspected Coolia Ostreopsis was shown to produce ., 2001). Palytoxin or its and spp. as tumor promoters in et al lenticularis . ., 1998), are toxic to mice (Tindall mice to toxic are 1998), ., Coolia O borbonicum borbonicum

et al et , and molluscan larvae (Rhodes and Ostreopsis are necessarily involved in ciguatera, there ciguatera, in involved necessarily are species produce hemolysins, ostreotoxins, hemolysins, produce species 218 , such as Artemia ., 1995; Ukena ., 1995). Two neuroactive compounds isolated produces toxins that are lethal to mice (Tindall Coolia et al et al Prorocentrum Herklotsichthys quadrimaculatus Ostreopsis Ostreopsis , and , ., 1995), monotis ., 1999). Palytoxin is also a potent tumor promoter (Fujiki promoter tumor potent a also is Palytoxin 1999). .,

et al et al et were demonstrated to interact with nicotinic cholinergic recep- to interact with nicotinic cholinergic were demonstrated Coolia Ostreopsis was characterized as a palytoxin analog, which implicates , ., 1995). Several 1995). ., ., 1995). Some ., 2002). et al et siamensis species as a likely biogenetic origin for palytoxin (one of the most lethal et al lenticularis . . O et al O ., 1984; Holmes ., 1990), and have been demonstrated to have neurotoxic effects on chick ., 1984b), and so the potential role of Prorocentrum Because top-level carnivorous fish may have consumed a variety of herbivorous of variety a consumed have may fish carnivorous top-level Because is a potential for their toxins to reach human tropical consumers benthic via dinoflagellate toxic communities fish. include Because numerous toxic genera, reef- dwelling organisms have the potential to be exposed to numerous toxins. Most of macroalgae, corals, as such substrates, of variety a inhabit dinoflagellates benthic the and sand (Carlson and Tindall, 1985; Anderson and Lobel, 1987), and are inadvert- ently consumed by benthic browsers combinations such of toxins as that are herbivorous potentially fish. available to Therefore, considerable. higher trophic the levels are fish species during the course of their lifetime, the CTX complex toxins that have fish may been biotransformed, bioaccumulated, and deposited in the muscle of the be present in different proportions and combinations. The families presence of of ciguatoxins different in ciguateric fish from the Caribbean Sea probably and underlies the Pacific the clinical differences reported in in the the two ciguatera regions syndrome (Vernoux as and Lewis, 1997). complex comprises multiple toxic components Because in fish, their effects on the humans can ciguatoxin though even 1993]), Swift, and [Swift identified been have symptoms (>175 varied be Human fish. toxic from characterized or identified been have toxins the of few a only consumption of toxic herbivorous fish is usually fish is illness or neurological symptoms, whereas consumption of toxic carnivorous associated with gastrointestinal borbotoxins that block postsynaptic nicotinic acetylcholinesterase receptors (Ten- Hage isolated D ostreocin Recently, 1). (Table toxins uncharacterized other and ostreocin, from analogs were recently implicated in a fatal human poisoning caused by sumption the con- of toxic sardines, (Yasumoto, 1998). Consequently, palytoxin has been inferred to be toxin the source possible in clupeotoxism, a fatal human intoxication (Onuma caused fish clupeoid by ingestion of Other considered. be also should systems aquatic been described (Faust, 1995, 1999; Faust and Morton, 1995), tested for toxicity. but Currently, the have potential harmful effects not of been species on aquatic organisms are unknown. Although it is not currently considered that LANDSBERGbeen described recently, these have not all been tested for 1993a, toxicity 1993b, (Faust, 1994). 1990, Recently, REVIEWS IN FISHERIES SCIENCE et al et al Thomas, 1997), but there is no indication that this species is involved in ciguatera (Holmes from tors (Type-I extracts) and voltage-dependent sodium channels (Mercado (Type-II extracts) Ostreopsis marine toxins known) (Usami in ciguatera cases in the Caribbean (Tosteson Caribbean the in cases ciguatera in et al embryo neurons (Rivera Rentas

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., 1988; Lewis, 1992). Lewis, 1988; ., et al et were fed toxic red snapper,

, , pure CTX-1, CTX-2, or PbTx-2 ., 1979). When mice are injected are mice When 1979). ., . (1988), who suggested that a et al et et al ., 1980). The relevancy of these earlier tests earlier these of relevancy The 1980). ., 219 ., 1986, 1988; Capra 1988; 1986, ., et al et Gambusia affinis et al et ., 1967; Bagnis 1967; ., ., 1985). Ciguatoxins are potent activators of voltage- of activators potent are Ciguatoxins 1985). ., et al et et al et ., 1966) suggested that there was no effect because when because effect no was there that suggested 1966) ., Acanthurus xanthopterus ., 1966; Swift and Swift, 1993). Earlier studies (Helfrich and ., 1966; Swift and Swift, 1993). Earlier ., 2000; Terao, 2000), and except for references pertinent to et al et et al et al , no overt pathology was detected. It was assumed that herbivorous ) in the water with a dose of 17 MU per ml died within two hours (MTX) For many years, it was generally assumed that ciguatera toxins were sublethal Recent experiments have shown that CTX can be lethal to fish when adminis- toxicus . more often associated with cardiovascular and neurological disorders (Bagnis, 1968). (Bagnis, disorders neurological and cardiovascular with associated often more Symptoms include paresthesia, arthalgia, myalgia, headache, diarrhea, nausea, asthenia, pruritus, chills abdominal pain, and vomiting, perspiration, giddiness tearing, (Swift and and Swift, 1993). In fatal cases, there are no remarkable gross or (Tonge changes histological Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS with lethal doses of acetone-extracted or methanol-extracted ciguatoxins, ciguatera symptoms observed are loss of activity, diarrhea, breath- gasping, choking-gasping penile labored cyanosis very and and/or ataxia, erection, incomplete and transitory ing. Respiratory failure is the cause of death because the heart is still beating after (Vernoux arrest respiratory dependent sodium channels in a variety of tissues, results from binding of especially the ciguatoxins to site 5 nerves. on the sodium channel (see Activation Lewis and Holmes, 1993), a site also attacked by the brevetoxins (Baden, 1989). There is and pathology, pharmacology, epidemiology, history, the on information of wealth a Withers, (see dinoflagellates benthic ciguatera-associated and ciguatera of toxicology 1988; Bomber and Aikman, 1989; Miller, 1991; Lewis and Holmes, 1993; Swift and Swift, 1993; Lewis (Bagnis (CTX) minutes 70 to 60 and potential harmful effects on aquatic organisms, further. ciguatera will not be discussed or lethal to the humans at the top of the food chain because they are exposed to high doses of bioaccumulated toxin and organisms at lower unlikely to trophic be affected levels by lower concentrations were of toxin. Randall (1958) proposed a hypothesis to link ciguatera transfer via the food chain. Three established corollaries were to support the (2) hypothesis: consumption of toxin (1) has no effect fish on fish, and (3) acquire CTX bodies of is fish in an unaltered stored CTX state (Helfrich and Banner, within 1963). For the most the part, via their diet, although even now the potential effects the second assumption has been accepted, from benthic dinoflagellates on fish have been of ciguatera toxins (other than CTX) fish on effect no have to reported been have toxins Ciguatera minimally. investigated in the wild (Banner Banner 1963; Banner, herbivorous surgeonfish, Lutjanus bohar any without toxin the carry would and asymptomatic behaviorally remain would fish adverse effects (Helfrich and Banner, 1963). However, the method of toxin admin- istration, that is, carnivory by a herbivore, would not be a normal toxin exposure mechanism in the wild. Guppies of exposed to purified MTX and CTX (from wild G to field conditions was questioned by Capra added to water induced similar signs, including pronounced opercular movement (suggestive of respiratory distress), inactivity, darkening of the skin, bursts of dietary exposure route would be more appropriate. tered orally, by intraperitoneal injection, or when dissolved in aquarium water (Davin in swimming were fish which In fish bioassays with mosquitofish,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., ., ., et et al et al et al et al Oryzias ., 1993b; ., 1993) and 1993) ., et al ., 1994), but et al et et al , that persisted for the ., 1994; Goodlett et al ., 1992; Yasumoto were not studied (Durand-Clement studied not were et al ., 1992; Lewis, 1992; Magnelia ., 1988; Gonzalez . (1999) suggested that the sensitivity ., 1998). Scaritoxin (SCTX = CTX4A), ., 1987). This effect appears to be similar be to appears effect This 1987). ., Gambusia affinis et al et al et al et al toxicus cells) to abnormal swimming behavior, . 220 ., 1997c; 1998a). Because it is water soluble, et al et G . Of all the toxins known to be produced by ., 1994). In addition to behavioral effects, CTX et al (Capra showed responses ranging from death within 24 h toxicus . et al and produces other electrophysiological changes in + (48 h) to mosquitofish for CTX-1, CTX-2, and PbTx- G ., 1990; Legrand (Marquais cells or were exposed to lethal doses of CTX dissolved CTX of doses lethal to exposed were or cells 50 ., 1989; Kelly ., 1987, cited in Gonzalez /l, respectively, indicating that in this assay the ciguatoxins the assay this in that indicating respectively, /l, et al M et al et al ., 1987; Flowers 1987; ., toxicus Gambierdiscus . ., 1977; Satake G et al et Caranx latus et al ., 1997c). Recently, a new toxic component, C-CTX-1, was isolated from (Table 1), only CTX and CTX analogs have been identified repeatedly in Gambierdiscus toxicus Gambierdiscus toxicus et al ) embryos, those exposed to 0.1 to 0.9 pg/egg (ppb) had cardiovascular, s between 16 and 30 ., 1987, cited in Gonzalez The specific effects of each of the toxic components originating in benthic 50 toxicus ., 1986, 1988; Kohler . (LD al 1992; Durand-Clement 5-day duration of the experiment (Lewis, 1992). Tilapia juveniles fed four different strains of G carnivorous fish (Murata in aquarium water. Fish behaved abnormally, exhibiting erratic movement, disori- entation, inactivity, and loss of equilibrium as well as experiencing physiological distress such as blanching or darkening of the skin and loss of appetite (Davin (Capra nerves fish to the CTX effect ciguatoxins on were microinjected mammalian into the tissues egg (Capra yolk of and Japanese rice Cameron, fish ( 1985). When of embryonic fish to CTX via direct oocyte exposure transfer may of indicate low that levels of maternal ciguatoxin is an unrecognized threat to the reproductive success of reef fish. dinoflagellate species are not are these because and 1) (Table species dinoflagellate benthic tropical by produced well characterized. Because multiple transformed through the toxins food chain, fish may be exposed are to a variety of toxins in the and level, trophic each at transformed be can Ciguatoxins combinations. various their from modified somewhat be may thus ciguatoxins as to referred toxins of suite original structure in 2 were 0.5, 2.1, and 10 n 10 and 2.1, 0.5, were 2 were up to 20-fold more experiments, pathological changes potent took place in the livers than of carnivorous fish that the brevetoxins ingested were (Lewis, 1992). In carnivores would not necessarily be expected to have herbivorous, direct crustaceans, contact in with observed benthic been have changes Behavioral dinoflagellates. or carnivorous fish after these organisms were allowed to feed either on ciguatoxic on directly or flesh 1994). Sublethal doses of CTX-1 or CTX-2 induced signs of respiratory distress and a loss of righting reflex in mosquitofish, quiescence, loss of equilibrium, and loss of appetite. Fish showing minimal intoxi- cation recovered 21 days after the one-time dose was administered (Kelly latipes muscular, and skeletal abnormalities, and those injected with 1 to 9 pg/egg (ppb) had reduced hatching success. Edmunds Satake horse-eye jack, MTX, and CTX analogs have been isolated from herbivorous fish (Yasumoto 1976; Chungue (Endean muscle fish carnivorous in concentrated found not is MTX LANDSBERGpreced- reflex righting of loss and disturbed, when activity swimming uncoordinated ing death. The estimated LD REVIEWS IN FISHERIES SCIENCE 1992). Pathological changes in fish fed fish in changes Pathological 1992). et al increases the efflux rate of Na

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 / ., ); M and et al Scarus ., 1985). ., (as et al et ., 1988), and 1988), ., ; and 3.1 n Glabrilutjanus Scomberomorus et al et Caranx latus (as ., 1994). One study Lycodontis javanicus Lycodontis (Vernoux et al (as ., 1991). However, MTX is CTX-1/kg) (Lewis, 1992). Chlorurus gibbus Chlorurus latus M Sphyraena jello . C . The highest concentration of et al CTX-1/kg fish) to cause human cause to fish) CTX-1/kg bloom in a coral reef microcosm, reef coral a in bloom M , which resulted in dramatic behavioral dramatic in resulted which , . These levels are lower than the level toxicus . G 221 toxicus Symphorus nematophorus . Seriola dumerili G Gymnothorax javanicus Gymnothorax Variola louti ., 1966). By definition, ciguateric fish can accumulate /kg in the heavybeak parrotfish, heavybeak the in /kg ., 1992). Abnormal neurological behavior was reported in et al M /kg in the pickhandle barracuda, M et al /kg in the narrow-barred Spanish mackerel, and the yellowtail M ); 1.5 n 1.5 ); ; 1.0 n /kg in the chinaman fish, /kg in the giant moray eel, moray giant the in /kg ); 1.5 n M M ., 1977). Fish can retain CTX for long periods of time — ciguatoxic fish Thus far, there is minimal but compelling evidence that CTX can affect fish At least 400 fish species have been implicated in ciguatera (Halstead, 1967). bartholomaei . Because CTX can be highly toxic and lethal to fish, Lewis (1992) suggested that there that suggested (1992) Lewis fish, to lethal and toxic highly be can CTX Because may be an upper limit to the CTX levels fish can have before the toxins kill them, and that this factor associated may with ciguatera. Given contribute the generally infrequent occurrence to of ciguateric the fish low and the fact that those found were incidence no more than moderately toxic, it of was not human considered fatalities surprising that fish kills attributed to reported (Lewis, the 1992). ciguatoxins have not been mortalities. fish for responsible be also may circumstances, certain under and, health There is little information regarding the effects of other ciguatera although it has been toxins demonstrated recently that on MTX is lethal fish, to fish (Terao gibbus commersoni kg in the yellow-edged lyretail, of CTX-1 shown to be lethal to mosquitofish (4.7 n apparently apparently has no direct role in ciguatera (Holmes (Yokoyama known toxins nonproteinaceous lethal most the of one fish are routinely exposed to this toxin. Aspects of the mechanisms of toxin transfer, of are carnivorous modification in toxin of of accumulation of the and chain, food precursors the through CTXs to still unclear. The fate and deposition of the various ciguatoxin derivatives in sequestered, excreted, defined. How the toxins are accumulated, tissues is not well fish or chemically biotransformed by fish are toxins the of many and incomplete, is is toxins ciguatera the of structure chemical currently unknown. Knowledge of the present in only very low concentrations in fish tissue (Anderson and Lobel, 1987), which makes purification and identification of these toxins difficult and expensive. Also, in many cases only of CTX concentrations heavy consumed is tested for in tissues. In acanthurids that had demonstrated that CTX was present in all tissues (liver, ovary, skin, muscle, gills, bones, and viscera [other than liver and ovary]) of the jacks (Yasumoto reported been has parrotfish a of liver the in g/tissue per MU 1000 to Up et al maintained in nontoxic water and fed a nontoxic diet maintained toxicity for up to 30 months (Banner 1.9 n nematophorus Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS changes that indicated a toxic effect, CTX could not be detected in muscle or liver tissues (Magnelia a to exposed were that fish herbivorous but CTXs were not detected in the muscle tissue (Goodlett n 0.1 > at (estimated CTX of levels high sufficiently intoxication (Lewis, 1992). Selected reports of the levels of CTX extracted from the CTX that indicate 1992) Lewis, in references (see fish ciguateric toxic highly of flesh (assuming all toxicity stems from CTX-1) can be present at levels of up to at least n 1.3 C 3.6-kg a of liver the in g/tissue per MU 14 was toxin

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., The

et al ), chloride M ., 1990). , MTX was 2000 was MTX , 2+ et al and a higher pH ., 1983), which can 2+ Ca M et al in mitochondria or intracel- 2+ ., 1992), could high doses of MTX of doses high could 1992), ., entry into the cells triggers a series a triggers cells the into entry . (1988) noted degeneration of the ., 1996). The mode of action of MTX of action of mode The 1996). ., 2+ , exposed to CTX via intraperitoneal via CTX to exposed , et al et ., 1991). The severity of cellular damage cellular of severity The 1991). ., ., 1999). Such experimental information et al et al et et al et 222 et al ., 1987) or DTX-1 (Terao ., 1996). In both hemolysis and ichthyotoxicity and hemolysis both In 1996). ., entry into the cells activates calmodulin, which calmodulin, activates cells the into entry 2+ et al et al et nitida Chromis strains be sufficient to cause pathological changes in changes pathological cause to sufficient be strains ions into the cells (Takahashi of ciguatera-associated toxins (see also page 289). 2+

to exert its activity. Ca activity. its exert to 2+ toxicus , was exposed to water containing MTX (0.58 n . G ., 1999). When the freshwater fish white cloud mountain minnow, mountain cloud white fish freshwater the When 1999). ., et al et ., 1996). At a pH of 8.0 and a concentration of 2 m 2 of concentration a and 8.0 of pH a At 1996). ., et al et The potency or activity of lipid-soluble toxins, such as CTX, that accumulate in Recently, Landsberg (1995) discussed the possibility that CTX or other benthic lamina propria, swelling of the villi tips, and an absence of the brush border in the chromis, reef barrier of intestine injection. These pathological changes in the intestines of reef fish may with associated be mice likened in symptoms diarrhetic the and pathology intestinal the both to exposure to either CTX (Coombe then lead to cell apoptosis (Terao apoptosis cell to lead then Ca required MTX assays, were 1994) Sylva, (de Florida in chain food ciguatera the in implicated fish of species 1994 to 1993 and 1980 the during affected were that species fish reef those to similar mortalities in the same area. The behaviors and signs demonstrated by fish experi- fish tropical diseased of those to similar were above) (see CTX to exposed mentally during the 1993 to 1994 event. Pathological changes in diseased fish included fluid accumulation and sloughing and detachment of the intestinal epithelium down to the basal lamina (Landsberg, 1995). Capra carnivorous fish, may be expected to differ from the potency or activity of water- soluble toxins, such as MTX and SCTX, that are concentrated in herbivorous (Magnelia fish. excreted routinely is MTX Although the intestine before these toxins can be excreted to fish of responses of range from the Theoretically, disease. to susceptible more fish? become If so, the fish could different benthic dinoflagellate strains (presumably due to the acute, between difference effects the explain could potencies) of various of components different toxic toxic mortalities and the mortalities caused by chronic toxicity that leads to patho- logical effects and subsequent invasion by 1995) opportunistic (see also page pathogens 289). (Landsberg, or particularly toxic particularly or cells in the epithelium organelles cell the and cells, chloride the of of cytoplasm the in appeared vacuolations the gills were affected showed lamellae gill the of capillaries the in after cells Blood destroyed. eventually were 10 to 30 (Terao exposure h min. 4 after apoptosis marked Marked Ca hemolysis. to lead that events of in turn promotes phospholipase A2 activity, which finally leads to the destruction of the cell membrane through the hydrolysis of membrane lipids (Igarashi lular vacuoles can result in a decrease (Hassan death in cell and ATP cytolysis causing and other nucleotides, eventually in fish gills was proportional to increased concentrations of Ca (Terao times more toxic than PbTx-3 (Igarashi needs to be extended to field situations in order to evaluate potential mechanisms by which fish may be affected by MTX. microalgal toxins could be involved in tropical reef fish kills. She postulated that a due immunosuppression by triggered was event mortality mass and disease fish reef to the consumption by fish LANDSBERG Igarashi 1996; REVIEWS IN FISHERIES SCIENCE is an increased influx of Ca Tanichthys Tanichthys albonubes 1999). The entry and subsequent accumulation of Ca

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ) ., ., et al et al Oryzias grindleyi . ) (Table 1) G ., 1997b). In Patinopecten ., 1997). When et al et al Gonyaulax polyedra Gonyaulax ., 1999), its biogenetic its 1999), ., have been implicated been have (= ., 1998b). Mussel extracts Mussel 1998b). ., et al et ., 1997b; Ciminiello ., 2001). Orally administered Orally 2001). ., et al et et al , 1990; Satake et al et et al. Protoceratium reticulatum (= ., 1997; Draisci 1997; ., ., 1997). 223 ., 1988), Chile, New Zealand (Yasumoto and (Yasumoto Zealand New Chile, 1988), ., et al et et al in (Daiguji Norway in Lingulodinium polyedrum Lingulodinium et al et Lingulodinium polyedrum Lingulodinium ., 1999). The biological effects of YTX are different are YTX of effects biological The 1999). ., and ., 1998). YTX and homoYTX were confirmed recently edulis et al et . M et al , in the Adriatic Sea (Satake ., 1998; Yasumoto and Satake, 1998), and 1-desulfoyessotoxin and 1998), Satake, and Yasumoto 1998; ., Gonyaulax grindleyi et al et ., 1996b; Yasumoto and Tazikawa, 1997). Because YTX occurred et al ., 1999). ., 1987), Norway (Lee Norway 1987), ., ., 1997a; Draisci 1997a; ., , in Mutsu Bay, Japan, where DSP incidents were frequent (Murata Gonyaulax grindleyi Gonyaulax et al et al et et al et studies with human lymphocytes, YTX appears to interact with calcium , exposed to 0.5 and 1.0 ppm YTX died during a 24-h exposure, but three fish three but exposure, 24-h a during died YTX ppm 1.0 and 0.5 to exposed , Both The potential effects of benthic dinoflagellate toxins on fish need to be critically be to need fish on toxins dinoflagellate benthic of effects potential The Other analogs of YTX — homoyessotoxin (homoYTX), 45-hydroxyhomoYTX, in marine animal mortalities (Table 2), but little is known about their possible toxic in marine animal mortalities (Table 2), but little is known about their possible mechanisms and the potential role of YTX and its analogs on the health of aquatic organisms. One incident in 1966 suggests that under certain situations can be lethal to marine fauna. During December 1966, in South Africa, a red tide (Draisci contaminated by homoYTX induce neurological symptoms in mice similar to those provoked by YTX. No evidence of diarrheogenicity of homoYTX was obtained at the doses tested (Tubaro dinoflagellate the in present be to was isolated from mussels mussels from isolated was 1998, 2000; Tubaro Tubaro 2000; 1998, and carboxyhomoyessotoxin — were isolated recently from Mediterranean mussels, Mediterranean from recently isolated were — carboxyhomoyessotoxin and Mytilus galloprovincialis from those of DTX and OA. YTX did not cause intestinal fluid accumulation in mice, in accumulation fluid intestinal cause not did YTX OA. and DTX of those from whereas OA and DTX-1 (associated with DSP) did (Ogino evaluated evaluated through experimental and field studies. currently used to test for It known ciguateric toxins are may not able to detect the be broad that the or to evaluate the potential that may affect or accumulate in fish spectrum of toxins methods web, food the of predators top the on CTX of effect The toxins. these of interactions that is, humans, has been studied, but the potential needs effect to be on examined. aquatic organisms I. YESSOTOXINS45,46,47- and (45-OHYTX) 45-hydroxyyessotoxin analogs, two and (YTX) Yessotoxin trinoryessotoxin (norYTX), were first isolated from Japanese scallops, yessoensis 1987; Satake together with DTX-1 and DTX-3, Japan in it shellfish from reported been has it Although 1998). Satake, and (Yasumoto was tentatively included in the DSP (Murata family (Ciminiello Italy and 1997), Tazikawa, isolated was toxin, polyether disulfated a YTX, recently. only discovered was origin from the dinoflagellate (Satake administered i.p., YTX shows a potent mouse lethality, but the toxicologic effects (Terao on human health are virtually unknown latipes exposed to bisdesulphated YTX died after 6 h. Ichthyotoxicity due to YTX was not considered to be significant (Ogino Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS YTX was not lethal to mice at 1.0 mg/kg, and did not appear fish, rice Japanese to the of None 2A. cause phosphatase protein inhibit to or hemolytic effects in vitro Rosa la (de maitotoxin to similar way a in channels

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , sp., blooms Calanus very low very was the

, polyedrum Mytilus . L , this species was grindleyi polyedrum . polyedrum . . G L blooms in Los Ange- L , were washed ashore. cells on the gills but no cells on the gills but grindleyi Chloromytilus meridionalis Chloromytilus . G suppressed feeding in ., 1975). cells/ml, and dissolved oxygen 3 was unknown. Gonyaulax et al 10 × for a prolonged period were starving. In starving. were period prolonged a for grindleyi grindleyi . . 224 G G , larvae were exposed to exposed were larvae , Chorisochismus dentex bloom in 1975. In 1975, Lingulodinium polyedrum , and resulted in gastrointestinal illness in eight grindleyi . G extended through St. Helena Bay to about 50 miles Bay to about 50 through St. Helena extended have been recorded along the southern California, USA, California, southern the along recorded been have polyedrum . L Mytilus edulis , a large number of black mussels, black of number large a , Engraulis mordax Engraulis polyedrum . L ., 1986; Sykes and Huntley, 1987). When these original studies were Gonyaulax grindleyi Gonyaulax et al Donax serra Donax . Copepods exposed to exposed Copepods . Both cells and cell-free filtrate of Blooms of Blooms Although no direct evidence was available, Huntley (1989) hypothesized that the Although no direct evidence was available, J. AZASPIRACID consump- the by caused was Netherlands the in event poisoning shellfish a 1995, In tion of toxic mussels, and numerous species of invertebrates (Table involved, and only a few sucker fish, 2). Almost no fish species the affected area had many Dead mussels from were identifiable cells in the gut contents, although unidentified red-pigmented material revealed mussels black and white the both on out carried tests Toxicity present. was was occurred poisoning shellfish human of cases no That toxic. highly were they that and area the in food of source popular a not were mussels that fact the to attributed to warnings that were issued over the radio and in the press. Although at the time there was no information on toxin production by considered highly suspect (Grindley and Sapeika, 1969; Grindley and Nel, 1970). levels were above 5.0 ppm. There was no evidence of oxygen depletion associated 5.0 ppm. There was no evidence of levels were above with a decaying bloom. This bloom resulted in the death mussels, of thousands of white produced copepods production, egg of absence and rate mortality high the to addition lethargic after about 10 days. When very few fecal pellets and became conspicuously anchovy, Californian LANDSBERG caused by REVIEWS IN FISHERIES SCIENCE north. Maximum cell counts were 5.0 to 6.7 pacificus those than lower were that rates observed, were survival and growth, feeding, of rates in the control larvae that (Huntley were in seawater without conducted, dinoflagellate the production of food YTX by sources the and smothered, mussels the clogged, became gills The pilings. on settled had that animals fell to the bottom and decayed (Oguri coast for many years (Sweeney, 1975), but rarely have mass mortalities have been noted. One of the earliest reports species described (Table 2) the four days mortality after the of “red streak” numerous had reached marine the 1902). No shore evidence (Torrey, was found to suggest that the California were toxic (Torrey, 1902; Sweeney, 1975), but organisms were not tested for toxins. for tested not were organisms but 1975), Sweeney, 1902; (Torrey, toxic were Similar blooms have also been organisms associated in the Adriatic with Sea along the marine coast of Croatia (Table mortalities 2). of demersal 1962-1977, during year worst the was which 1976, in anchovy northern of recruitment followed a large-scale dominant food organism in the south California Bight, and Lasker (1981) concluded that its poor nutritional quality was to exposed primarily fish responsible that for confirmed the studies poor Experimental 1989). year-class (Huntley, of 1976 showed no growth (Huntley, 1989). mussels, of gills the to damage mechanical caused Harbor Beach les–Long

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tests et al , and ., 1998a, (Gribble, mikimotoi et al

Heterocapsa in vitro , ., 2000; James Chrysochromulina et al Gamberdiscus toxicus Gamberdiscus ., 1995). Several other Amphidinium carterae Ostreopsis ovata , et al ., 1995a; Bodennec ), Gymnodinium Gymnodinium pulchellum (= et al Protoperidinium crassipes Protoperidinium tamarense Gambierdiscus toxicus . in part), , A , 225 ., 1999a, 1999b; Draisci sp.; the prymnesiophytes et al Karenia mikimotoi Karenia monilatum , . ., 1999a). A = Gyrodinium aureolum ( , Nitzschia et al Karenia brevis Karenia ., 1998a). Cochlodinium polykrikoides , , ., 1990). Most of the hemolysins produced by harmful microalgae are microalgae harmful by produced hemolysins the of Most 1990). ., et al ; the diatom ., 1998b, 1998c; Ofuji et al et et al Alexandrium catenella The bioorigin of azaspiracid is associated with associated is azaspiracid of bioorigin The When mice were injected with acetone-extracted hepatopancreas from toxic Several species of microalgae produce hemolysins that can adversely affect siamensis nagasakiense = Gyrodinium aureolum operculatum ., 2000). In October 1997, a second shellfish poisoning event in Arranmore, along Arranmore, in event poisoning shellfish second a 1997, October In 2000). ., . . . A circularisquama Gymnodinium aureolum al risk serious a as recognized Now people. 12 affected Ireland, of coast northwest the to human health the shellfish poisoning syndrome has poisoning or been AZP (Ofuji termed azaspiracid 2000; Satake health the on have may toxin this effects any, if what, stage this at unclear is It 2002). of aquatic organisms. G K. HEMOLYSINS by determined are compounds bioactive of properties hemolytic the Experimentally, their ability to lyse mammalian (or other animal) erythrocytes in people. Symptoms included nausea, vomiting, severe diarrhea, and stomach cramps stomach and diarrhea, severe vomiting, nausea, included Symptoms people. — all symptoms similar to DSP. The mussels had originated from Killary Harbor, Ireland (McMahon and Silke, 1996), where toxicity persisted in the shellfish for at least 8 months. The toxin was identified recently and characterized as azaspiracid (Satake mussels, the clinical symptoms in mice were clearly different from those caused by DSP. Mice experienced neurological symptoms, including respiratory distress, limb paralysis, and death within 20 min at higher doses. Unlike DSP, which only targets the digestive tract, pathological changes in mice necrosis lymphocyte and included liver, the in accumulation necrosis fat intestine, small the of in propria the lamina in the thymus, spleen, and Peyer’s patches of the small intestine (Ito (Yasumoto fatty acids (Table 12). Many species have numerous fatty acids occurring in different proportions a in each species, and not all specific fatty acid profile, with of the these fatty acids are necessarily hemolytic. Hemolytic fatty acids tend acid to octadecapentaenoic be as the (such lipids and acids fatty polyunsaturated longer-chain aquatic target of function membrane the altering by act to appear which [C:18:5n3]), more is and species algal in encountered rarely is acid Octadecapentaenoic species. potent than the fatty acids 20:5n3 or 22:6n3 (Arzul species also produce toxins that are hemolytic, but the toxins’ mode of action are by produced MTX example, For acids. fatty the from different causes lysis of cells due to the influx of calcium (see ciguatoxins). aquatic organisms. Species reported to produce hemolysins include the dinoflagel- lates O Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1995). Therefore, hemolytic activity, depends on the production of high concentra- tions of specific fatty acids, and like toxins, their production may vary according to environmental conditions and genetic factors (Bodennec

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; has not been not has has been associated been has , these toxins may have monilatum . A Chattonella marina 226 Alexandrium monilatum Alexandrium -associated fish kills in the last few years (K. ; the raphidophyte species produce PST, produce species Gymnodinium pulchellum sp. (Table 1). Numerous fish kills associated with and monilatum . brevis A

Alexandrium Synechococcus Prymnesium parvum Karenia and different actions — hemolytic include that toxins the multiple producing Species hemolytic. major are components toxic derivatives are provides section following The 1). Table (see sections neurotoxic, other in discussed are activity whereas other examples of the major species currently known to produce hemolysins. Although most toxic most Although 1. Dinoflagellates a. Alexandrium monilatum cyanobacterium blooms of these hemolysin-producing species have been documented world-wide, the in frequency in increasing been have to appear they species HAB other like and last 30 years (Tables 2 to 7). In some cases where species produce multiple toxins, for example, toxins. these produce to demonstrated polylepis with fish kills principally in the Gulf of Mexico and along the Atlantic coast of Florida of coast Atlantic the along and Mexico of Gulf the in principally kills fish with have there areas, these in ago years 50 to 20 frequently recorded Although 2). (Table been few reports of Steidinger, personal communication). LANDSBERG REVIEWS IN FISHERIES SCIENCE TABLE 12. Hemolytic lipids reported from microalgae

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ) , and may be klebsii . cells/ml, 7 A ., 1985). Rats 1985). ., toxins showed et al et ). At lower concen- lower At ). ., 1995). cells at 10 , the fish first showed first fish the , ., 1995, 1996, 1997). Amphidinium et al (reported as et al Crassostrea virginica monilatum . A have been shown experimen- monilatum and was eight times more potent more times eight was and monilatum . . ., 1985). The extracts contained a (Paul A A operculatum et al . A Brachidontes recurvus Brachidontes klebsii . monilatum . ., 1990). 227 A A (as et al ., 1983), but the hemolysin has still not been Aspergillus niger Aspergillus A. operculatum ., 1990). The compound hemolyzed mouse blood were exposed to exposed were et al

, and et al ., 1990). Seven bioactive compounds, amphidinols 2 to 2 amphidinols compounds, bioactive Seven 1990). ., ., 1980a,b; Erker spp. have been demonstrated to produce ichthyotoxic et al et m) and the difficulties of obtaining samples of them, it will et al µ Ischadium recurvum Ischadium ., 1990). The minimum concentrations of C18:4n3 and hemolysin-1 ., 1990). Ichthyotoxicity of hemolysin-2 has not been tested. In fish cephalus Mugil et al were estimated to be 1.9, 0.8, and 0.24 SU/mg, respectively, where one et al Amphidinium ., 1967; Clemons Hemolytic activities of a fatty acid (C18:4n3), hemolysin-1, and hemolysin-2 of Whole cells and crude extracts of An active compound isolated from The hemolytic compound identified from this species was thought to be a carterae . cells at 84 ng/ml, indicating that it was 120 (Nagai (Merck) saponin times more potent than commerical hooked mussels, hooked trations, byssus production was inhibited in bent mussels, and oysters When 1969). closed (Sievers, effects extracts’ the to the resistant were Crustaceans valves. shell mullet, striped horizon- or down upside turning equilibrium, of loss activity, frenzied then distress, 1960). Wilson and (Gates death to prior just activity of burst violent final a and tally, involved in unexplained fish kills. A SU (saponin unit) stands for the hemolytic potency of (Yasumoto 1 mg of Merck’s saponin required to kill killifish were 10 and 7 to 18 ppm, respectively (Watanabe, 1986, in Yasumoto killed by hemolysin-1, vigorous congestion after min 50 in to 24 in the occurred death and gills off, peel to occurred began scales the first, open, the opercula stayed convulsive movements (Yasumoto fungus the of growth the inhibited 8, were characterized recently from b. Amphidinium carterae Although than amphotericin B (Nagai there were significant mortalities of eastern oysters, hemolysins (Table 1), reports of their Because effects of their in benthic habitat natural and the environments lack of are not are species these with human associated events mortality many that possible is it areas, rare. presence in remote reef reported until some time after the events have occurred. Additionally, because of their small size (< 20 require a concerted effort to investigate the extent to which Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS tally to be lethal to mice, rats, fish, and cockroaches (Gates and Wilson, 1960; Aldrich 1960; Wilson, and (Gates cockroaches and fish, rats, mice, to lethal be to tally et al water-soluble glycosidic substance constric- abdominal sedation, that produced it and mice, was adult young to i.p. not administered STX or GTX. The extract tion, fecal clumping in the perianal area, ataxia, was tremors, cyanosis, loss of reflexes, convulsions, and death. Gross and microscopic pathology included an acute active hyperemia of the viscera, multifocal areas of necrosis intestinal wall of and diaphragm, and the presence of cytoplasmic vacuoles in cells in the musculature of the (Erker pancreas the of portion acinar the of margins peripheral the exposed to extracts from mussels contaminated by increased disaccharidase activity in the intestine (Taboada nonaromatic glycolipid (Bass characterized. When experimentally exposed to

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . . ., ., et et et et A G , or , et al et et al [= niger .

A Karenia mikimotoi in North had closer . . (2000), the (2000), . Gyrodinium Gyrodinium K ., 1991; Blasco Gymnodinium described from described et al et nagasakiense ., 1989; Taylor ., 1995; Hansen and et al cf. aureolum

. et al

et al complex to be harmful be to complex G . (2000) (= (2000) . ATPase) bound therein bound ATPase) + , has also been reported /K et al et + aureolum . (Taylor G mikimotoi than with the North American (Steidinger ). To further confuse matters, a mikimotoi . . K Gymnodinium ., 2000), suggesting that the species / ., 1996). Amphidinols bind to sterols K ., 1988; Nagasaki nagasakiense Gymnodinium aureolum . Hansen and Moestrup in Daugbjerg G mikimotoi (Hulburt, 1957), is morphologically and Gymnodinium aureolum Gymnodinium et al et al , et al identical to the North American strain has strain American North the to identical . Miyaki and Kominami ex Oda 228 K Hansen in Hansen in Hansen aureolum Karenia mikimotoi . G . 2000) and mikimotoi aureolum and will be considered as a separate species from the

Karenia mikimotoi mikimotoi et al cf.

Karenia mikimotoi Gyrodinium aureolum ., 2000; Hansen sp. and cause hemolysis of human erythrocytes (Paul ., 1995). Reports of Karenia mikimotoi (Iizuka and Irie, 1969; Partensky and Sournia, 1986; Partensky 1986; Sournia, and Partensky 1969; Irie, and (Iizuka

. (1991). Amphidinols are active against the fungus et al aureolum et al . Gyrodinium aureolum et al G Nitzschia Gymnodinium type ‘65 type Gymnodinium Gymnodinium aureolum Gymnodinium (Partensky, 1988; Partensky Takayama and Adachi]) complex have been variously confused or , and because of their potent antidiatom activity, it was suggested that (= ., 1997). Amphidinols were detected in the culture medium of

(Taylor Hulburt) and as conspecific with et al ., 1988; Takayama and Adachi 1984; Takayama and Matsuoka, 1991). Some ., 1996; Gentien, 1998), whereas others treated the European At the current state of knowledge, the knowledge, of state current the At aureolum aureolum, Gymnodinium aureolum, . (2000) ., 1995; Daugbjerg ., 1996, 1997). The mode of action of amphidinols mimics that of the antimycotic ., 2000) (described as . operculatum (Paul microbes benthic other against compounds allelopathic as function they aureolum al nagasakiense synonymized, so some taxa may have been Because of reported uncertain identifications, in under addition to an incorrect name. to attributed been have 2) (Table mortalities invertebrate and fish some cf. al Species in the in Species c. Gymnodinium aureolum and the diatom (Paul 1997). et al aureolum responsible for fish kills in northern mikimotoi Europe, Australasia, and Japan is America have indicated that this species is toxic to marine animals (Mahoney to marine animals will be referenced together and, unless otherwise stated, will be Hansen Following description. original their by 2 Table in denoted North American LANDSBERG Amphidinol 1 was originally reported to be activity by an Satake antifungal agent with hemolytic REVIEWS IN FISHERIES SCIENCE al channel in results membrane cell the at complex a of formation The B. amphotericin formation, increased ionic exchange, and increased permeability of the cell mem- brane, which can lead to hemolysis (Paul affinities with the Pacific G 1990; Smolowitz and Shumway, 1996) (Table 2). North America as genetically different from the European al of this in North America (K. Steidinger, personal communication). For the purposes the in reported species discussion, in plasma membranes of target cells, ergosterol in fungi, and cholesterol in animal cells, thus disturbing the arrangement of the lipid bilayer structure by nullifying its Na as (such enzymes deactivating and/or function barrier of strain Danish also been described (Hansen et al researchers recognized that the European Gymnodinium

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., ., ., ), et et et al et al et al et al ., 1979) ., ) (Braarud ) sp. (Arzul et al et — and are Gyrodinium that demon- Gyrodinium

) nagasakiense sp. contained (as (as . The composition (Romdhane , and reduction of cf. was demonstrated ., 1990).

) in Japan (Iizuka and aureolum

contained 7.6% MGDG et al Gymnodinium mikimotoi ., 1998). Hemolysis of red ., 1995a). The exact mode . mikimotoi K aureolum mikimotoi .

. Gymnodinium K et al et al K have been identified as exotox- as identified been have ., 1995a). The fatty acid C18:5n3 acid fatty The 1995a). ., Gyrodinium aureolum Gyrodinium Gymnodinium was determined for several differ- several for determined was nagasakiense ., 1998). Although the fatty acid Gyrodinium et al et (Mahoney Gyrodinium aureolum Chaetoceros gracile et al cf. (as Karenia mikimotoi Gyrodinium mikimotoi

Karenia mikimotoi 229 . (reported as (reported ., 1990; Arzul 1990; ., mikimotoi Karenia Gyrodinium ., 1992). However, field measurements of dissolved . et al et K mikimotoi . et al ., 1990), C18:5n3 (Parrish mikimotoi ., 1982). K . K et al et al Gymnodinium aureolum Photobacterium phosphoreum Photobacterium bloom in 1984 amounted to about 4.4 billion yen. Some human to about 4.4 billion yen. Some bloom in 1984 amounted . (1984) classified PUFAs extracted from the toxic hepatopancreas toxic the from extracted PUFAs classified (1984) . (reported as ., 1989) that fish and invertebrate mortalities were possibly due to ., 1989), but only in small quantities that were not considered to be et al et et al et al ) (Yasumoto ) were shown to be hemolytic (Table 12) and ichthyotoxic (Yasumoto mikimotoi

were C20:5n3, C20:4n6, and C22:6n3 (Arzul mikimotoi . ., 1995b), which was also reported as reported also was which 1995b), ., A digalactosyl monoacylglycerol (MGDG) and a polyunsaturated fatty acid Since the mid-1960s, when blooms of Hemolysins associated with associated Hemolysins K ., 1990). The origin of both of these hemolysins could have been the more ., 1998). When the percent lipid compositions of the two species and their of action of PUFA in the inhibition of diatom growth is still unknown (Arzul Takagi 1995a). of contaminated scallops according to their toxicity to mice (cited in Arzul strated hemolytic activity against mouse blood cells and inhibited growth of of growth inhibited and cells blood mouse against activity hemolytic strated gracile 1995a). In October 1994, in the Gulf of Gabè, Tunisia, a et al blood cells, growth inhibition of the diatom aureolum in bioluminescence (Yasumoto C18:5n3 including made up 25.1% of the total fatty acids isolated from of additional fatty acids from ent PUFAs produced by produced PUFAs ent components of both the hemolytic MGDG and DGDG were proportionally different proportionally were DGDG and MGDG hemolytic the both of components in each species, the dominant fatty acid was, as in aureolum al common type of glycolipid, a glycosyl diacylglycerol. Hydrolysis glycolipid would yield a free of fatty acid and a glycosyl monoacylglycerol (Parrish this type of al hemolytic components were evaluated, and 8.9% digalactosyl diacylglycerol (DGDG), and 14.2% MGDG and 20.8% DGDG (Parrish or in Scotland (Jones from isolated [C18:5n3]) acid (octadecapentaenoic (PUFA) (Partensky other environmental stressors, such as depletion of dissolved oxygen (Helm 1974; Tangen,1977; Heinig (Ottway Ireland southern in case the be to this indicate not did oxygen high enough to be lethal. At the organism level, no ichthyotoxin was isolated and preliminary fish bioassays were negative (Tangen, 1977). There were suggestions (Partensky species previously referred to as the European referred to in the text as as (described waters Norwegian in and 1966) Irie, mortalities invertebrate and fish multispecies problems, became 1970) Heimdal, and have been reported from Europe, Australasia, Japan, Africa (Table South 2). Significant economic America, losses to fish and farms have occurred North (Tangen, 1977; Dahl and Tangen, 1993; Honjo, 1994); for Karenia example, damage costs from a and lung congestion, have as nausea, sore throat, eye irritation, health effects, such been associated with Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS cultures from cytotoxin fat-soluble a of presence The 1990). Arzul, and (Gentien ins of

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 2 4 ) et 10 × ) also cells/ 5 ., 1979; ., (Tables Mytilus 10 , presum- aureolum mikimotoi . aureolum × . Only after . ehrenbergii

et al et diversicolor G . . K S ) to increased F ; softshell clams, were unable to ., 1995a), whose 1995a), .,

, mikimotoi nagasakiense were more resistant more were . mikimotoi in laboratory tests, , were not adversely not were , et al et ) to the development K . (Lesser and Shumway, K

?) cultures at concentra- cf. ) Gyrodinium ., 1995b). There is some Haliotis discus (as (Arzul et al aureolum ., 1998a, b). Several plankton . ., 1990; Nielsen and Strømgren, bloom. An offshore bloom of G at densities exceeeding 5 aureolum . Spisula solidissima Crassostrea virginica et al G nauplii were documented (Helm et al

, Favella ehrenbergii Gymnodinium Arenicola marina Arenicola mikimotoi (as Sulculus diversicolor Sulculus . , whereas mortalities of K cells/ml were mortalities of 5 230 aureolum mikimotoi Mercenaria mercenaria . Argopecten irradians

K Chaetoceros gracile Chaetoceros mikimotoi . (reported (reported as aureolum K . Artemia salina G died, whereas whereas died, ., 1998a). When Mediterranean mussels, cells were further concentrated to around 1.4 Gymnodinium et al mikimotoi mikimotoi cells/ml, two species of showed different responses: . , were exposed to K cells/ml showed a significant reduction in growth rate (Nielsen and 3 ; and northern quahogs Haliotis discus Haliotis was transported shoreward by southerly winds and tidal action. 10 all ×

., 1979). When bivalves were exposed to Effects on invertebrates, including plankton, have been documented both in both documented been have plankton, including invertebrates, on Effects In September 1988, a large mortality event occurred in Maquoit Bay, Maine, USA, Maine, Bay, Maquoit in occurred event mortality large a 1988, September In Blue mussels exposed to aureolum ., 1974). . noted (Matsuyama Karenia mikimotoi G Gymnodinium aureolum ml by onshore winds and weaker-than-normal tidal flushing. The high number of mortalities may have been due to the exposure by produced toxin and mucus of the to as well as concentrations oxygen the animals to low dissolved clearance of cells (Widdows 1991), which was probably due to reduced Strømgren, et al species-specific differences in pathological effects were documented. Pathological effects ranged from none (in Atlantic surfclams, of mantle and gill lesions (in eastern oysters, mortalities, decreased height of absorptive cells, and increased lumen diameter in the digestive gland (in bay scallops, 1993; Smolowitz and Shumway, 1996). When exposed to concentrations of 10 h, 24 after and demonstrated paralysis only. Avoidance behavior such as contraction tentacles of and escape the locomotion was also exhibited by galloprovincialis indication indication that al when an estimated 30 to 40% of softshell clams, blue mussels, and marine worms were exposed to a (Heinig and Campbell, 1992). Mya arenaria a 48-h exposure at concentrations of 10 LANDSBERG1998), was demonstrated to produce C18:5n3 (Arzul REVIEWS IN FISHERIES SCIENCE embryos and , produces (ROS) (Gentien, 1998). (Widdows 11) to 9 1, (Tables experimentally and 2) (Table field the Heinig and Campbell, 1992; Erard-Le Denn diatom the including 11), and 10 accounted for about 90% of the total biomass, the growth rate of Lugworms, 25%. than less by reduced was affected by exposure to tions of 9 1991; Dahl and Tangen, Smolowitz 1993; and Shumway, 1996; Lesser Matsuyama and Shumway, species have also been shown to be adversely affected by 1993; Hansen, 1995; growth was inhibited. Tintinnid ciliates, ably because of exposure to toxic exudates. However, of direct cultures attempts dense to of dem- filtrates using exudates, of effects toxic onstrate lack of toxic effects may have been due to failed. Hansen (1995) suggested that the rapid turnover of the toxin. When sustain growth when exposed to high concentrations of

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 4 5 2 at (as 10 10 10 × × × , that ), and ., 1983; et al aureolum ) bloom were mikimotoi . ) above 1 cf.

. K

G 3 cells/ml) may cause may cells/ml) 3 10 × aureolum 10 . Histopathological exami- ×

aureolum G . ). cells/ml. Behavioral effects Behavioral cells/ml. G 3 (as 10 (as Gymnodinium aureolum × by avoidance (Dahl and Tangen, ) (0.5 to 2 to (0.5 ) (as showed “stress rings” on the shells aureolum . G mikimotoi . mikimotoi K (as 231 . K aureolum aureolum . mikimotoi whereas animals injected with flesh extracts G ., 1990). . cf. K cells/ml, filtration rates were affected and growth 3 ., 1982). Pathological examination of affected salmon (as et al ) bloom in a salmonid fish farm in Scotland, distressed Scotland, in farm fish salmonid a in bloom ) mikimotoi densities exceeded 1 exceeded densities 10 . et al ., 1998b). Wild great scallops, × K et al Gyrodinium mikimotoi (Jones to 1.5 . aureolum 2 . K G K. mikimotoi (as G. aureolum) mikimotoi K. 10 × (as ) cells/ml, fish immediately increased their rates of opercular move- ., 1982). Concentrations of ., 1987); conditions similar to those observed in natural mortalities of Atlantic clearance clearance of cells was reduced noticeably. Mussels also retracted their

Salmo salar et al et al Young animals and larval stages of many fish and invertebrate species are also mikimotoi aureolum . . fish came to the water’s surface and circled around before turning over and sinking. and over turning before around circled and surface water’s the to came fish Death appeared to take place within a few hours. At the time of the fish kill, water in the ponds was turbid and viscous, with (Jones considerable foaming on the surface ment. At low concentrations, fish were hyperactive and attempted to leap out of the of out leap to attempted and hyperactive were fish concentrations, low At ment. container. large In secreted fish the the treatments, highest all In concentration, h. 1 the within dead fish or became comatose were inactive, and color, turned dark in quantities of mucus. Negative effects on the gills and digestive tract of salmonids were also associated with that could be used for dating growth inhibition periods. A bloom of up to 8 response pathological a swimming; surface increased and feeding reduced included was a mild sloughing of the gill epithelium K (Dahl and Tangen, 1993). During a cells/ml may cause acute mortality due to secondary severe sloughing gill in lamellae the and primary damage and to oxygen the uptake, chloride osmoregulation, cells. Damaged and gills blood affect suggested pH. that chronic Dahl exposure and (several weeks) Tangen of (1993) salmon of concentrations to also low to moderate liver necrosis. When experimentally exposed to up to 10.9 G cells/ml, mantle edges, showed intermittent shrinkage of their exhalent siphons, and were unable to completely close their valves during the 10-min experiment. Eighty-five percent mortalities (N = 20) were noted after 5 days’ exposure to 1.3 to 2.5 cells/ml (Matsuyama survived a bloom of 3.0 mm) scallops and growth mortalities of postlarval (0.25 to cells/ml caused mass after 1 month. When (5 to 30 mm) until the red tide vanished cessation in juveniles to exposed experimentally were scallops juvenile and postlarvae was inhibited (Erard-le-Denn nation of the gills of moribund fish sloughing of showed the epithelial tissues, swelling a and pyknosis of the severe primary lamellar epithe- and acute toxic necrosis, lium, lamellar hypertophy, and congestion of the branchial vessels (Roberts Turner salmon, showed that death was likely to have resulted from asphyxiation and osmotic shock as gut or gut of extracts Acidic intestine. and gills the to damage cellular extensive of result a contents from salmonids killed during a Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS susceptible to the presence of naturally occurring blooms may well influence their development and recruitment. stages and species numerous of individuals of number the in declines bloom, a After have been reported (Potts and Edwards, populations 1987). of Fish usually respond to 1993). dense In fish farms, adverse effects were observed in salmon, rainbow trout, and algal when cod lethal to mice 24 h after injection, densities of 5

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., (as et al et al Numer-

. Nassarius cells/ml and mikimotoi . K ., 1989; Underdal 1989; ., bloom caused the et al et polylepis , were exposed to ., 1999). Exposure to . C ), common periwinkles

3 ), dog ( et al branchial or intestinal Mg- intestinal or branchial Anomia ephippium 10 ) were completely eradicated × species to result in mortalities of in vitro in ., 1989; Lindahl and Dahl, 1990). Sea are currently known or suspected to be ., 1999; Sola Nucella lapillus et al 232 et al exposures. Postexposure examination of the of examination Postexposure exposures. Dicentrarchus labrax Chrysochromulina polylepis undatum , and saddle oysters, Apporhais pespelicani in vivo in Chrysochromulina (Underdal ) (Fossat , had retracted their tentacles and were losing mucus , were severely affected and to a large extent, gastropods. extent, large a to and affected severely were , Chrysochromulina ., 1989; Lindahl and Dahl, 1990). Effects on marine fauna mikimotoi et al Hiatella arctica ), waved whelks ( ., 1989). In other areas, dogwinkle mortalities were as high as 98 to cf. Raniceps raninus Metridium senile et al ), and pelican’s foot ( Echinus esculentus Echinus ., 1989; Lindahl and Dahl, 1990; Eikrem and Throndsen, 1993; Granéli . (1999) suggested that other observed morphological changes in the gills may In laboratory tests, sea bass, Littorina littorea (Table 4) ranged from extreme to slight. Numerous symptoms, including discolora- tion, changes in behavior patterns, and urchins, “loosening” of organs were noted. Sea first known toxic outbreak of a (Dahl Norway in invertebrates and tons) (800 fish of thousands et al a salinity of 30 ppt were killed within 4 h. damage There was that evidence would of have gill epithelial increased permeability and resulted failure in (Underdal osmoregulatory In some areas, the Atlantic dogwinkles ( 2. Prymnesiophytes a. Chrysochromulina spp. At least eight species of aquatic of mortalities in implicated been have species few a only but 1), (Table toxic organisms (Table 4). In early 1988, a ( octadecapentaenoic octadecapentaenoic acid (C18:5n3) and other PUFAs produced by Gymnodinium of inhibition in resulted acid octadecapentaenoic ATPase activity, but there was no demonstrable change in Na-ATPase or K-ATPase during fish of gills the in activity cells, chloride and mucus less were there morphologically, that determined gills fish and a reduction in the amount of mucus produced. The presence of fatty acids did not inhibit the production of the enzymes Na-ATPase or K-ATPase in the chloride cells, so enzyme activities related to ionic transport were not influenced, but Sola et al be sufficient to induce gill pathologies and ultimate death in fish. Salmon million. $4.5 at estimated were farms fish to losses Economic 4). (Table 1993) of 4 to 8 that were kept in water with concentrations reticulatus ous mortalites occurred among the fish groups Labridae and Gobiidae and in the tadpole fish LANDSBERG remained healthy. Ether extracts of flesh, liver, and gut from affected fish resulted in sublethal effects on mice within minutes of injection. Mice showed signs of mild problems (Jones hyperactivity, and respiratory paralysis, loss of coordination, REVIEWS IN FISHERIES SCIENCE species several HABs, with reported rarely and Unusually, 1990). Dahl, and (Lindahl red as such macroalgae, dying and dead bloom; the by affected were macroalgae of (Underdal 99%, which reduced dogwinkle distribution, local population sizes, and reproduc- tion levels (due to the high mortality rates of 1 and 2 year olds, 3 years of recruits were lost) (Robertson, 1991). Mortality rates were also Arctic rock high borer, among the bivalves anemones, 1982).

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , polylepis polylepis Ceratium . polylepis (Underdal . . C C C

3 cells/ml, no Oncorhynchus 4 ., 1992; Eikrem 10 × 10 ., 1988). During a et al × has also been docu- spp. has been difficult ., 1988), and copepods et al in many cases caused a et al ., 1995). Highest cell counts et al polylepis , and rainbow trout, rainbow and , . events have been from Scandinavia from been have events ., 1989). At the height of the bloom, the of height the At 1989). ., C , reduced the growth of blue mussels, cells/ml), and there was no evidence et al et 5 Chrysochromulina showed an increase in plasma osmolality 10 233 × Salmo salar Salmo was responsible for the death of almost 750 polylepis spp. in Denmark, an almost complete absence . densities reached 6 to 7 C cells/ml reduced growth by up to 60% when ., 1995). In bialgal cultures of 1 ., 1995). In bialgal cultures of 1 8 polylepis ., 1990). ., 1992). The bottom fauna was generally unaffected generally was fauna bottom The 1992). ., . , was common. Adverse affects such as frayed and Chrysochromulina has been reported from New Zealand (Chang, 1995), (Chang, Zealand New from reported been has ), blue mussels (Granmo Chrysochromulina leadbeateri et al C ., 1993; Edvardsen and Paasche, 1998). At minimum 10 × et al et al et ., 1991, cited in Hansen bloom demonstrated that toxins affected eggs and larvae and eggs affected toxins that demonstrated bloom polylepis (up to 6.14 leadbeateri

et al . cells/ml, Skeletonema costatum 4 C cells/ml (Heidal and Mohus, 1995). Mice injected with fish et al 4 polylepis 10 × parva 10 . . cf. . polylepis × Chrysochromulina C . C C ., 1995). , Atlantic 1995). ., changed color from red to orange to green, indicative of pigment ., 1990). The fertilization of gametes and successful development of Ciona intestinalis ., 1992). et al et Delesseria Delesseria sanguinea et al that were exposed to

et al Chrysochromulina , ., 1989). Laboratory experiments showed that in water containing In May-June 1991, Subsequent reports of toxic of reports Subsequent In laboratory experiments, toxicity of In laboratory experiments, ., 1984), and and 1984), ., sanguinea . and Throndsen, 1993; Heidal and Mohus, 1995), resulting in an economic loss that (Aune million $5 exceeded by the bloom, but in a few localities a high level of mortality of observed (Johannesen sea urchins was did not exceed 1 (Leivestad and Serigstad, 1988, cited in Nielsen, 1993). metric tons of salmon and trout in Norwegian fish farms (Aune mykiss for low dissolved oxygen or other potential environmental stressors. Although when et al D breakdown (Lindahl and Dahl, 1990). Empty thecae of the dinoflagellate (Dahl area bloom the inside noted also were inhibited the activity of planktonic bacteria, ciliates, structure and web copepods food (Tables pelagic 10 normal and the bloom, the of height the after weeks Two 11). was reestablished (Nielsen (Nielsen ascidians and mussels was completely inhibited (Granmo natural bloom of cells/ml and the diatom 96% decrease in the abundance (Myklestad of diatoms when compared with the control any muscle extracts from moribund fish exposed to the bloom did not demonstrate exposed been had that mussels blue from extracts with injected Mice effects. adverse of narrowing scratching, as such patterns behavioral deviating showed bloom the to eyes, and progressive limpness after 30 to 60 min, but they recovered subsequently (Aune or North (Estep Atlantic America North the (Table of areas other 4). in and 1993) (Hallegraeff, Australia in mented al et but thus far none of mortalities in these these areas. A species freshwater fish mortality event have in Denmark coincided been associated with a bloom with of aquatic animal discolored thalli were reported, particularly amongst benthic rhodophyceans benthic amongst particularly reported, were thalli discolored and concentrations of 1.1 of ascidians ( of ciliates was noted (Hansen , seaweed, potential grazers were present in the subsurface bloom, and bacterial production was extremely low. Field and laboratory experiments showed that to demonstrate (Granéli compared with controls (Nielsen and Strømgren, 1988 the during 1990). Experiments conducted Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS concentrations of 5

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , . 4 ., et et et 10 (see et al × et al cells/ in vitro cells and polylepis spinifera . (Stabell . C leadbeateri (Meldahl

C . , C polylepis . ., 1989; Meldahl C

polylepis Chrysochromulina Chrysochromulina hirta . 4 polylepis . C range wide a affects . et al C cells/ml. Although the Although cells/ml. C 10 4 × bloom in Norway, were Norway, in bloom ., 1993). Yasumoto 10 × Gyrodinium aureolum et al polylepis . polylepis C . ., 1995). C species may be toxic. Further studies Further toxic. be may species et al 234 ., 1989; Stabell ., 1990). In addition to hemolysins, other toxin acts at the cellular level, interfering lysed rat hepatocytes (Aune, 1989; Underdal 1989; (Aune, hepatocytes rat lysed has also been shown to inhibit the have not yet been characterized (Meldahl et al et al l saponin solution, then the hemolytic potency of ., 1989). Mussel extracts exposed to exposed extracts Mussel 1989). ., µ , and maximum total cell concentrations were 5 et al et Chrysochromulina leadbeateri . leadbeateri . ., 1992). At concentrations of 1 to 2 to 1 of concentrations At 1992). ., C , was associated with a mixed bloom of C ., 1994). In April-May 1992, a mortality of caged rainbow trout, rainbow caged of mortality a 1992, April-May In 1994). ., brevifilum et al et , naturally exposed to the 1988 the to exposed naturally , . (Underdal and et al et C , and Chrysochromulina polylepis polylepis polylepis polylepis Chrysochromulina polylepis . . C C Ostrea g saponin was used to define one hemolytic unit [HU]). The results indicated activity, nonspecific this of Because 1993). ., ., 1989; Aune 1989; ., The Intraperitoneal injections of hepatopancreas extracts of blue mussels and oys- ericina µ ., 1993). ., 1994). As well as demonstrating hemolytic activity, crude extracts of toxins .,1993). . C 1992). vesicles synaptic and synaptosomes into neurotransmitters of uptake al with cell membrane functions and ionic balance (Underdal with cell membrane functions and spp. in the Lillebaelt, Denmark. Dominant species included (1990) extracted hemolytic compounds from water containing bloom. As with from blue mussels exposed to the 1988 page 229), the hemolytic compounds Extracts from mussels exposed to a octadecapentaenoic acid (C18:5n3). were identified as MGDG and bloom had a a higher hemolytic activity (80 PUFA, HU/g) than did those from uncontami- nated mussels (16 HU/g). (A 0.1% saline solution of commerical saponin was used as the standard against a hemolysis took 0.4% place with a 10- mouse red blood cell 10 suspension. If H2LC as complete characterized later substance, hemolytic a accumulated had mussels the that (digalactosylglycerol) (Yasumoto uncharacterized toxins are also suspected to be present in al al et of aquatic organisms from protists to fish (Edvardsen and Paasche, 1998). The toxic compounds in al from doubled ml, most of the rat hepatocytes were totally destroyed. Cell size almost and the cell membranes lysed completely. The effects of prymnesin at concen- trations of 500 HU/ml on rat hepatocytes were similar to the effects of extracts from resulted leakage Enzyme hepatocytes. rat to toxicity dose-dependent a displayed from complete destruction of cell membranes and lysis of cells. Extracts from salmon livers and stomach contents also affected rat hepatocytes (Aune species (Hansen species Oncorhynchus mykiss cells/ml. Fish swam apathetically near the water’s surface prior to death. The total loss of fish was about 50 tons. Fish died principally during the early phase of 3 the to 1 between ranged concentrations cell when bloom, bloom was not toxic to the plankton community, the effect on the fish suggested these of several or one that lethal to mice. Toxicity of the bivalves to mice decreased within 20 days after the bloom had disappeared (Underdal al et LANDSBERGbe to proven if and, suspect highly is species this conducted, not were tests toxicity toxic, would represent the first report of toxicity in a freshwater REVIEWS IN FISHERIES SCIENCE on these four species are required (Hansen ters,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 and are , was cells/ml 8 parvum . calathiferum . P P patelliferum . sp. after 170 min , P calathiferum . P blooms occur in low- and ., 1995). Ichthyotoxicity 1995). ., cell densities exceed 5 to spp. blooms have been et al et Gambusia (approximately 10 Prymnesium parvum . P has been implicated in fish kills in Prymnesium , Prymnesium Prymnesium , now commonly reported in Scandinavia in reported commonly now , are confirmed toxic by the fish bioassay 235 saltans calathiferum are currently recognized as being toxic to .

(Table 4), it is possible that some of these P parvum calathiferum . . parvum patelliferum have taken place yearly during July-August, and have and July-August, during yearly place taken have P P . . P P Prymnesium Prymnesium Prymnesium was implicated as the causative organism of fish kills in . can become dominant within 3 to 5 days (Sarig, 1971). patelliferum , have helped fish farmers take preventative measures and avoid saltans parvum was highly suspected to be a toxic species. In an experimental spp. have been responsible for significant economic losses caused losses economic significant for responsible been have spp. . ., 1993) but not recognized as a separate taxon until 1982 (Edvardsen 1982 until taxon separate a as recognized not but 1993) ., . P . are identical (Larsen, 1999). For the time being mortality reports will reports mortality being time the For 1999). (Larsen, identical are P P parvum . P et al et and cells/ml. Since 1989, in the Ryfylke fjords, Norway, mixed blooms of 5 10 The biological spectrum of activity of prymnesins is extremely broad (Table 1) Although Prymnesium × parvum calathiferum patelliferum . . . exposure (Chang, 1985). (Shilo, 1981), but because of the difficulties of purifying (Igarashi the entity molecular toxin, single a it to due was is this unclear whether P centrifuged at 3000 rpm) was lethal to mosquitofish, b. Prymnesium spp. At least four species of aquatic organisms (Table 1). Ichthyotoxic documented since the late 1800s, mostly from Europe, brackish-water Ukraine, systems North America, in China, earlier Israel, the of many and Although species. particular identified Australia wrongly have may reports (Table 4). Some of these blooms were attributed to caused extensive damage to farmed fish (Edvardsen and summer Paasche, of 1998). 1983, In an the unusual fish and coincided shellfish with mortality the event presence in of New a Zealand newly (Chang, reported 1985; Chang and Ryan, dissolved 1985). oxygen At levels the in height the of area P the of bloom, the minimum fish kill were bioassay, 3.8 a to cell-free 6.3 supernatant ppm, of and Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS to due were events harmful (see Larsen (see and Paasche, 1998). However, the possibility that possibility the However, 1998). Paasche, and Germany (Kell and Noack, 1991), Moestrup (1994) noted that the kills were more likely due to actually one species is differences at still the light microscopic level, it appears that genetically being discussed. Apart P from slight morphological be provided as described in the original reports, but readers should be aware that the taxonomy of this genus is still in flux. China (Wang and Wang, 1992). Recently, associated with fish mortalities a in New Zealand and is the first report new of this genus species, from marine waters (Chang, 1985; Chang and Ryan, 1985). by fish kills in aquaculture facilities and fish farms around the world (Sarig, 1971; Moestrup, 1994). Except for salinity (usually between 1 to 12 ppt), brackish waters in ponds, shallow lakes, or lagoons (Edvardsen and Paasche, 1998). Under suitable conditions in fish ponds, blooms of since problem significant a been have Israel in ponds fish brackish in kills Recurrent using suspect pond water and mosquitofish, the early 1940s. Routine fish bioassays Gambusia affinis unnecessary economic losses. Ponds are chemically treated when low with concentrations copper of sulfate (Sarig, 1971). Fish kills usually occur when 10

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and , ., 1995, ., 1995). ., 1982). was kept Rana et al patelliferum Artemia et al . et al P , were affected at a concentration a at . However, it was has been shown to , and this effect may effect this and , + -glutamate and GABA L Acartia clausi were toxic to , and K and , + patelliferum . , there was no mortality of late patelliferum P , Na , . . It was concluded that toxic 2+ P toxins; their tails curve, they become they curve, tails their toxins; Tanichthys albonubes Rhodomonas baltica patelliferum in situ . P 236 patelliferum was offered singly as food and when it was . parvum . P P contained fatty acids, amino acids, phosphate, and ., 1982) either as a mixture of two galactoglycolipids two of mixture a as either 1982) ., ., 1994, 1995). The toxic extract of et al et et al ., 1996). ., 1994). An extract of parvum parvum . . Prymnesium P P et al . (1995) purified two toxins, prymnesin-1 and prymnesin-2, et al et al ., 1982), or as proteolipids (Shilo, 1981). Hemolysin-1, identified as a on other animal groups. When the copepod (see references in Shilo, 1981). Gill-breathing animals, such as fish, cells/ml caused a rapid inhibition of glutamate accumulation in the 4 et al 10 × tadpoles are highly sensitive to sensitive highly are tadpoles There have been relatively few reports or studies on the potential impact of Purified toxin from -aminobutyric acid), and (2) it enhances the calcium-dependent release of acetyl- γ suggested that laboratory studies may significantly underestimate the adverse effects adverse the underestimate significantly may studies laboratory that suggested of potentially toxic prymnesiophytes copepodids or adults. Strong negative effects on egestion and reproduction were documented, both when fed in conjunction with the ( of extract An 1995). Fonnum, and (Meldahl choline of 5 Ca to synaptosomes of permeability the increases be responsible for the plasma membrane depolarization neurotransmitter and processes (Meldahl the and Fonnum, 1995). disturbance of Prymnesium in dense cultures of a toxic strain of synaptosomes. The effect on this transport system is probably due to impairment of impairment to due probably is system transport this on effect The synaptosomes. one or more gradients (sodium, potassium, and calcium) and depolarization of the plasma membrane (Meldahl LANDSBERGis enhanced in the presence of cations (Ulitzer and Shilo, 1964). Killifish immersed when ppm) 1.0 to (0.5 concentration lower a at died solution prymnesin 2-ppm a in calcium ions (0.2 to 0.5%) were added to the solution (Igarashi REVIEWS IN FISHERIES SCIENCE Prymnesins act on the cytoplasmic membranes of cells and have been demonstrated been have and cells of membranes cytoplasmic the on act Prymnesins to affect fish, bird, and mammalian erythrocytes; human tumor liver cells (Ehrlich and ascites and HeLa); amnion bacterial protoplasts cells; and spheroplasts; and Mycoplasma permeability increased the of because toxins to sensitive are molluscs, and tadpoles, of the gill membranes of these animals, especially in activated, conditions pH-dependent, that and are sodium cation- chloride-inhibited (Shilo, 1981). noticeably. Vacuolations appeared in the severity of cytoplasm cellular damage was of proportional to increasing the calcium concentrations chloride cells. and higher pH The (Terao and hexose sugars (Ulitzer Kozakai 1981; (Shilo, and identified Shilo, 1970). Six hemolytic (Kozakai components were mixture of galactolipids, with the major fatty acid component being C18:4 and the minor component C18:5, was present in high amounts (Kozakai have two different effects membranes: (1) on it inhibits the sodium-dependent uptake of the transport of neurotransmitters across nerve paralyzed, and they eventually die when immersed in toxin. After metamorphosis, these amphibians are totally refractory to toxins (Shilo and Aschner, 1953). When exposed to prymnesin-2, chloride cells in the epithelium of the gill filaments of the freshwater white cloud mountain minnow, Bufo Recently, Igarashi 1996). Cultures of both crude lipid extracts had toxic effects on human erythrocytes and synaptic vesicles of rat brains (Meldahl that have hemolytic and ichthyotoxic properties (Shilo, 1981; Igarashi

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., et al et al ., 1989; Pseudo- ., 1998), ., 1999), (Takemato et al et al et al ., 1988; Bates ., 1991; Dickey , 3 people died, g/g DA (Addison et al µ ., 1990; Teitelbaum 1990; ., et al ., 1998; Bates, 2000), et al et neuronal swelling, and . (1997), in mammalian

., 2000; Campbell , et al 2+ et al et al ., 1996; Arévalo Chondria armata Mytilus edulis (Subba Rao ., 1990a, 1990b, 1990c; Xi ., 1990; Haya et al ., 1989; Perl 1989; ., ., 2000), New Zealand (Mackenzie et al ., 1996), Japan (Kotaki et al ., 1998). Nerve cells located in the et al et et al et al et al , may not only cause occasional conspicu- occasional cause only not may , ., 1998). Neurotoxicity of DA to rats and ., 1990). DA causes massive depolarization 237 (Table 1) (Bates et al et al ., 1990; Gilgan Nitzschia ., 1989; Wright 1989; ., patelliferum . ., 1992; Adams et al P ., 1994), Scotland (Gallacher ., 2001); Spain (Miguez ., 1990; Bates et al et ., 1995; Rhodes ., 1993). Domoic acid causes extreme neurodegenerative et al Pseudo-nitzschia multiseries et al et al et al et al et al ., 1989; Sutherland ., 1989; Martin et al and one species of et al ., 1992; Garrison ., 1993; Chang ., 1989). Edible tissue of blue mussels contained up to 900 ., 1990; Todd 1990, 1993; Nijjar and Nijjar, 2000). Prior to the PEI event, DA was DA event, PEI the to Prior 2000). Nijjar, and Nijjar 1993; 1990, Todd 1990; ., The production of DA has now been identifed in nine species of DA is an analog of glutamate, an excitatory neurotransmitter that binds to the cell death (Novelli hippocampus are associated with memory retention, hence associated the with loss ASP of cases memory (Bates monkeys has been demonstrated (Tryphonas prymnesiophytes, such as such prymnesiophytes, on impact may also have a substantial kills, but they effects such as fish ous bloom feeding zooplankton’s the constraining by waters coastal in chain food plankton the (Nejstgaard concentrations sub-bloom occurring commonly at even reproduction and and Solberg, 1996). L. DOMOIC ACID Although originally isolated from the red macroalga microalgae by produced be to known not was (DA) acid domoic 1958), Daigo, and in Prince Edward Island shellfish poisoning event. In 1987, prior to a recent human (PEI), Canada, DA was implicated for the first time in Amnesic Shellfish Poisoning (ASP). After the consumption of toxic blue mussels, Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 19 were hospitalized (12 of whom were in intensive care for a time), and more than more and time), a for care intensive in were whom of (12 hospitalized were 19 100 suffered from varying degrees Clinical of symptoms gastrointestinal of DA and intoxication neurologic are diarrhea, illnesses. nausea, memory loss, decreased vomiting, level of consciousness, abdominal seizures, confusion, and cramps, (Debonnel disorientation Wright et al et al of the neurons, with a subsequent increase in cellular Ca 1997). In rodents, DA causes inactivity, seizures, response and (Work a characteristic scratching systems, DA acts as a glutamate agonist, causing damage to neurons by binding to by caused is toxicity the Thus, neurons. the overexciting and receptor glutamate the ampli- involves mechanism this Because mechanism. receptor-mediated indirect, an disorders in mammals and birds. As discussed by Shaw et al et not suspected as a hazard to public health. DA toxicity in mussels was eventually linked to the diatom et al toxin. the of dose high extremely an to consumers exposed which 1989), Stewart, and nitzschia and DA has also been confirmed in shellfish in North America (Bates Denmark (Lundholm 2001), France (Amzil and Portugal (Vale and Sampayo, 2001). kainate type of glutamate receptors. DA is a water-soluble, heat-stable excitotoxin that binds to receptors in the brain, in particular in the hippocampus (Debonnel

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ), et g/ µ Mya over et al ), rock ., 1993; ., (Miguez glacialis Recently, Tigriopus .

Palinurus (Rhodes , there was there , C et al et , the rotifer Scrobicularia T. longicornis Pseudocalanus (Rhodes and and Pecten maximus Pseudo-nitzschia Tiostrea chilensis (Drum ., 1996); Dungeness and ., 2001). This apparent This 2001). ., multiseries ), softshell clams ( et al . Callinectes sapidus P ., 1998); et al et ., 1996); the sea scallops, longicornis and at 72 h was 135 and 38 at 72 h was 135 and . T 50 et al ), and spiny lobster ( et al novaezealandiae Mytilus galloprovincialis Mytilus Siliqua patula Siliqua . (Kotaki ., 1997). Because compounds such compounds Because 1997). ., subtriangulata P (Lincoln ), peppery furrow shells ( shells furrow peppery ), et al et ), dredge oysters ( ., 1984). Feeding rates, egg production Pseudo-nitzschia multiseries Pseudo-nitzschia longicornis ., 1997). Presumably, there are species- . Temora longicornis T et al ) (Rhodes 238 Modiolus modiolus et al ., 1996). Menippe adina P. pungens P. ., 1994a); blue crab ( ., 2001) and Ensis siliqua Ensis et al et al et al Crassostrea gigas Patinopecten yessoensis (Gilgan 1996 in Stewart suggest that the mechanism for action is the same in this ., 1990); Pacific razor clams, razor Pacific 1990); ., (Wekell ), stone crab ( ., 1995). As with other bivalves and certain phycotoxins, there phycotoxins, certain and bivalves other with As 1995). ., Perna canaliculus assimilated DA rapidly for 27 h. Toxin levels then declined in 24-h experiments, the copepods copepods the experiments, 24-h in et al et were not significantly different when copepods were fed toxic et al et ., 1998; Campbell or a nontoxic strain of strain nontoxic a or at very low concentrations. The experimental results for the effects of ., 1994a); razor clams ( clams razor 1994a); ., californicus et al . ) (Gilgan ) T , but only at relatively high concentrations (LC , but only at relatively et al et multiseries ) (Altwein ) . ) (Vale and Sampayo, 2001); clams, Cancer magister P ., 1996); northern horsemussels ( Acartia tonsa A few studies have investigated the effects of DA on zooplankton. Domoic acid Domoic zooplankton. on DA of effects the investigated have studies few A In addition to blue mussels, DA has been found naturally in numerous species multiseries ., 1996); Pacific oysters ( . crab, acuspes toxic fed When seawater. in respectively) ml, greenshell mussels ( Placopecten magellanicus (Arévalo 1996); Japanese scallops, crab ( elephas LANDSBERGfication of the initial concentration response for toxicity. to receptor binding, DA has a low is toxic threshold to the small estuarine copepods REVIEWS IN FISHERIES SCIENCE are species-specific differences in the response of some species to species some of response the in differences species-specific are less than 90% mortality of the copepods plana al a 9-day period. About 50% copepods can act as toxin of vectors (Windust, 1992, cited in Bates, 1998). the When fed ingested DA toxic was retained, suggesting that arenaria Wekell showed no adverse effects, such as decrease in feeding rate, egg-hatching success, unusual feeding behavior, or mortality. As pointed out by Bates (1998), the lack of toxicity to copepods is surprising, given that junctions in crayfish and insects (Maeda DA can depolarize neuromuscular rates, egg hatching success, and mortality of the calanoid copepods and lack of effect on copepods suggests that 1997). Tester, and (Turner copepods consumers zooplankton higher-level to transfer could act as vectors for DA that DA was toxic to the copepods however, another study demonstrated californicus DA on P (Shaw systems mammalian in is it as copepod as PST and -LR copepod (MC-LR) can cease can feeding and be thus detected avoid pounds. the DA, however, by does potential not appear to be toxicity detected chemoreceptors, and avoided of and therefore these the com- probably acts as a neurotoxin (Shaw specific differences in susceptibility to DA. When fed et al Brachionus plicatilis rapidly to a minimum after 75 h of exposure. The mean fecundity of the rotifers was rotifers the of fecundity mean The exposure. of h 75 after minimum a to rapidly reduced considerably, from 0.70 to fecundity 0.39 was eggs attributed to per the female, feeding and inhibition nutritional insufficiency caused (Whyte this by change DA rather in than to of invertebrates, including Mediterranean mussels, Mediterranean including invertebrates, of

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ) ., ., ., the et al et et al et al ., 1990). ., multiseries et al et f. are physiologi- are ., 1993; Wekell 1993; .,

) et al et g DA) in the gonad the in DA) g ., 1994b). In animals In 1994b). ., µ g/g tissue) (Campbell ., 1995b). Mytilus californianus pungens µ . rubida ) (J. Whyte, personal et al et . P C et al ., 1991). After a 72-h period, Pseudo-nitzschia pungens, Pseudo-nitzschia g/g tissue were observed in sea et al , DA concentrations in all tissues µ ., 1992). Because DA is essentially ., 1992). In the digestive system of et al et al 1.9% of the toxin (Grimmelt toxin the of 1.9% Chlamys hastata ± 239 , with the majority of toxin being found in the Pecten maximus ., 1995), there is some indication that Pacific oysters ., 1995), there is some ., 1997), and pink scallops ( scallops pink and 1997), ., spp.), or California mussels ( et al et al et . (1995b) discussed the possibility that because DA causes DA because that possibility the discussed (1995b) . ., 1989), it is more likely that toxin will be excreted than Mytilus et al et et al g/g tissue) and adductor muscle (0.38 to 0.82 g/g tissue), with less than the regulatory limit (20 limit regulatory the than less with tissue), g/g ., 1995a), spiny scallops ( µ µ ., 1995a; Whyte ., 1995a). When Pacific oysters were exposed to to exposed were oysters Pacific When 1995a). ., Placopecten magellanicus et al et al et al et ., 1994b). In king scallops, ., 2001). In blue mussels, DA is found only in the viscera (Wekell viscera the in only found is DA mussels, blue In Maximum DA concentrations of up to 3100 cally stressed (Bates, 1998). Within 4 h of exposure to exposure of h 4 Within 1998). (Bates, stressed cally shells of Pacific oysters closed tightly, which is a typical initial stress response by bivalves to some microalgae (see tissue soft remaining and muscle, PST). gill, mantle, examined: tissues the all in increased Before the shells closed, levels (Jones of DA another initial stress response was a marked increase in the number and activity of hemocytes after a 4-h exposure to the algae. Toxin levels in the oysters increased during the 48 h of exposure, and the number and activity of hemocytes declined from the 4-h peak values to values significantly lower than those of controls after a 24-h clearance period. The suppression in hemocytes following initial toxin response was rectified only after a 48-h clearance number and activity of circulating period, when DA levels in the oyster tissue had declined to trace levels, allowing blood cells to regain their normal characteristics (Jones spp. and to DA. Jones DA. to and spp. Douglas in communiction, encapsulated food a as or form dissolved in mussels live to presented was DA When in liposomes, less than 1% of the dissolved DA and up to 6% of the foodborne DA was incorporated into mussel tissues. body of proportion Domoic relative their For acid kidney. and gland ingested digestive the in as concentrated food was mostly propor- expected than larger retained gills and gland, digestive kidneys, the weight, concentration of toxin in mussel tissues did not tions of the total toxin burden. The decrease consistently over a depuration period of 48 h, or translocated did to DA any tissue appear for storage to (Novaczek be Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS observed excitoxic effects on Although toxin. of levels mammalian high concentrate they because expected be might shellfish systems, physiological aberrations (Douglas scallops in sea on DA of effects adverse obvious no be to appear there (Jones from a naturally contaminated area in PEI, the hepatopancreas, which composed 93.4 contained weight, mussel the of 30% was 50% than more mussels, of lumen gut the in present was DA the of majority the eliminated within the first 24 h (Novaczek hydrophilic (Wright 1997), blue mussels ( (Jones that it will be bioaccumulated (Novaczek mussels domoic acid can be converted to isodomoic acid isomers (Wright 1990). scallops, digestive gland. Levels remained high up to 15 days after exposure (Douglas 1997). In razor clams, DA can also occur in the foot and the siphon as (Drum months six well to up for as retained be can DA in and viscera, the et al except gonad and adductor muscle accounted for 99% of the total individual burden individual total the of 99% for accounted muscle adductor and gonad except 760 to (580 (8.2 to 11.0 et al

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., 1992; , and 95 and , ., 2001). et al et al Pseudo-nitzschia (Work Pelecanus occidentalis Pelecanus was detected, and DA was ., 1992). , died from ingesting anchovies was demonstrated to produce DA produce to demonstrated was et al Engraulis Engraulis mordax australis 240 . P ., 1995), and gut content of sardines (Vale and . At least 150 birds were found during a period . (1997) reported another DA event in Cabo San et al ., 1992; Garrison et al sp. were found in the stomach contents of Pseudo-nitzschia australis et al died, and others suffered severe neurological dysfunction.

, g/g total fish weight resulted in severe neurotoxic symptoms, neurotoxic severe in resulted weight fish total g/g Phalacrocorax penicillatus µ , and Californian anchovies, Scomber japonicus ., 1994a) has been responsible for significant bird mortalities. Although mortalities. bird significant for responsible been has 1994a) ., ., 1995; McGinness Pseudo-nitzschia ., 1992; Fritz ., 1992, 1993). In this case, this In 1993). 1992, ., et al et et al et al et et al are able to vector DA, and toxin transfer via Californian anchovy Recently, Sierra Beltran In May 1998, near Monterey Bay, California, more than 400 California sea lions, sea California 400 than more California, Bay, Monterey near 1998, May In Domoic acid has been detected in the flesh and viscera of bay anchovies, g/g were measured in anchovy viscera, with brain and muscle levels 3 orders of DA was implicated in the 1987 September 1991 human that the first ASP aquatic animal mortality was event attributed to this in toxin. Canada, pelicans, brown 43 least at California, Cruz, Santa In it was not until (Wekell Brandt’s cormorants, contaminated with DA. The birds displayed a characteristic slow, side-to-side head motion, held their wings partially extended, and were unable to fly for more than 10 m without having to either lie land. and reflex, righting Vomiting lose torticollis, was display surroundings, their common, of awareness and the birds would lose on their back or necro- side and hemorrhages were observed with lesions histopathologic and gross their consistent feet paddling slowly prior phosphokinase creatinine to and nitrogen urea blood Serum muscle. skeletal the of death. sis The only detected was acid Domoic controls. in were they than birds affected in higher were in the stomach contents of the sick and Californian dead anchovies, birds, in and the in flesh plankton and samples viscera of dominated (Work by (Buck Sampayo, 2001), but fish mortalities or neurotoxic effects in the field have not been not have field the in effects neurotoxic or mortalities fish but 2001), Sampayo, reported. In experiments, intracoelomic injection of anchovies with DA concentra- 14 to 1 ranging tions magnitude lower, indicating low but measurable DA uptake (Lefebvre including spinning, spiraling, and swimming 1175 upside as high down; as circling levels at DA the mortality. eventual surface, with school; to inability gaping; mouths µ pelicans and mackerel; mouse bioassay of bird stomach contents and HPLC con- firmed the presence of DA. In the mouse bioassay using pelican stomach extracts, mice showed symptoms corresponding to DA toxicity, including akinesis, prostra- tion and scratching, diarrhea, convulsions, and loss of lateral movement and motor coordination. Zalophus californianus Characteristic clinical signs were scratching, dullness, ataxia, seizures, and convul- hippocampus. the of necrosis neuronal were lesions histological Predominant sions. Domoic acid was detected in urine, A feces, concurrent and serum bloom from of examined animals. Altwein DA-contaminated on feeding after killed were pelicans brown where Mexico, Lucas, chub mackerel, of 5 days in January 1996. Live pelicans showed symptoms of intoxication, such as disorientation and agitation, difficulty in swimming, and not themselves being if able they to had right turned upside down drown. At least 50% of during the pelican colony died as swimming a after result months of DA 2 intoxication, causing and disorientation and weakness them of symptoms showed to still birds surviving the event. LANDSBERGAnchoa mitchilli REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , et ., 1992) ., ., 1977) ., 1993); (Sivonen ., 1990). In et al et

et al et al ) ., 2000). The planctonica . et al ., 2000). A et al g/kg body weight, , µ ., 1999). In contrast, et al (Devlin ) (Edwards ) Oscillatoria et al (= in Japan (Park circinalis . A Oscillatoria ., 1999; Scholin (= Planktothrix et al Anabaena flos-aquae ., 1977) that is highly neurotoxic. It is a 241 sp., from the same area (Scholin

et al Planktothrix Microcystis aeruginosa ., 1998; Lefebvre Cylindrospermum et al sp., g DA/g), exceeding values in the body tissues by sevenfold and suggesting and sevenfold by tissues body the in values exceeding DA/g), g µ 5 ± intraperitoneal mouse dose for purified toxin is about 200 The presence of DA in shellfish and fish certainly provides an opportunity for 50 ., 1989), and in benthic mats of mats benthic in and 1989), ., this report remains to be confirmed (Codd, 1998). Anatoxin-a is a low-molecular- weight secondary amine (Devlin postsynaptic cholinergic nictotine agonist that blockage of neuromuscular transmission causes and subsequent respiratory paralysis. The death via a LD depolarizing al (Table 1). Most recently (and uncommonly) anatoxin-a microcystin-producing strains of was also reported from Aphanizomenon and has also been reported from strains of 1. Anatoxin-a Anatoxin-a was originally isolated from with a survival time of 4 to 7 min (Carmichael, 1988, 1992). Symptoms of anatoxin- a toxicity in mouse bioassays include muscle fasciculation, gasping, loss convulsions, and of death by coordination, respiratory staggering, include arrest animals (Carmichael domestic and wild in poisoning of signs the reports, field muscle fasciculations, gasping, convulsions, and opisthotonus (in birds). Death by respiratory arrest occurs within minutes to a dosage, and prior few food consumption. Animals hours, 1988). need to (Carmichael, ingest bolus only depending lethal a a few receive milliliters on to bloom surface toxic species, the of liters few a to M. ANATOXINS There are three main anatoxins that have been isolated from cyanobacteria: ana- toxin-a, anatoxin-a(s), and homoanatoxin-a (Table 1). In addition to the terrestrial animal mortalities associated with drinking anatoxins (Table 7), documented reports there of the effects are of anatoxins on a aquatic life. few Because several of also produce microcystins or PST (Table 1), the known anatoxin-producing species than rather toxins these with associated are water in mortalities most that appears it (see anatoxin-a(s) with associated was birds of mortality recent a but anatoxins, the below). confirmed confirmed in Californian anchovies Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS of DA in muscle tissue (Lefebvre minimal bioaccumulation primary route of sea lion exposure to DA was considered to be through consumption through be to considered was DA to exposure lion sea of route primary of toxic fish. The highest DA concentrations in anchovies occurred in the viscera (223 analyses Histopathological DA. of amounts trace only or no contained mussels blue anterior the in lesions brain revealed DA to exposure acute suffered that lions sea of ventral hippocampus. This was the first proven case of fatal DA toxicity in marine mammals (Gulland high the at accumulated not is DA if even and chain, food the through transfer toxin lethal doses recorded during mortalities of birds and mammals, there could still be or neurological selective the with as However, effects. health chronic for potential a gastrointestinal impacts of other toxins on vertebrates, it is unclear effects what could be chronic anticipated.

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., was Ana- et al minor ., 1992; ., 1991). Anabaena ., 1998). In et al Brachionus var. NRC 525-17 et al , et al g/l) in the water appeared to be Cryptomonas µ ., 1997a). Although 1997a). ., and two clones of et al et ) (Edwards flos-aquae fed . (Barros A lemmermannii . Planktothrix A Asplancha girodi ., 1997) confirmed for the first (Carmichael and Gorham, 1978) Gorham, and (Carmichael Oscillatoria et al Asplancha girodi ., 1992; James 1992; ., g/kg, about 10 times more lethal than lethal more times 10 about g/kg, (= did not release extracellular toxin and µ (IC-1) that produces anatoxin-a. In the In anatoxin-a. produces that (IC-1) et al et ., 1995). ., 1990). , reproduction was suppressed at an g/ml; the other species’ reproduction was g/ml. 242 Daphnia pulicaria Synchaeta pectinata µ µ ., 1997). The strain et al et al et al Anabaena strain NRC 525-17 was recently shown to inhibit , and Planktothrix Anabaena flos-aquae Anabaena ., 1997a). After drinking surface water, dogs displayed ., 1997a). After drinking calyciflorus ., 1990). Anatoxin-a(s) was first reported from a culture . B (i.p. mouse) of 20 of mouse) (i.p. et al isolated from Buffalo Pound Lake, Saskatchewan, Canada 50 et al Anabaena flos-aquae Anabaena flos-aquae (Henriksen . A ., 1997). Extracts of field samples were neurotoxic to mice and were not inhibited reproductively by filtrates of very dense very of filtrates by reproductively inhibited not were Keratella cochlearis ., 1997a). , et al ., 1992; James acetylcholinesterase activity in et al lemmermannii et al dry-mass concentration of 0.5 . A There is little information on the effect of anatoxin-a(s) on aquatic animals. Recently, several dog mortalities have been associated with the production of Few studies have determined or investigated effects by anatoxin-a on aquatic Anabaena flos-aquae calyciflorus . subsequently showed an anticholinesterase activity similar to anatoxin-a(s). Neither anatoxin-a nor saxitoxin or its derivatives were detected by HPLC, which together with the pharmacological evidence (Henriksen and can simultaneously produce anatoxin-a(s) and microcystins (Harada Anatoxin-a(s) is a potent acetylcholinesterase inhibitor (Mahmood and Carmichael, LD an with 1987) 1986b, of (Carmichael and label), [s] Gorham, the — toxin 1978). the for terminology In the gives (which salivation mouse viscous marked or rat bioassays, symptoms include lachrymation in mice, chromodacryorrhea (red-pigmented tears) incontinence, muscular in weakness, fasciculation, convulsion, rats, defecation, and death urinary from respiratory failure (Carmichael July 1993 and June-July 1994 at lakes in Denmark, deaths of wild birds coincided with massive cyanobacterial blooms dominated by (Onodera 2. Anatoxin-a(s)by produced is Anatoxin-a(s) anatoxin-a (Carmichael Anatoxin-a(s) from in vitro time that anatoxin-a(s) was the cause of deaths of wild animals (Onodera LANDSBERGthose from distinguish to difficult often are vertebrates in anatoxin-a of signs Clinical of PSP. Both result in all or some of the coordination, following staggering, and symptoms: collapse and death trembling, by respiratory loss failure. Anatoxin-a of additionally causes salivation (Negri REVIEWS IN FISHERIES SCIENCE severe respiratory discomfort, coma, rigors, cyanosis, limb twitching, hypersaliva- (Gunn dying before convulsions and tion, anatoxin-a from benthic mats of anatoxin-a from benthic Gunn no surface blooms were detected, benthic mats of linked to the presence of high levels of anatoxin-a (up to 444 1997). Although the previous involvement of the toxin in the poisonings of dogs (James organisms. Reproduction in the planktonic rotifers calyciflorus baena B suspensions, which showed that thus could inhibit only rotifers that ingested it (Gilbert, 1994). suppressed at a concentration of 4.0 inhibited by a strain of strain a by inhibited most susceptible species,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ), was 32 ., 1992;

, flos-aquae et al . g/ml lupenyl g/ml commersoni A ) rather than

µ C ) were exposed Aphanizomenon flos- ., 1997). The toxin is Planktothrix formosum Planktothrix ., 1992). In preliminary In 1992). ., . (1968). Young-of-the- (1968). . ., 1989) was suspected tamarense cells/ml. Survival times Rana clamitans 2

Catostomus commersoni et al et et al et et al et al Danio rerio 10 × ., 1975; Skulberg et al , from 15 to 60 min for min 60 to 15 from , . When injected intraperitoneally with intraperitoneally injected When . Alexandrium g/ml lupenyl acetate, larvae had edema in edema had larvae acetate, lupenyl g/ml µ 243 ), white suckers ( g/ml mueggelone, there was no effect on larval µ ) were exposed to lyophilized cells of cells lyophilized to exposed were ) was described by Sawyer by described was blooms on the basis of toxicological properties and it is unclear if other mechanisms are operating. Lepomis gibbosus Lepomis

, Lebistes reticulatus Lebistes g/ml mueggelone for up to 32 hours, there was a 45% NRC-44-1 (Carmichael µ ., 1968). Whether these mortalities were due to the potent ) strain NIVA-CYA-92 (Skulberg NIVA-CYA-92 strain ) ., 1997). When zebrafish larvae ( Lepomis gibbosus et al flos-aquae et al flos-aquae . . flos-aquae A A

., 1988) and of birds and swine (Cook flos-aquae . ., 1997). Lebistes reticulatus Lebistes A ., 1997). Field reports of toxicity are currently unknown. et al et al C (Sawyer o et al (Papendorf Another experiment in which fish and green frog mortalities were associated Oscillatoria formosum Oscillatoria varied from 30 to 240 min for min 240 to 30 from varied for min 240 to 60 from and a dosage of 60 mg/kg, the survival time for the green frog, of case the In unknown. is toxins undescribed other or PST, mueggelone, of effects exudates toxin by affected be can fish that appears it blooms, dinoflagellate and PST (see section on uncharacterized toxins and from just PST, which Aphanizomenon are normally ingested (see PST), but in the case of N. MUEGGELONEacetate lupenyl known previously the and mueggelone, lipid, C18 identified newly A were isolated from a field-collected freshwater sample of (Mahmood (Mahmood to be related to chromatographic results, the causative agent (Onodera was not characterized chemically 3. Homoanatoxin-afrom isolated was homoanatoxin-a, neurotoxin, new a Recently, (= aquae min at 12 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS experiments, homoanatoxin-a was shown to be toxic to mice when i.p. injected — phrenic isolated the in neurostimulation by induced contractions muscular blocked it nerve-hemidiaphragm preparation of the rat (Lilleheil readily absorbed from the gastrointestinal tract in induced mice after that oral exposure than to algal less was exposure such from resulting toxicity the but suspensions, death caused and quickly very acted Homoanatoxin-a admininstration. parenteral by by respiratory arrest within a few minutes, an action similar to that of produced anatoxin-a by Lilleheil to a concentration of 10 mortality. In larvae surviving from 24 to 32 developed, h, whereas no after blood 3 circulatory days system there had was region. edema At a and concentration thrombosis of 1 in the heart obtained from lake water containing approximately 4 development. At a concentration of 100 of concentration a At development. ( guppies and the heart region and showed tail bending (30%) after 3 days. After 5 days, all larvae all days, 5 After days. 3 after (30%) bending tail showed and region heart the 10 of concentration a At tails. bent and region heart the in edema had acetate, there were no obvious effects on development. Lupenyl cause acetate any did larval mortality. not to exposure with year pumpkinseeds (

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., after et al et were found were (Harada , and rainbow racbiorskii cultures proved . C Cylindrospermopsis raciborskii . C ., 1997) (Table 1). An out- raciborksii . ., 1998). At the ultrastructural C ., 1985). Umezakia natans Umezakia et al et al ., 1985). The majority (139) of cases of (139) majority The 1985). ., et al was implicated in mortalities of cattle Cherax quadricarinatus et al et ., 1992), ., (Banker 244 et al et g/g) at a concentration five times that of the µ , exposed to a natural bloom of bloom natural a to exposed , was 64 mg of freeze-dried culture per kg mouse. (Ohtani 50 ., 1985, 1997). Extracts of ., 1998). After drinking from a contaminated area, a calf ., 1983; Hawkins 1983; ., . The accumulation of CY in the flesh of crayfish raised some raised crayfish of flesh the in CY of accumulation The . et al et al et et al g/g) in crayfish, but no mortalities occurred. No histopathological Cylindrospermopsis raciborskii died within 7 to 8 h of inoculation. Livers were reddened, swollen, in Lake Griffin, Florida, suggest a possible connection, but this has not has this but connection, possible a suggest Florida, Griffin, Lake in µ raciborskii . ., 1994). Aphanizomenon ovalisporum C in aquaculture ponds in Australia. Toxin accumulated in the hepatopan- et al Melanotaenia eachamensis Melanotaenia ., 1983; Hawkins Recently, Reports of the effects of CY on aquatic organisms are few, but it is likely that raciborksii raciborskii . . in the gut contents, indicating that ingestion of cells was one mechanism of accumu- CY toxin exposures, experimental In 1999). Eaglesham, and (Saker accumulation lated in the hepatopancreas (4.3 muscle (0.9 abnormalities were observed in crayfish exposed either of cultures to case extracellular the in or As 1999). Eaglesham, pure and (Saker effects long-term possible about concern Affected mice were huddled, anorexic, and had mild limb diarrhea; occasional exhibited those and respiration dying gasping in slow, had the h) 9 to (6 times shortest paddling. The principal lesion produced was centrilobular to massive hepatocyte necrosis in the liver, but various degrees of injury were also seen in the kidneys, adrenal glands, lungs, and intestine (Hawkins 1994), and in Australia (Thomas became weak and was seen staggering before it died. Gross pathology indicated severe abdominal and thoracic hemorrhagic effusion, hyperemic mesentery, pale and swollen liver, and an extremely distended gall bladder containing dark yellow bile. Histopathology of the liver revealed extensive fibrosis and bile duct prolifera- tion. Two mice that received a C 1.0-ml-i.p. dose of freeze-dried pure culture of fish, raciborskii of Trichomes fish. the of viscera and crayfish the of creas LANDSBERGO. CYLINDROSPERMOPSIN Cylindrospermopsin (CY) is a raciborskii Cylindrospermopsis hepatotoxin that was isolated recently from REVIEWS IN FISHERIES SCIENCE that livers had mice Control weight. body total the of 13.0% to 12.6 represented and were 5.7 to 7.7% of total body weight (Thomas (148) were in children. The liver was enlarged in all cases, and the initial symptoms initial the and cases, all in enlarged was liver The children. in were (148) resembled hepatitis accompanied by abdominal profuse pain. bloody Kidney diarrhea malfunction followed. Symptoms and copper occurred sulfate to after a dense the algal application bloom in of the et water al supply (Blyth, 1980; Bourke protein of inhibition showed CY to exposed experimentally mice of livers the level, synthesis, membrane proliferation, fat-droplet accumulation, and finally cell death (Terao effects do occur. Recent C mortalities of alligators and yet been concurrent confirmed (J. Burns, personal blooms communication). Cylindrospermopsin was of shown to accumulate in redclaw crayfish, break of human hepatoenteritis was associated with a bloom of northeast- Island, Palm on contaminated became reservoir water drinking domestic a (Bourke Australia ern to be lethal to mice; the LD

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., 1998; . Other -treated ., 2000). is almost et al et al viridis Anabaenopsis . of MC-LR i.p. or M 50 spp., (Lanaras and Cook, Microcystis and ), but it is thought that g/kg (Dawson, 1998, and Cylindrospermopsis µ agardhii Anabaena ) , g/kg, and the inhalation toxicity µ Oscillatoria = 43 43 = 50 and Oscillatoria -amino acids at positions 2 and 4. The most The 4. and 2 positions at acids -amino L 245 blooms are real (Humpage (= Microcystis aeruginosa ., 1994; Kaya 1996; Carmichael, 1996, 1997; tumor initiation was demonstrated when Swiss Anabaena et al Anabaena flos-aquae Anabaena in vivo Planktothrix 180 mg/min/mm, LD 50 Cylindrospermopsis ., 1994; Sivonen, 1996; Carmichael, 1997; Namikoshi et al spp., and followed by either dimethyl sulfoxide or tetradecanoyl phorbol acetate phorbol tetradecanoyl or sulfoxide dimethyl either by followed produces only microcystins (Carmichael, 1992, 1997). Unlike other toxic other Unlike 1997). 1992, (Carmichael, microcystins only produces ., 1998; Sano and Kaya, 1998; Sivonen and Jones, 1999) (Table 1). The Nostoc , et al As with other microalgal toxins, most information regarding the mechanism of Until recently, it was known that other cyanobacteria can produce both Cylindrospermopsin Cylindrospermopsin was demonstrated recently to have carcinogenic activity. raciborskii . animals showed histological evidence of showed frank tumors, including neoplastic one fibroblastic osteosarcoma, one processes; hepatocellular three small statistically of a represent results these Although lymphoma. one and carcinoma, which sample, Falconer and Humpage (2001) recognized public health significance of such findings and recommended the further investigation. potential biological and references therein). Symptoms of microcystin intoxication are diarrhea, vomiting, piloerection, weakness, and pallor (Bell and Codd, 1994). Microcystin targets the Microcystis planktonic cyanobacteria, which may have nontoxic strains, always toxic. Microcystins microcystins of are variants structural the 60 About most 1996). (Carmichael, common cyanobacteria by cause caused of water-based toxicosis been have which of half than more 1999), Jones, and (Sivonen known currently are isolated from species and strains of by produced are microcystins are acids amino variable the where (MC-LR), microcystin-LR is microcystin common leucine (L) and arginine (R) (Dawson, 1998). action has been obtained from rodent bioassay models. The LD i.v. in mice and rats is in the range of 36 to 122 Albino mice were orally C exposed with a crude (known saline tumor promoters). After 30 weeks, 5 extract out of 53 of freeze-dried P. MICROCYSTINS been have microcystins of pathology and toxicity, biology, the on reviews Excellent published recently (Rinehart Dawson, 1998). As with other cyanobacteria, most they after wildlife, and livestock including animals, terrestrial in documented toxicity microcystin reports concern ingest contaminated pond water (Table 7). effects of This microcystins in section aquatic organisms. briefly reviews known microcystins and anatoxins (e.g., two the in principally differ microcystins of PST produced by dinoflagellates (see PST), there is a high transfer likelihood up of the toxin food chain. Cylindrospermopsin induces cytogenetic damage via two mechanisms: one at the DNA level inducing strand breaks, and the other findings at These (aneuploidy). the chromosomes whole level of loss of inducing function, the spindle kinetochore/ substantiate the concern that potential public health risks associated with drinking water sources and Preliminary evidence for milleri 1994; Rinehart Sano in mice is similar: LCT Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al et ., 1993; ., ., 1981; et al et et al ., 1999). ., 1981; Eriksson et al et al ., 1990; Falconer, 1991; ., 1990, 1994; MacKintosh 1994; 1990, ., ., 1996). The incidence of et al et al et et al ., 1989) (up to 100% increase) ., 2001). ., 1997). Microcystins do not easily ., 1991; Carmichael, 1992; Dawson, et al et al et al et al 246 ., 1990; Honkanen Honkanen 1990; ., ., 1990; Matushima et al et et al ., 1990); by depolymerization of intermediate filaments ., 1990; Hooser ., 1991, 1992; Ito ., 1998; Carmichael et al et al et al et al ., 1990; Yoshizawa ., 1990; Honkanen ., 1990; Fujiki and Suganuma, 1993). In countries where water supplies contain ., 1990; Fujiki and Suganuma, 1993). Microcystins are also potent tumor promoters that are mediated through the When quail were exposed to microcystins, the toxic effects observed were quite were observed effects toxic the microcystins, to exposed were quail When In aquatic systems, most documented cyanobacterial effects are those on (Dawson, 1998). Microcystins mediate their toxicity by uptake into liver hepatocytes liver into uptake by toxicity their mediate Microcystins 1998). (Dawson, protein serine/threonine of inhibition by systems; transport acid bile multispecific via (Eriksson 2A and 1 phosphatases LANDSBERGwhich liver, the in blood of pooling and necrosis, damage, cytoskeletal causing liver, consequently increases liver weight (Hooser which leads to loss and by disruption of the liver cytoskeleton, and microfilaments; (Hooser necrosis cellular and adhesion, cell-to-cell of loss morphology, cell of REVIEWS IN FISHERIES SCIENCE et al 1991; Falconer and Yeung, 1992; Eriksson and Goldman, 1993; Runnegar 1993; Goldman, and Eriksson 1992; Yeung, and Falconer 1991; of shrinkage or rounding cause microcystins doses, toxic acutely At 1996). Sivonen, the of disorganization This structure. hepatocyte normal of loss and hepatocytes the animals of death the by followed often hemorrhage, hepatic massive to leads tissue from hypovolemic shock or hepatic insufficiency (Falconer et al 1998). Death can occur within a few hours after a high dose (Falconer Bell and Codd, 1994; Dawson, 1998). More poisoning recently, was a disastrous caused case by of the acute unusual hemodialysis: exposure about of 50 humans people to died microcystins in Brazil via when received patients untreated at water a dialysis contaminated center with reservoir microcystins (Jochimsen from a nearby water inhibition of protein phosphatase type 1 and 2A as such inhibitors, phosphatase protein activities. other of modes from different be Their to appears mode of action okadaic acid (Falconer, 1993), and their effects are organ specific (liver) (Falconer and Buckley, 1989; Honkanen cyanobacteria, the potential threat of primary liver cancer by human exposure to microcystins has been recognized (Yu, 1991; Ueno penetrate epithelial cells and they do not promote tumors on mouse skin (Matushima skin mouse on tumors promote not do they and cells epithelial penetrate et al primary liver cancer in the populations of Qidong County, Shanghai, China, where is about eight times higher than in populations people drink pond and ditch water, drinking well water (Yu, 1991), and microcystins were found in the cancer areas incidence where was high (Yu and Chen, 1994). different from those observed in rodents. injection, Quails whereas rats usually and mice died died in 1 14 to 3 to h. After 18 variant the microcystin-RR, quail administration spleens h were of enlarged to the double after the size of those of were necroses hemorrhagic Slight liver. the in change no was there but controls, the necroses, Such tissue. subcutaneous and intestine, liver, gizzard, the in observed also except for those in the liver, were not seen in mice and rats (Takahashi and Kaya, 1993; Kaya, 1996). Although the experimental effects in of birds appear to be different from those in mammalian models, a recent mortality birds such as the quail in Japan also indicated hepatotoxicity (see below, Matsunaga Nishiwaki-Matsushima zooplankton. There have been a large number of studies of individual species and of specific trophic levels in which 1987; 1982, Lampert, see (e.g., investigated were the habitats freshwater in zooplankton fate of microcystins and their effects on

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ) ., ., et was et al Aedes et al ., 1991). g/ml) and g/ml) to aquatic ), daphnia strains that strains µ Oscillatoria et al have ranged have pulicaria was the most . (= D , 11.6 , sp. were toxic to was exposed to 50 ., 1991; Jungmann, ; nevertheless only agardhii Microcystis ., 1994, 1995a, 1995b, . Intermediate survival Intermediate . .) et al O ., 1991; Peñaloza Artemia salina ., 1998). The lethality of et al (=

(48-h LC (48-h . et al P Planktothrix Microcystis Microcystis et al birgei Microcystis and or was associated with low sensitivity low with associated was Microcystis aeruginosa Microcystis was associated with greater physiological greater with associated was ., 1990; Jungmann g/ml). Survival in the presence of toxic 247 pulicaria Daphnia hyalina Daphnia µ . Microcystis Microcystis D et al pulex ., 1992; Reinikainen . D g/ml), et al , 21.4 µ 50 ., 1991). In some cases, in addition to microcystins, other g/ml) were intermediate in sensitivity, and sensitivity, in intermediate were g/ml) µ et al was positively associated with food selection capabilities and ., 1986; Kiviranta and Abdel-Hameed, 1994; Reinikainen , 0.45 to 1.0 to 0.45 , , 9.6 , ., 1980; Lampert, 1981a; Jungmann 50 ., 1995b). Fractions obtained from 50 et al ), and aquatic larval stages of the yellow fever mosquito ( et al ., 1991; Watanabe et al was strongly influenced by both physiological sensitivity and feeding ., 1991). When the copepod may affect aquatic organisms even though they have no known effect on et al (48-h LC (48-h ) (Nizan et al ., 1990; DeMott Until recently, little information regarding the toxicity of Diaptomus birgei pulex ., 1997). As previously noted, it is important to recognize that the interpretation . Daphnia pulex the least sensitive (48-h LC agardhii that shown have Bioassays mammals. sensitivity and nearly uninhibited feeding on toxic on feeding uninhibited nearly and sensitivity by (De feeding uninhibited and sensitivity physiological high with associated negatively Mott microcystin-LR, no toxicity was demonstrable even at high concentrations (Shaw al (and organisms whole to is exposure when different be may organisms on effects of therefore potentially to multiple toxins) rather than purified toxins. animals was available, yet their impacts systems. Studies have largely concentrated ecologically significant than to terrestrial in aquatic systems are probably on more the effects of microcystin-producing cyanobacteria on terrestrial mammals, and same the by caused was zooplankton to toxicity that assumed been sometimes has it compound (Lampert, 1987). As Jungmann (1992) recently stated, “toxicity to higher animals cannot be of evolutionary adaptive value to a few publications deal with the response of MC-LR in lake ecosystems” (Peñaloza et al bioactive compounds produced by were nontoxic to mice were highly toxic to brine shrimp ( ( 1992, 1995; Jungmann and Benndorf, 1994; Kiviranta Reinikainen and Abdel-Hameed, 1994; of that to similar weight molecular a had fractions these although and zooplankton, to purified toxin and a rapid inhibition of feeding in the presence of toxic cells. In of survival poor very the contrast, Infante and Abella, 1985; Fulton and Paerl, 1987; De Bernardi and Guissani, 1990; De Mott 1999; Christoffersen, 1996; Hanazato, 1996; Sbiyya microcystins to zooplankton has been 1981b; well Fulton and Paerl, documented 1987), but the actual (e.g., dose of ingested Lampert, toxin depends on 1981a, feeding behavior as well as cell concentration in the water (De Mott D Microcystis by survival good Relatively behavior. aegypti been recently has microcystins to addition in compounds toxic for Evidence 1995b). found for several species (Peñaloza Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS For example, reactions of daphnids exposed to exposed daphnids of reactions example, For some in dying; to cells, the avoiding completely to source, food a as cells using from cases, responses depend upon (De the Bernardi toxicity of individual microalgal strains 1990). Acute toxicity experiments with purified toxins, toxic cyanobacteria, and cell experiments with purified toxins, 1990). Acute toxicity extracts showed that four species of zooplankton differed markedly in their physi- ological sensitivity to microcystin. The copepod LC (48-h sensitive

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 Cyclops ., 1996). ., g/g. The PCC7806 Daphnia Lymnaea ., 1997b). ) or white or ) µ Microcystis Microcystis Microcystis ]), and tad- et al et et al ., 2001). g/g dry weight) dry g/g ., 1997; Williams µ et al ) were exposed in Esox lucius Esox et al ., 1995b). Some strains Some 1995b). ., were high because of toxin/l), swan mussels . Other dominant species, dominant Other . et al et and the copepod Helisoma trivolvis fatalis in which the toxin was present was toxin the which in Microcystis flos-aquae ., 1996a; Williams . [as B produced microcystin-RR, whereas microcystin-RR, produced ., 1992). In a recent field study in et al Anodonta cygnea Oscillatoria ]) in amounts of up to 120 g ., 1997; Watanabe et al µ Bosmina fatalis Bosmina . (1996b) confirmed the presence of MC-LR Artemia 248 et al et al ) larvae survived acute exposures to toxic to exposures acute survived larvae ) ), although the HPLC method used would not have not would used method HPLC the although ), Physa gyrina was less than 10,000, and the molecular weight of ., 1996b). A new compound that is toxic to toxic is that compound new A 1996b). ., Oscillatoria agardhii Oscillatoria Planorbella trivolvis [as are common, microcystin (75 to 1387 to (75 microcystin common, are et al et Diaphanosoma brachyurum PCC7806 is 35,000 (Reinikainen (Reinikainen 35,000 is PCC7806 g toxin per mussel. The highest concentrations were present were concentrations highest The mussel. per toxin g agardhii µ that were toxic to toxic were that ., 1992; Hanazato, 1996). In Lake Kasumigaura, Japan, where .) O Microcystis et al (=

. ., 1990; Bury 1990; ., Microcystis P Procambarus clarkii Procambarus Physella gyrina , while juvenile crayfish tolerated toxic strains better than nontoxic ones. nontoxic than better strains toxic tolerated crayfish juvenile while , rubescens et al et .) ), marsh rams-horn ( O Catostomus commersonii Catostomus , that co-occurred with the bloom did not accumulate microcystin because (=

C, it was inferred that other bioactive compounds are produced by this species are produced bioactive compounds inferred that other C, it was ., 1989; Vasconcelos, 1995; Prepas ., 1997a; Amorim and Vasconcelos, 1999), and although there do not appear to ., 1997a; Amorim and Vasconcelos, 1999), and although there do not appear Among the zooplankton, there is an apparent differential toxicity of there is an apparent differential Among the zooplankton, Crayfish ( Crayfish There have been a few reports of microcystin accumulation in bivalves (Eriksson bivalves in accumulation microcystin of reports few a been have There . o P (Jungmann and Benndorf, 1994; Jungmann, 1995). The molecular weight of a toxin 1994; Jungmann, 1995). The (Jungmann and Benndorf, isolated from several lakes in Alberta, Canada, Kotak was isolated and purified from water extracts of from toxin the of (Feuillade compounds uncharacterized different, produced strains other to rotifers, copepods, and cladocerans (Gilbert, 1990), effects and have been found a (Tables 10 and 11). range Copepods avoid handling of sublethal behavior (Fulton and Paerl, selective chemosensory feeding because of their highly microcystins, to exposure by unaffected apparently are species certain Because 1987). this allows for accumulation of toxins and for their potential transfer up the food chain (Watanabe vicinus of feeding avoidance. Because mortalities predation by fish and of prawns, it was suggested that microcystins would be trans- ferred to higher trophic levels (Watanabe natural blooms of blooms natural in phytoplankton, zooplankton, and in the gastropods swamp lymnaea ( be any short-term lethal effects it chronic is health effects. unclear When whether swan mussels these ( animals experience (Peñaloza cladoceran the in accumulate to found was stagnalis pole physa ( et al LANDSBERGMC-LR, toxicity was destroyed by boiling. Because microcystins are heat-stable to 160 REVIEWS IN FISHERIES SCIENCE such as the cladoceran Because crayfish accumulate microcystins primarily in the intestine and hepatopan- Because crayfish accumulate microcystins primarily in the intestine and creas, this is not considered to pose a risk removed from to the animals human prior health to consumption if (Vasconcelos these organs are absence of detectable MC-LR in numerous other macroinvertebrates (e.g., damselfly (e.g., macroinvertebrates other numerous in MC-LR detectable of absence from up taken not is toxin the that suggested chironomids) and larvae dragonfly and the water or that, given its high water solubility, it is rapidly eliminated. The MC- LR present in gastropods was thought ( pike northern of livers to the in detected not was have MC-LR system, been ingested. In the same lake ( sucker detected covalently bound microcystin (Kotak aeruginosa et al of strain a to aquaria laboratory at low concentrations (up to 40 to 60 accumulated up to 280 to up accumulated in the hepatopancreas, and low levels were present in the intestine, gonad, muscle, gonad, intestine, the in present were levels low and hepatopancreas, the in

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ) g µ . (1997) minimum . et al P ., 1997a). The ., 1997). When g DTX-1, 0.2 g/l) concentrations g/l) µ et al g/l) after 4 days of µ et al µ Pyganodon grandis (= , were experimentally ex- (Prepas over a 4-day period, they g OA, 30 µ ., 1993; Williams et al 249 grandis simpsoniana . Ondatra zibethicus A ) has led to the suggestion that “VSP” may in fact ., 1993). The need to identify the bioorigin of such of bioorigin the identify to need The 1993). ., Mytilus galloprovincialis Microcystis aeruginosa et al et . (1993) detected 15 ., 1993; Luu g per 100 g of shellfish. Williams From cultivated mussels collected from Prince Edward g MC-LR without any detectable ill effects. After 2 weeks µ

µ et al et al ., 1997a). cells/ml Prorocentrum 5 et al ., 1997a). , it was determined that the freshwater clams accumulate microcystins accumulate clams freshwater the that determined was it , et al g/l) accumulated measurable levels (between 13 to 21 µ ., 1989). However, recently it has been determined that microcystins can occur The recent detection of microcystins in marine mussels (species not identified) When the freshwater clams aeruginosa . detection of microcystins in bivalves, the reports of venerupin shellfish poisoning toxic consumed people after Japan in hepatoxicity human associated the and (VSP), characterized chemically been not has venerupin that fact the and clams, and oysters (see section under dinoflagellate the with not and microcystins with associated been have marine microcystins in the apparent absence of known microcystin producers is an marine microcystins in the apparent absence of known microcystin producers important priority. (Williams Island, Canada, Chen microcystins of concentrations low to humans of exposure chronic that concerns raised that likelihood The cancer. of risk the increase could consumption shellfish through raised; been has inhibtors phosphatase protein produce cyanobacteria marine many for example, 60% of 15 species Chen in results unpublished produced such inhibitors (Holmes and Taylor, without further exposure, low levels of microcystin were still present in the feces (Amorim and Vasconcelos, 1999). from Vancouver Island and Prince Edward Island, Canada, and (proposed from intoxication shellfish of type new a for potential the the identified has lands, Nether- as hepatotoxic shellfish poisoning [HSP]) that could represent a serious hazard to human health (Chen microcystin-LR, and 0.02 kidneys, and was it connective mice, to lethal were tissue. toxins the because Because and tissues mussel swan high the in present concentrations of the toxin were concluded that microcystins were not 2 months in clean water, toxins were still metabolized detectable in the swan mussels (Eriksson in the swan mussels. et al After either as a covalent complex microcystin will detect or only low levels as of free toxin, free whereas levels of toxin. covalently bound toxin Enzyme can be thousands of times and higher and can be detected HPLC only by other assays methods. for Therefore, earlier studies in which only free toxins were measured may have significantly underestimated the concentrations tissues of (Williams microcystins present in were exposed either to dissolved microcystin M (MC-LR) or to natural blooms of principally by ingesting (55 high relatively toxic to exposed clams Freshwater phytoplankton toxin. dissolved and minimally whereas toxin, of levels significant via accumulate not did days 3 uptake for MC-LR dissolved of of the phytoplankton intact in present concentrations toxin lower far the to exposed those (< 4 days 21 than more for clams freshwater in microcystins of persistence The exposure. was considered to be important because mammals such of as the the potential muskrat, transfer of marine Mediterranean toxins mussels, to posed daily to 10 accumulated up to 16.0 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., ., ., et al et et al et et al et al ., 1994; ., 1997, ., 1994; ., 1994). flos-aquae et al et al et al . et al A . As recently ., 1995, 1997b; . Intraperitoneal . et al Anabaena flos-aquae Anabaena ., 1993; Johnston 1993; ., ., 1996; Bury , did not die and showed Microcystis et al et ., 1990; Johnston et al scum was highly hepatotoxic ., 1991; Tencalla Gambusia affinis et al , died when when died , et al ., 1994). Analyses of the water and ., 1994; Williams ., 1998a). et al flos-aquae . et al et al A ., 1998a). Histopathological evidence was 250 ., 1991; Andersen 1991; ., ., 1996a; Sahin ., 1994). Salmo trutta Salmo Oncorhynchus mykiss ., 1995, 1996a) that may be detrimental to their et al ., 1985; Sugaya ., 1995). When exposed to freeze-dried toxic et al et et al ., 1994), (2) fish exposed to cyanobacterial extracts cyanobacterial to exposed fish (2) 1994), ., et al et al ., 1985; Råbergh . (1990) demonstrated that a soluble fraction obtained fraction soluble a that demonstrated (1990) . et al or by gavage), and bath immersion (Phillips et al et al et et al et al et ., 1994; Tencalla libitum

., 1994; Bury et al ad ., 1995). Consequently, the exact cause of natural death, that is, ., 1990; Råbergh 1990; ., sp. was lethal to mosquitofish, et al ., 1994; Bury et al et al et ., 1996a, 1996b; Kotak in water, rainbow trout, ., 1996b; Zambrano and Canelo, 1996). Fish have been experimentally et al et al Microcystis et al Although algal blooms producing microcystins have been linked directly with Recent research has advanced a number of explanations for the fish kills (Bury Approximately 1000 brown trout, brown 1000 Approximately ., 1998a): (1) fish may ingest toxins or toxic cyanobacteria, which may then result then may which cyanobacteria, toxic or toxins ingest may fish (1) 1998a): ., fish determined that the most likely cause of mortality was exposure to microcystins to exposure was mortality of cause likely most the that determined fish and the consequent hepatotoxicity. The to mice in intraperitoneal assay. In addition to changes in the liver consistent with hepatoxin exposure, some pathological changes to the gills caused acute irritation and mucus production. The cause to the density of the bloom, high pH, (2) physical irritation due (1) irritation due to of the gill pathology was (3) a direct suggested toxic effect of to the toxins, or (4) be a combination of all of Hepatic these factors. pathology was indicative of toxic damage and may directly have through ingestion resulted or absorption either of toxins or secondarily after gill damage and a consequent increase in epithelial permeability that allowed toxin to accumu- late in the fish’s tissues (Rodger such of majority The unclear. is exposure of route exact the 7), (Table mortalities fish oxygen high the from resulting conditions hypoxic to attributed been have kills fish (Bury senescence bloom and/or night at respiration bloom by caused demand bloomed in Loch Leven, Scotland (Rodger Leven, Loch in values normal of 90% were levels oxygen dissolved However, 1998a). Scotland, when moribund brown trout were found after lysis of an bloom (Rodger found for gill and liver damage that was similar to damage observed in fish treated with microcystins (Phillips Immersion trials using concentrations found in eutrophic environments did not cause cyanobacterial cells similar to those of aqueous extracts of deaths (Bury the hepatotoxic the biochemical mechanism underlying death following exposure to cyanobacterial the biochemical mechanism underlying blooms, has yet to be established (Bury et al et (Tencalla malfunction liver in display a stress response (Bury health, or (3) toxic compounds affect fish-gill ion transport by activities in inhibiting the plasma membranes ATPase of the branchial epithelium (Gaete Bury exposed to microcystins via a number of routes, including intraperitoneal injection, oral ingestion (fed LANDSBERG REVIEWS IN FISHERIES SCIENCE 1985; Sugaya 1985; 1998a, 1998b). In general, fish are microcystin solutions (Phillips insensitive to short-term immersion in toxic Tencalla Microcystis 1994; Keshavanath Carbis no significant clinical or histological changes over a 96-h period (Tencalla 1994). However, Peñaloza However, 1994). from sensitive also are fish that possible is it above), (see zooplankton with demonstrated by produced are that compounds bioactive additional to

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 + ., ., ., et al et al et et al ., 1994; , and K + aeruginosa et al . M , Na (see section on 2+ ., 1991; Carbis 1991; ., ., 1997). Lethal doses Lethal 1997). ., ., 1994; Tencalla 1994; ., , microcystin damaged microcystin , g microcystin/kg body ., 1994; Zambrano and µ et al et et al et ., 1996a, 1996b). In some ., 1996a), and death was et al et et al et al et al ., 1996b, 1998a), again providing Cyprinus carpio Cyprinus ., 1996a; Bury 1996a; ., ., 1997). One suggested mechanism for et al ., 1995), there are inconclusive findings ., 1985; Råbergh Råbergh 1985; ., 251 includes MGDG and DGDG (Bury et al et et al Chrysochromulina polylepis et al et al et ., 1991; Kotak . (1998a) could find no evidence that inhibition and ., 1991). Unlike in mammals, hemorrhage of the et al et al . (1998a) suggested that lipids, rather than MC-LR, et al aeruginosa . et al ATPase (Bury M ., 1994; Kotak 1994; ., produces additional bioactive compounds that can harm + ., 1994; Bury /K ., 1994). Following intraperitoneal injection, the main uptake + et al et et al et al ., 1996a). In common carp, common In 1996a). ., ., 1995, 1998a; Carbis Microcystis Karenia mikimotoi et al et ., 1996a). et al ., 1998b), and toxicity is manifested as massive hepatic necrosis (Råbergh et al et al ., 1991; Tencalla 1991; ., Although the skin epithelia of freshwater fish form a barrier to microcystin equivalent microcystins of amount an with gavaged when h 96 within died Trout inhibited fish-gill Na of ionic exchange in tilapia gills was due to microcystin activity, but rather found that it was due to the action of fatty acids. Fatty acids produced by interfere with gill basolateral membrane ion-extrusion mechanisms Interestingly, bloom. and cyanobacterial a of thus lysis after seen may deaths fish the to contribute the fatty acid profile of Bury al et of MC-LR induced a total loss of parenchymal architecture of the liver and necrosis of the hepatocytes (Råbergh evidence that aquatic organisms. Bury Canelo, 1996). However, Bury 1998a) (Table 12), both of which are implicated in the hemolytic and ichthyotoxic properties of route of microcystin in salmonids is the gastrointestinal tract (Tencalla liver was rare in fish (Råbergh ATPases, which are the major enzymes involved in the transport of ions. Microcystins ions. of transport the in involved enzymes major the are which ATPases, dephospho- aspartic the inhibit and proteins phosphorylated of hydrolysis the block rylation step of the sodium pump enzymes (Gaete exposure exposure to microcystin causes tissue (Phillips neurons optic and cerebellar damage in the gills, liver, Kotak 1996b; kidneys, and the branchial epithelium, so impaired health or death may result from respiratory failure or an imbalance in ionic homeostatis (Carbis treatments, exposure of common carp to microcystin via gavage caused changes, indicating mild wild liver in occur to damage unlikely is toxicity and acute that indicated exposure changes gavage from results in the cation-anion equilibrium. The carp populations, but chronic poisoning may follow repeated sublethal exposures (Carbis transport (Tencalla (Gaete exposure of routes possible as gills the regarding 1994; Bury hemolysins). to that which passed through the (i.e., 1440 mg gills freeze-dried algae/kg body within weight or 6600 18 h in weight) (Tencalla aqueous exposure tests Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS toxicity is that microcystin inhibits the enzymes of the gill microsomal fraction that are involved in ion pumps and exchanges. Fish may die during a bloom because microcystins affect the ability of the gill to medium. Freshwater fish maintain a maintain large ion concentration gradient between their homeostasis of the from the osmosis by internal lost Ions environment. aquatic dilute the and fluids extracellular branchial circulation into the water are replaced medium through active by transport sites on the gill epithelium. Microcystin inhibition active ion uptake from the of the transport processes exchanging calcium or sodium via the gill chloride cells can lead to osmoregulatory disruption. MC-LR inhibited the Ca

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al ., 1993; ., ., 1996a, Microcystis et al et et al — decrease g/kg/day) for µ concentrations – was begun in the ., 1995), microcystins was 13% of the total (Beveridge and Cl and

+ et al g/l) was confirmed in the µ increase aeruginosa aeruginosa . . M M Microcystis Hypophthalmichthys molitrix 252 ., 1995, 1996a). Some fish species may be were examined, hepatic lesions were found in et al g/g microcystins were found in the viscera and liver, ., 1997) and frogs (Gromov µ et al appeared. Microcystin (12 and silver carp,

, ., 1994). aeruginosa ., 1998); and to influence growth rates, stress responses, and ionic et al g/g and 3 to 15 . ) (Oberemm ., 1996a) rather than death by hypovolumic shock, as reported for mice µ M Microcystis aeruginosa et al et al In August 1996, routine monitoring for ., 1997), which confirmed previous experimental findings (Carbis When monthly samples of common carp that had been exposed to a natural In addition to the association of microcystins with acute mortalities, a number Danio rerio grazing activity as concentrations of toxic of concentrations as activity grazing Keshavanath Jacarepaguá Lagoon, Rio de Janeiro, Brazil, with bimonthly collections sediments, of and fish water, samples. By the first week of November, bloom an atypical of surface plankton mass). By the end of November, microcystins were detected in Ranges 1997. the January, by liver ovaries the in and species) (unspecified fish of viscera and of 1 to 150 susceptible to toxicity via ingestion of microcystin, but others — Oreochromis such niloticus as tilapia, concentrations microcystin that demonstrated findings significant More respectively. in fish muscle were close to or above the recommended limit (0.04 1996b). Previous investigations indicated that branchial injury is more likely to occur to likely more is injury branchial that indicated investigations Previous 1996b). particularly not are carp because bloodstream, the to access have microcystins when susceptible to branchial injury from microcystins dissolved in water (Carbis during a 4-h exposure (Bury plankton samples (the relative abundance of LANDSBERG considered to be by general hepatic failure caused by massive hepatocyte necrosis (Kotak REVIEWS IN FISHERIES SCIENCE (Carmichael, 1992). When common carp were gavaged with a single sublethal bolus sublethal single a with gavaged were carp common When 1992). (Carmichael, dose of microcystin, damage to the renal proximal tubular cells of the kidney and to the hepatocytes of the liver was after 12 hours (Fischer in the intestinal mucosa were reported pathological changes seen as early as 1 h and after Dietrich, 2000). treatment, while bloom of more than 50% of the fish from February to April. This pathology was considered to be consistent with the possibility that microcystins were absorbed through the intestine. However, degenerative changes in the branchial epithelium of carp were also consistent with toxic injury caused by microcystins. The branchial injury was accompanied by a reduction in serum sodium and chloride concentrations (Carbis et al 1996a). Therefore, it appears that the differences noted in the effects of microcystin which by route habitat, species, the on depend may gills the via fish on and the active compounds to which fish reaches the gills (i.e., internal or external), are exposed (pure microcystin or other bioactive compounds). of sublethal effects on poikilotherms have also been ( demonstrated. In zebrafish mortali- high and malformations gross produce to shown experimentally been have ties during embryonic development, to affect behavior and reproductive (Baganz success increasing include solutions microcystin in immersion of effects Sublethal regulation. Na plasma decreasing and levels glucose and cortisol plasma human consumption. The confirmation 1998; concern about potential human health implications (Magalhães and Azevedo, of microcystins in fish fillets warrants

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . ., ., ., et al et al et al steel- × ., 1988; . submit- ., 1998). et al aeruginosa . et al et al M ., 1994). As with , chinook salmon, et al ., 1993; Williams et al been associated with several Salmo salar

has . Apparently, there was a significant influx 253 ., 1995), and MC-LR was detected in copepods spumigena in Lake Alexandrina, Australia, was the first . N et al Microcystis , and Donaldson steelhead trout (rainbow trout ., 1997a). ., 1988; Kent, 1990; Andersen , along the Pacific Northwest since 1986 (Kent et al et al ., 1989; Baker and Humpage, 1994; Jones ., 1999). Also in the summer of 1995 in Jehay, Belgium, mortalities mykiss ., 2001) and suggests that fish for consumption should be tested for ., 1993; Williams ., 1993). Latest findings indicate that a zooplankton vector is involved is vector zooplankton a that indicate findings Latest 1993). ., et al . . Lyophilized algal cell powder from Shin-ike contained large amounts large contained Shin-ike from powder cell algal Lyophilized . Nodularia spumigena O et al et al et al et al et During the summer of 1995, about 20 spot-billed ducks died from unnatural In several reports, ill health in salmonids from what appears to be chronic aeruginosa . 1997b). Although the etiology of NLD has not been definitively proven, field and experimental evidence strongly suggests that NLD is caused by exposure of fish to area this in microcystins of bioorigin the However, microcystins. occurring naturally is not known and cyanobacterial blooms are typically uncommon (Horner bloom from microcystins of presence the confirmed assays hepatocyte rat and HPLC material, but animal tissues were apparently not examined (Wirsing 1997). NLD is characterized by severe necrosis and megalocytosis of the liver and has been observed in netpenned Atlantic salmon, ted, cited in Williams of microcystins that showed acute toxicity in mice, whereas algal samples from a necropsies Bird toxic. acutely not were mortalities) bird no (with pond neighboring showed necrotic livers that were severely jaundiced microcystin toxicity. (dark-green), The suggestive results of suggested that the cause sudden appearance of of toxic the deaths was the of untreated sewage into the pond following the Hanshinn earthquake of January 1995. Eutrophic conditions likely contributed to the (Matsunaga development of the bloom of about 30 ducks and herons coincided with a massive bloom of causes in a pond (Shin-ike) in Nishinomiya, Hyogo Prefecture, Japan. The suspected The Japan. Prefecture, Hyogo Nishinomiya, in (Shin-ike) pond a in causes cause was the sudden M appearance of a cyanobacterial bloom identified as Oncorhynchus tshawytscha and crab larvae collected from NLD-affected salmon farms (McReady head trout), of analysis bioassay phosphatase protein chromatography-linked Liquid 1990). Kent, liver extracts taken from Atlantic salmon affected by NLD showed the presence of from indistinguishable chromatographically was that inhibitor phosphatase protein a the recreated salmon Atlantic healthy into MC-LR of injection Intraperitoneal MC-LR. pathology of NLD, which included (Andersen diffuse necrosis and hepatic megalocytosis (Andersen Q. NODULARIN A bloom of livestock, canine, and wildlife mortality events (Table 7), principally occurring in brackish waters in Australia, New Zealand, and the Baltic Sea (Carmichael documented report of an animal mortality (Francis, 1878). event Since associated that with time, cyanobacteria 1988; Sivonen Magalhães Magalhães toxins in waterbodies known for microcystins. exposure to microcystins has been deemed to be a new syndrome, “netpen liver disease” (NLD) (Kent Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 and ., 1990; ., bloom , tested ., 1994). ., ., 1999). initially blooms was the Daphnia spumigena et al et et al et et al . Anabaenopsis N sp. than on the on than sp. ., 1988; Sivonen, Nodularia pulicaria sphaerocarpa Nodularia Nodularia

. Mytilus edulis et al N by mysid shrimp Nodularia Daphnia ., 1994). Several nodularins ., 1991). Sublethal effects such ., 1999). Both in experimental et al spumigena et al et al . g/ml) and the cladoceran N µ ., 1994). In rodents, nodularin induces ., 2001). et al 254 et al bloom ended. It was recommended that edible that recommended was It ended. bloom has been determined to produce nodularin (Bolch nodularin produce to determined been has g/ml) (De Mott ., 2000), but when the copepod had been fed toxic µ ., 1988). Nodularin promotes liver tumors (Yoshizawa tumors liver promotes Nodularin 1988). ., , 0.52 to 1.25 are toxic (Bolch (Engström (Engström 50 et al et al et Nodularia g/ml) were the most sensitive; , 14.1 bloom in Peel-Harvey Inlet, western Australia, were shown were Australia, western Inlet, Peel-Harvey in bloom µ 50 spumigena . Some tissues from edible blue mussels, . N (48-h LC Nodularia fed less actively on the toxic strains of strains toxic the on actively less fed , 3.9 50 , were also present in the bloom, so it is unclear to what extent ., 1991; Carmichael 1992; Rinehart ., 1992). Mussels that were found to be ingesting spumigena ., 1983). . ., 1993), but the biogenic origin of these toxins is currently unknown. et al Nodularia N sp. for 3 to 5 days, their rates of mortality increased (Koski alone was responsible for the fish kills (Devidze, 1998). Fish and crabs et al (48-h LC et al et al , when compared with grazing on nontoxic strains of ., 1999), which is a cyclic pentapeptide hepatotoxin that is, like microcystin, a A few studies have demonstrated effects of nodularin on zooplankton. Acute Thus far, only far, Thus There are potential human health risks associated with shellfish that have Few reports have associated nodularin with aquatic animal mortalities, although mortalities, animal aquatic with nodularin associated have reports Few ., 1990) and is also considered to be a direct liver carcinogen (Ohta carcinogen liver direct a be to considered also is and 1990) ., Rhizosolenia Not all strains of enlarged hemorrhagic livers, centrilobular necrosis, lysis of hepatocytes, and death (Runnegar h 2 to 1 within al et animals and based on observations in field reports, the toxicity and pathogenicity of nodularin is very similar to that of microcystin (Runnegar 1996). Along with microcystin, nodularin has also been found in marine (Chen mussels toxicity experiments with purified toxins showed that four species of zooplankton differed markedly in their physiological Diaptomus sensitivity birgei to nodularin. The hyalina copepod least sensitive (48-h LC et al (Yoshizawa tumor-promoter and inhibitor 2A and 1 phosphatase protein Honkanen as reduced feeding and fecundity have also been noted (Table 10). The copepod affinis Eurytemora nontoxic strains (Engström LANDSBERGother cyanobacteria, most of these nodularin by terrestrial events animals (Table 7). are caused by the consumption of REVIEWS IN FISHERIES SCIENCE have now been characterized (Rinehart Nodularia Similarly, there was reduced grazing on toxic mixta as such plankton, other that noted was it However, 7). (Table blooms or Nodularia have been reported to avoid the waters of the estuaries during (Potter consumed an during (Falconer showed little sign of ill health or exposure, slight hepatopancreatic pathology. tubular lesions However, developed, and after extensive secre- several months of tion of algal pigment by diapedesis of engorged hematocytes was evident. It was suggested that long-term effects on the health of mussels should be investigated to be toxic to mice. Mice showed characteristic hepatic hemorrhage and hepatocyte and hemorrhage hepatic characteristic showed Mice mice. to toxic be to any in result not did tissues nonintestinal of extracts of Administration degeneration. mouse toxicity. Only the gastrointestinal tracts in the mussels retained toxicity; this the after declined toxicity Aphanizomenon flos-aquae recently, fish mortalities in the Black Sea, Georgia, coincided with a during consumption human for collected be not should mussels

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., et al et al ., 1995; Akashiwo , et al Heterosigma , ., 1999a, 1999b, Vibrio alginolyticus Vibrio et al is the most toxic ROS ., 1992; Yang Cochlodinium polykrikoides Cochlodinium ., 1989, 1991; Tanaka ., 1999a), but to what extent et al marina et al . has many small and large verru- C et al Prorocentrum micans ., 1993). All raphidophytes tested from Orielton Lagoon, Tasmania, et al ) Hansen and Moestrup (Daugjberg (Kim antiqua, Fibrocapsa japonica Fibrocapsa antiqua, . 255 spumigena ., 1999b). The production of small quantities of . and the dinoflagellate the and N et al ., 1994). Low concentrations of nodularin were also Chattonella marina et al ., 1992a, 1992b, 1994, 1995, 1997; Kim Chattonella marina, C marina, Chattonella ., 1999b). Alexandrium tamarense et al Gymnodinium sanguineum ., 1996a, 1996b), although other organisms may also be affected Olisthodiscus luteus Olisthodiscus et al (= , but additional information was not provided (A. Goodsell, personal et al ., 1992b, 1997; Kim , and , et al ., 2000), and ., 1992). The hydroxyl radical generated by Thus far in the wild or in aquaculture, raphidophytes have, with the exception the with have, raphidophytes aquaculture, in or wild the in far Thus elicited histological alterations in the livers and pyloric Salmo salar ceca of Atlantic salmon, communication, in Jones detected recently in livers of flounder and cod caught in the Baltic Sea (Sipia merits animals aquatic in exposure nodularin of effects chronic potential The 2001). further investigation. (Langdon, (Langdon, 1990). Extracts from R. REACTIVE OXYGENraphidophytes The SPECIESakashiwo known (Kim ciform protrusions on the cell surface, and these are production of superoxide probably anions (Shimada involved in the has also been detected in sanguinea et al Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS been also has ROS An unclear. currently is significant be may concentrations these generate reactive oxygen species (ROS) (e.g., superoxide anions, hydroxyl radicals, species (ROS) (e.g., superoxide generate reactive oxygen singlet oxygen, and hydrogen peroxide) (Shimada 1992, 1994; Oda 2000b; Twiner and Trick, 2000), which can have adverse effects on aquatic organ- isms. During , superoxide and hydrogen peroxide are generated as a result of the photoreduction of dioxygen that occurs in illuminated the chloroplasts disturbs which production, ROS trigger can stress Environmental 1951). (Mehler, steady-state balance of prooxidants and antioxidants (Okamoto 1998). The and toxicity of Colepicolo, oxygen radicals to biological systems has been mented (Fridovich, 1978; Cunningham and Capone, 1992). At the cellular well level, the docu- damaging effects of ROS include denaturation of polysaccharides, DNA enzymes, damage depolymerization that results of in genetic 1998). mutation Colepicolo, and (Okamoto death (Cunningham or injury cellular severe and and 1992), Capone, Lipid molecules are susceptible to removal oxygen-radical-induced of hydrogen peroxidation radicals, and such via damage can lead the to destruction of mem- brane integrity (Cunningham and microalgae appears Capone, to principally affect fish (Tanaka 1992). The production Ishimatsu of ROS by (Tables 10 and 11). of one case (Table 5) attributable be (and may one 5) (Table unconfirmed mortalities case, fish Some see ichthyotoxic. Section only T.3) be to been mented docu- to the production of brevetoxin-like compounds (see brevetoxins), to ROS, or to a combination of both. ROS contribute to pathological changes in the gills of fish (see below), and the brevetoxin-like neurotoxins impair cardiac function (Endo et al bacterium marine the of growth the inhibited ROS produced that (Oda

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et al Gentien,

., 1992b; ., 1996a). ., , Chattonella Heterosigma et al et ., 1985, 1989; et al spp. (Ishimatsu spp. et al Chattonella ., 1989; Oda ., 1998; Munday and Hallegraeff, and Munday 1998; ., et al Gymnodinium aureolum Gymnodinium depends on the growth phase; the phase; growth the on depends cells/ml were recorded. were cells/ml ., 1995). 2 et al et (as ., 1985; Toyoshima spp., a series of histopathological and spp., a series of histopathological 10 et al × et al marina g per 100 g of tissue. A likely scenario for 256 . µ ., 1997). A decrease in the oxygen partial C ., 1992). Yellowtail were exposed to two strains et al Chattonella , in Boston Bay, Australia, coincided with a bloom spp. is that fish die because of the direct toxic effect toxic direct the of because die fish that is spp. et al ., 1990; Hallegraeff 1990; ., Karenia mikimotoi mikimotoi Karenia , worth more than 7 billion yen (Okaichi, 1989; Imai et al et . Counts of up to 6.6 to up of Counts . ., 1983, 1991; Endo Chattonella Fish were observed to be distressed, swimming in a haphazard

spp. et al ., 1996a; Hishida that had been subjected to differing light-dark regimes, filtering, marina

Thunnus maccoyi red tide caused a mass mortality of about 14 million cultured yellowtail, cultured million 14 about of mortality mass a caused tide red spp. have caused significant fish mortalities in the Far East, particularly ., 1986; Kobayashi, 1989; Kobayashi et al red tides totaled 2 billion yen from 1972 to 1987 (Honjo, 1993). Blooms marina et al . antiqua ., 1990, 1991; Tsuchiyama In April-May 1996 a mortality of an estimated 1700 tonnes of cultured southern C . Chattonella C akashiwo 2. Heterosigma akashiwoIn the Seto Inland Sea, Japan, the economic damage caused by cultures tested positive for brevetoxin-like compounds (0.03pg PbTx-3 equivalent/ cell in log phase). manner on the surface, and in some cases, gasping. No other finfish or shellfish in Gill fish. the of gills the on present was mucus Excess dead. reported were area the pathology showed marked epithelial swelling and separation Breve-like toxins were detected in the livers of the tuna by using a sodium-channel of the epithelium. receptor binding assay with up to 142 by caused mortality of oxygen radicals on the (Ishimatsu brevetoxins gills in combination with the 1998). cardiotoxic effects of 1. Chattonella Chattonella Sea, Inland Seto eastern Harima-Nada, in 1972, of summer the In 5). (Table Japan in a LANDSBERGin presumptively identified REVIEWS IN FISHERIES SCIENCE Seriola quinqueradiata 1998). 1998). When fish are exposed to 1998). When fish are and cells goblet mucus the of stripping The gills. the in occur changes physiological the mucus coat, along with the structural alteration in the chloride cells, leads impaired to osmoregulation, the development of reduced oxygen edema transfer, which results in in asphyxiation (Matsusato and Kobayashi, the gill lamellae, 1974; Shimada and Sakai Ishimatsu pressure of arterial blood to exposure after and fish in observed changes an physiological increase in plasma catecholamines are et al early of highest the agitation; mechanical by disturbance or freezing, disruption, mechanical fish mortality (19 out experimental various the by affected significantly was of radicals oxygen of production of 20) and production radical occurred oxygen between correlation clear in a was there and the treatments, presence of intact (Ishimatsu fish to toxic not were cells. filtrate cell-free or cells Dead toxicity. The by radicals oxygen of generation The rate of superoxide and hydrogen exponential-growing phase and subsequently decreased peroxide to one-fifth of its maximal generation was highest during level in the the stationary-growth phase (Oda bluefin tuna, of

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 cells/ cells/ Salmo cells/ 4 5 5 ., 1995). ., 10 10 ., 1996c). ) cultures 10 × × × passes over passes et al et et al luteus . O was associated with associated was

) akashiwo . (as H and Atlantic salmon, salmon, Atlantic and

, akashiwo cells/ml (Khan 5 , were exposed to cultures of . cf. . akashiwo . 10 H H reach several million per l, and some × (Uye and Takamatsu, 1990). That ROS That 1990). Takamatsu, and (Uye than others, such as the chinook salmon, chinook the as such others, than was fed different plankton species, and ) have also been associated with fish kills been associated ) have also ) has been reported to affect numerous 257 Pagrus major luteus akashiwo . (originally as (originally . Oncorhynchus mykiss Oncorhynchus H O Heterosigma (as Olisthodiscus luteus Olisthodiscus akashiwo cells/ml were recorded. Histopathology of affected salmon . generates reactive oxygen intermediates that probably first Olisthodiscus luteus Olisthodiscus 3 H 10 obtained from an ongoing 1995 red tide that was responsible for responsible was that tide red 1995 ongoing an from obtained × (Taylor and Haigh, 1993). In early January 1989, a dense phy- (or as Pseudodiaptomus marinus probably first attack the membranes of the epithelial cells of the cultures were disturbed, the release of superoxide increased, thus , no abnormal behavior was noted at a cell density of 3.4 akashiwo are more sensitive to sensitive more are

. H akashiwo — cells/ml. When fish were experimentally exposed to a density of 3.0 3.0 of density a to exposed experimentally were fish When cells/ml. . ., 1990). Heterosigma akashiwo 6 When juvenile red seabream, akashiwo akashiwo tschawytscha akashiwo H . . . . 10 The copepod extremely high mortalities were observed to exposed (32% was copepod by the second day) when were involved this in the mortality still needs to be verified. S. SPECIES WITH MULTIPLEThere are numerous species or strains TOXINS of produce microalgae multiple that toxins have or been bioactive shown compounds to (Table 1). In some cases, the Heterosigma akashiwo attack the membranes of the epithelial cells of the secondary gill lamellae. When H containing water when occurs that situation the mimicking the secondary lamellae of finfish. The reactive oxygen intermediates generated by H produced teratologies or mortality in echinoderm eggs and larvae (Wilson, 1981). 3. Olisthodiscus luteus secondary lamellae, causing lipid peroxidation damage, inactivation of ATPase of the of ATPase of inactivation damage, peroxidation lipid causing lamellae, secondary (Yang lamellae gill the in edema and membrane, plasma cell epithelial 10 (Tables sublethally and lethally both and zooplankton of species Egloff, 1986 and references therein). Undefined and 11) (Verity and Stoecker, 1982; cell concentrations or cell-free filtrate from of including vasculature, and epithelium branchial the of changes degenerative showed swelling of the respiratory epithelium, mucus cells by edema. Impairment of respiratory and secondary epithelium from the pillar discharge, and separation (Chang death of of cause the be to concluded was the gills the of function osmoregulatory et al H 1.2 exceeded density cell the when died eventually and paralyzed were fish but ml, × of ml wide-scale fish kills, these fish showed a transient but not fatal paralysis. Fish were dying in the wild at concentrations of 1.0 Volume 10 (Issue #2) 2002 2 than more of bloom a 1986, EFFECTS In OF 5). ALGAL (Table BLOOMS 1976 ON since AQUATIC ORGANISMS Columbia British in ml led to losses to the salmonid industry of at least $2.5 million, which at the time represented a third of the value normally produced (Taylor, 1993). Penned salmo- nids die when concentrations of O of bloom toplankton species — such as rainbow trout, rainbow as such — species salar concentra- cell mortality, the During 5). (Table Zealand New in kills salmon chinook tions of up to 2

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., et al cells/ , were 2 Seriola type ’78) type Tubastrea 10 × Cochlodinium , Alexandrium monilatum Crassostrea virginica Cochlodinium polykrikoides sp. red tide (4 to 5 heterolobatum , and the azooxanthellate coral azooxanthellate the and , . C spp. have been noted to coincide with 258 (= Cochlodinium damicornis sp., occurred from Weitou Bay to Quanzhou Bay, . P , Cochlodinium polykrikoides . (97% by relative abundance); C cells/ml, calcium uptake was negligible, and the larvae died 4 spp. cells/ml, calcium uptake rate in the oyster larvae was depressed was larvae oyster the in rate uptake calcium cells/ml, sp., were demonstrated to be ichthyotoxic in experimental 2 10 Cochlodinium × , showed unusual behavior, possibly because of toxin exposure 10 × species, catenatum . C Pocillopora elegans Pocillopora . At Uva Island, numerous pocilloporid corals had bleached and were ., 1993a). Although recognized to be a potential risk to aquaculture facilities in Japan, two Japan, in facilities aquaculture to risk potential a be to recognized Although Cultured straight-hinge larvae of eastern oysters, et al In late August 1985, a small-scale Cochlodinium to approximately 10% of that of the control, and high mortality occurred. At densities At occurred. mortality high and control, the of that of 10% approximately to greater than 1.0 promptly (Ho and Zubkoff, 1979). Cochlodinium but after the water was filtered they were able to reach apparently normal umbo stage. In experimental exposures, when densities of the dinoflagellates were 5 proximately ap- and 1. Dinoflagellates a. Cochlodinium Occasionally, blooms of mortalities of aquatic organisms (Table 2). In 1985, coral reefs at Cañon Island, Costa Island, Cañon at reefs coral 1985, In 2). (Table organisms aquatic of mortalities Rica, and Uva Island, Panama, were affected during a dinoflagellate bloom domi- nated by were m 3 to 2 of depths to waters surface Island, Cañon At present. also was (1.4%) bloom. the by produced presumably was which foam, viscous by red-yellow colored Numerous fish were affected, with pomacentrids, and hundreds tetraodontids, as of well as affected, scarids, also were corals hermit shallow-water Some beach. the crabs, balistids, on dead found gastropods brachyuran acanthurids, crabs, and especially coccinea sloughing tissues. It was suggested that the mortality combination was of possibly toxicity, caused oxygen by depletion, a and smothering during the blooms. by Adhesion of mucus to the mucus polyps, which interfered with produced polyp expansion, was deemed the most likely cause of coral mortality (Guzmán ml) occurred in Harima-Nada, quinqueradiata Japan, where cultured yellowtails, (Yuki and Yoshimatsu, 1989). During mid-June 1990, an unusual a red toxic bloom of tide event, China. A large variety of fish, shellfish, and species were killed (Table 2). Interestingly, crustaceans such as crabs, lobsters, and (Qi shrimp were not affected initially deformed when cultured in water containing LANDSBERG coincidental occurrence of a particular species for which toxins have already been make to researchers led have may event mortality aquatic an of site the at described assumptions about the cause of the mortality. However, more and more examples are being found in which other minor (or less well studied) toxins are now being Many involved. are compounds which unclear often is it cases, some In implicated. of the species producing multiple reference text its and 1 Table to refer should toxins reader the sections; other in discussed and bioactive compounds have discussed. are toxins been or species particular article this in where determine to column REVIEWS IN FISHERIES SCIENCE 1990).

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 × . When mortalities

Paralichthys Oplegnathus sp. cells/ml; polykrikoides . C Pagrus major , beakperch, , Cochlodinium type ’78) and were charac- 3 ., 1985; Yuki and Yoshimatsu, 10 -hydroxyneosaxitoxin sulfate (Ic- sulfate -hydroxyneosaxitoxin × , were exposed to 1.74 to 6.51 . Hemolytic activity produced by α type ’78 Yatushiro). Ichthyotoxicity Yatushiro). ’78 type et al , to the effects of caused fisheries losses amounting to ., 1999a). The production of bioactive Pagrus major Pagrus and their potential roles in , lipid-induced peroxidation was linearly was peroxidation lipid-induced , to 1.79 1 Cochlodinium ., 1999a). When live flatfish -sulfo-11 et al 259 N 10 × et al (as polykrikoides . Cochlodinium C 1.0

polykrikoides . polykrikoides C . ., 1999a). Because oxygen radicals are generated from generated are radicals oxygen Because 1999a). ., polykrikoides , red seabream, red , C Leiognathus Leiognathus nuchalis . sp. (as sp. C Malakichthys wakiyae . (1999a) suggested that it would be reasonable to assume to reasonable be would it that suggested (1999a) . et al et polykrikoides (hemolytic unit) was estimated to be 0.5 HU per mg in . et al et 50 C cells/ml and g), they died from respiratory paralysis. The lethal dose was 2 to g), they died from respiratory paralysis. was shown recently to be associated with the fatty acids µ , Kim , . The reaction of fish to the hemagglutinin fraction was similar to that to similar was fraction hemagglutinin the to fish of reaction The . epimer (epi-Ic-1). Intraperitoneal injection of Ic-1 into mice resulted Cochlodinium β were exposed to exposed were , and seaperch, polykrikoides . ., 1999a). The increased susceptibility of pelagic fish such as black scraper, More recently, blooms of C polykrikoides

3 polykrikoides . . that lipid peroxidation of permeabil- fish solute increased to gill linked been has tissue peroxidation lipid Further, is oxidation. a result of oxygen-radical-dependent ity of the vesicle cells gill in membranes changes functional and structural Because that gills. the in edema control for account swelling and lysis, transfer which oxygen for capability the reduce could also may effects these ROS, then by induced are in the gills. Additionally, because hydroxyl radicals also induce mucus secretion in fish gills, the combination of effects induced by ROS will result in a fish kill (Kim et al septentrionalis fasciatus C were investigated (Lee, 1996; Kim olivaceus proportional to algal cell density. Edema and histopathological changes in the gills (Kim observed also were terized as a zinc complex of carbomoyl- of complex zinc a as terized $95.5 million in Korea (Kim, 1998; Kim compounds such as ROS in 4 mg/kg (Onoue and Nozawa, 1989a). Two unique paralytic shellfish poisons were 4 mg/kg (Onoue and Nozawa, 1989a). also separated from 1) and its 11 in signs characteristic of PSP: ataxia, convulsion, and respiratory paralysis. When exposed to 150-200 ppm Ic-1 in seawater, juvenile red sea bream displayed loss of was Ic-1 hydrolysis, acid mild On arrest. respiratory and breathing, labored balance, converted into GTX1 and epi-Ic-1 into GTX4 (Onoue and Nozawa, 1989b). To my knowledge, since these initial studies there identifying have PSP-like toxins have from been no reports further (C20:5n3)(15.3%) acid eicospentaenoic and (25.3%) (C22:6n3) docosahexaenoicacid (Lee, 1996) (Table 12). Cochlodinium of fish to the hemolytic fraction. When mice were injected i.v. with the hemagglu- tinin fraction (20 assays. Juvenile slipmouths, 10 after 48 h, mortalities were 20 to 40% and 80%, respectively. Seawater filtered from the cultures had no effect on the fish (Onoue exposed to 0.02% of the neurotoxic fraction, the fish behaved as though they were observed. was bodies their of whitening or change color marked a and anesthetized, Labored breathing and respiratory arrest resulted in symptoms: following the death developed fish fraction, in hemolytic the of 8 0.02% to exposed to 10 min. When violent convulsions, loss of balance, arrest. Fish died in 20 in the gill lamellae, and respiratory excess mucus production labored breathing, edema, hemorrhaging, to 30 min. The HD C Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1989). Three toxic fractions — neurotoxic, hemolytic, and hemagglutinative — were — hemagglutinative and hemolytic, neurotoxic, — fractions toxic Three 1989). from isolated of the three fractions was tested on juvenile red sea bream,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., et et al et , and cells/ 3 10 × ., 2000b) with Oncorhynchus ., 1971); it has et al et al ., 2000). toxins (Gerwick toxins et al brine shrimp. , are included]). Most of the of Most included]). are ) caused substantial mortali- substantial caused ) have been experimentally Paralichthys Paralichthys olivaceus Artemia is common in marine tropical cells/ml, fish showed signs of 3 Artemia Lyngbya 10 Lyngbya ., 1999; Milligan × , and chinook salmon, polykrikoides et al . C 260 ., 2000a). et al Lyngbya majuscula Salmo salar sp. (likely sp. cells/ml fish stopped feeding and when concentrations when and feeding stopped fish cells/ml (Kim 2 , the activities of the ion-transport enzymes, carbonic , was recently demonstrated. When experimentally ex- cells/ml mortalities occurred. Experimental field bioassays field Experimental occurred. mortalities cells/ml 3 10 ., 1997). ., 2001). × -ATPase, in the gills were significantly decreased in relation 10 + ) was also measured in red sea bream and flounder when × 2 than from any other microalgal species discussed herein (Table than from any other microalgal species et al et al /K was recently associated with lyngbyatoxin-a in Madagascar O + p

, polykrikoides Cochlodinium polykrikoides . blooms in southeastern Queensland, Australia, include localized is a problem for aquatic animals such as marine mammals or sea turtles. sea or mammals marine as such animals aquatic for problem a is C . C Sebastes inermis function in nature as mechanisms that protect this alga from predation by , in British Columbia that resulted in an economic loss of CAN $2 million. Several of the compounds isolated from Blooms of Blooms ., 1995a; Thacker majuscula majuscula . . A fatal human intoxication caused by the consumption of meat from a green turtle, Chelonia mydas posed to rockfish, anhydrase and Na toxic compounds have been verified through experimental study, and it is unclear to what extent these compounds affect aquatic organisms in the natural environ- ment. It has been speculated that many of the bioactive metabolites produced by Lyngbya a variety of groups (e.g., crustaceans, herbivorous fish, and gastropods) (Orjala al areas and has principally been associated with contact skin dermatitis in humans (Grauer and Arnold, 1961; Moikeha and Chu, 1971; Moikeha caused numerous outbreaks of field and dermatitis mammals small with in experimentation laboratory from the Apart 1984). (Moore, Hawaiian Islands and in Japan observations in humans, it L is unclear if contact dermatitis associated with (Yasumoto, 1998; Yasumoto and Satake, 1998). Observed L ecological impacts of seagrass loss and poor harvests of crab and fish (O’Neil demonstrated to be toxic to fish, molluscs, and potent are tanikolide and lyngbyabellin, kalkipyrone, There have probably been more Lyngbya majuscula potentially toxic compounds by produced be to known compounds all [not 1 identified from 2. Cyanobacteria a. Lyngbya majuscula to increasing algal cell density and exposure time. A drop in blood pH and oxygen cell density and exposure time. A drop to increasing algal partial pressure ( exposed to 2 than more reached with salmon smolts demonstrated lethality after 120 min exposure with over 90% 10.8 to 2.7 from varied concentrations cell when min 500 after mortality column. water the in stay to inability an and equilibrium, in loss distress, respiratory Pathological change to the gills inlcuded edema epithelium (Whyte and separation of the lamellar 1994; Graber and Gerwick, 1998; Singh unknown effects on natural zooplankton populations; barbamide is toxic to the LANDSBERGwhen compared to benthic fish such as flounder, REVIEWS IN FISHERIES SCIENCE ties of farmed Atlantic salmon, kisutch 5 exceeded cells When ml. In fish bioassays at densities of above 1

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., ., spp. et al et al = 0.003 resulted 50 g/ml), but tamarense µ . A ., 1999). = 5 ., 1995a). As well catenella et al 50 Alexandrium . A produces a suite of et al ., 1981]) (Orjala et al also produces reactive oxygen , showed that pure [Lin majuscula . g/ml (Orjala and Gerwick, 1996). L µ ., 1984a, 1984b, 1985, Moore 1984; spp. Media from ., 1999a) warrants further investigation further warrants 1999a) ., et al = 100 et al et tamarense . 261 Danio rerio A 100 ., 2000). Carassius Carassius auratus et al Alexandrium at LD ., 2001). Such activity may have significant implica- ., 1995). Supernatants from cultures of et al et al ., 1984a). Unlike okadaic acid, nodularins, and microcystins, and nodularins, acid, okadaic Unlike 1984a). ., spp. have been implicated in natural aquatic mortality events were also toxic to rat primary neuronal cells (Perovic et al et spp. g/ml) – it is exceeded in potency only by the brevetoxins (LD µ Biomphalaria Biomphalaria glabrata lusitanicum Alexandrium . COMPOUNDS A In March 1982, in Kitaura Bay, Japan, a massive bloom of In addition to other bioactive compounds, Malyngamide H was also shown to be toxic to goldfish (LC = 0.05 1. Dinoflagellates g/ml against the freshwater zebrafish species such as hydrogen peroxide (Kim peroxide hydrogen as such species events. mortality aquatic in play may compounds bioactive these role potential the of Kodama, and (Ogata bream sea red and yellowtail cultured of mortality large-scale in and 2000). The recent demonstration that Toxicity experiments on goldfish, (LD plants marine from date to isolated metabolites ichthyotoxic most the among is showed no activity against brine shrimp or molluscs (Orjala as being toxic to brine shrimp, kalkipyrone and hermitamides are toxic to goldfish (Graber and Gerwick, 1998; Tan T. SPECIES WITH UNCHARACTERIZED TOXINSa. Alexandrium OR BIOACTIVESeveral where toxic compounds other than PST were suspected (Tables 1 and 2). PST are not usually excreted into the surrounding medium, and exposed organisms are through generally ingestion. However, a few toxic additional that indicated have studies experimental with conjunction in case events histories of unusual mortality exudates are also produced by and (Ogata erythrocytes mammalian and fish against activity hemolytic had cultures Kodama, 1986; Simonsen mollusc 50 µ tumor promoters: , debromoaplysiatoxin, lyngbyatoxins — all of which bromoaplysiatoxin, have been shown to produce erythema, blisters, and and necrosis when applied to the skin (Fujiki Fujiki and Suganuma, 1996). Like okadaic acid, lyngbyatoxin-a has been demon- strated experimentally to induce papillomas in (Fujiki experiments two-stage mouse carcinogenesis lyngbyatoxins promote tumor growth through the activation of the fact protein The 1996). kinase Suganuma, and (Fujiki inhibition phosphatase protein through not C, that lyngbyatoxin-a was identified recently in role green potential their turtle for considered meat be (Yasumoto, also should 1998) OA, like toxins, these that confirms as tumor-promoters in sea turtle fibropapillomatosis (Landsberg Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1995b). It voltage- of activator was an is brevetoxin, demonstrated like and, cells, granule recently cerebellar rat of that cultures antillatoxin is neurotoxic in primary gated sodium channels (Li tions for natural resources and public health in marine environments.

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , , cells/ minutum minutum minutum . . . A (reported as A A Acanthopagrus ., 1998). When minutum . , were exposed to exposed were , A et al bloom, with concen- minutum toxic . 4 A 10 × minutum . cells/ml, had noticeable edema, , were exposed to cell-free filtrates cell-free to exposed were , A 4 10 × Rhombosolea taparina Rhombosolea on aquatic animals, and several of these of several and animals, aquatic on culture media. Test fish died after 3 to 5 h 262 ) bloom coincided with mortalities of rainbow of mortalities with coincided bloom ) to 2.05 3 minutum . 10 A × tamarense . excavata . A A cells/ml. Prawns and black sea bream, ., 1993) (Lush and Hallegraeff, 1996). In enclosed aquac- 4 , were exposed to toxic or nontoxic cells, or cell extracts and 10 et al × , was associated with a bloom of Hypomesus pretiosus japonicus pretiosus Hypomesus filtrate; control fish and prawns were exposed to pond water from , increased mortalities were observed with increasing concentrations , increased mortalities were observed (reported as (reported catenella by Su . A Chanos chanos , were experimentally exposed to pond water containing either containing water pond to exposed experimentally were , minutum minutum . . In June 1989 in southern Taiwan, a mass mortality of cultured grass prawns, In July 1984 in Tronisvagur fjord in the Faroe Islands, the appearance of an A A tamarense tamarense . . or three adjacent ponds where no bloom had occurred. After 9 of 12 10 prawns h died; the in remaining pond prawn was water, moribund, but it recovered when placed in fresh seawater with strong aeration. Dense accumulations of trations of up to 1 ml or to cell extracts at 5.13 trout and salmon that resulted in 77% four (27 and metric tissue, tons) g of 100 the per fish MU dying. 9000 Although contain to shown were PST for analyzed shellfish the fish behavior and pathology suggest that in human cases of PSP were reported, Two mortality. the for responsible been have may toxins extracellular also, case this with randomly swimming either were trout rainbow started, mortalities the after days or on their sides with their heads above the dorsal and caudal fins out of the water surface. There were acute histopathological changes to the gills, with necrosis and sloughing of the epithelia of the secondary lamellae, pyknotic break- nuclei, with cytoplasmic associated hemorrhage sometimes and hypertrophy, cellular vacuolation, down of blood sinuses. No pathological changes were detected in internal organs or were other infectious agents suspected (Mortensen, 1985). Penaeus monodon A ulture facilities, water-quality problems and environmental stressors can sometimes of effects stress the exacerbate salinity, low temperature, high of combination A involved. been have may stressors low ammonia-N, and a lack of other plankton competitors and predators appeared to be conducive to the development of a heavy schlegeli of in the first and second treatment death occurred within 3 h. Signs of stress included stress of Signs h. 3 within occurred death treatment second and first the in small mouth, and operculum the of heaving swimming, uncontrolled of bursts rapid damage Gill orientation. of loss and body, whole the of movements convulsive rapid was characterized by severe epithelial swelling. In addition, gills from fish exposed No cells. chloride vacuolated and necrosis, fusion, lamellar had medium cell-free to pathological changes were detected in internal organs (Lush milkfish, of each treatment. Fingerlings exposed to 1.5 to 3.0 hyperplasia, and necrosis of the secondary gill lamellae. consumption rate An was increase also observed in in oxygen fish exposed to algal extracts Chou, 2001). (Chen and A LANDSBERGsuspected was it planktivorous, not are fish of species these Because 2). Table 1986; experimental In mortalities. the in role a played have may PST than other toxins that smelt, surf studies, REVIEWS IN FISHERIES SCIENCE from both cells epithelial the Histologically, hypoxia. of signs had and exposure continuous of of the gills were swollen and exfoliated from the pillar cells (Ogata and Kodama, flounders, greenback juvenile When 1986). of medium culture solvent-extracted lipophilic or cell-free, cells, whole

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 (as or to Pagrus g), they g), . (1992) µ et al Alexandrium Alexandrium type 84 k). Karenia by producing . (2000) recently became inactive ., 1996). Both whole Both 1996). ., for the first time in et al et al et sp.) by Endo Alexandrium tamarense Gymnodinium cells per MU. Prawns became 4 , Daugbjerg Favella ehrenbergii (Bagøien (Bagøien (as ., 2001). was shown to inhibit the growth of growth the inhibit to shown was 10 × , eggs were exposed to Euterpina acutifrons et al Gymnodinium has been implicated in fish mortalities both mortalities fish in implicated been has culture medium were lethal to metanauplii to lethal were medium culture 263 species either to the genus minutum , exposed to cell-free filtrate water that had . (as A pulchellum . Gymnodinium pulchellum G minutum . Chlamys farreri A Gymnodinium pulchellum Mytilus edulis . , whereas pure PSP fractions produced no comparable Gymnodinium pulchellum G cells. Alexandrium lusitanicum Alexandrium Artemia . (1998b) reported Gymnodinium pulchellum Gymnodinium ) caused cell death of the ciliate . catenella . et al A cell cultures, hatching rate was 30% when compared with controls, and ., 1993). . When exposed to 0.02% of the neurotoxic fraction, fish behaved as though Culture filtrate of filtrate Culture et al There is additional evidence for the production of exudates by excavatum . tamarense larval survival decreased significantly (Yan b. Gymnodinium pulchellum With the exception of transferred ichthyotoxic Karlodinium Recently, 2). (Table Australia and Japan in particularly wild, the in and aquaculture in Steidinger (Table mortalities invertebrate and fish in species the implicated and America North hemolytic, neurotoxic, — fractions toxic three isolated (1989a) Nozawa and Onoue 2). and hemagglutinative — from suggested that these toxins were oxidized brevetoxins, but no further information contained cells were observed in the dead prawns’ gills or stomach. Toxicity as measured by mouse bioassay was determined to be 1.39 spp. Nauplii of the harpacticoid copepods and adults of mortality. The presence of a fast-acting toxin was proposed (Lush and Hallegraeff, 1996). When Farrer’s scallop, dinoflagellates (Table 11, Blanco and Campos, 1988). and ciliate the of membrane cell the on acted exudate The toxins. extracellular lethal induced continuous ciliary reversal. After some time, the ciliate swelled and subse- quently lysed (Hansen, 1989). Andrasi (1985) also demonstrated the toxins presence in of blue mussels, Ichthyotoxicity of the three fractions was tested on juvenile red sea bream, major anesthetized, and a marked min. color 4 change than or whitening less of in the body death was in observed. resulted arrest respiratory and breathe to labored fish The convulsions; violent exhibited fish fraction, hemolytic the of 0.02% to exposed When produc- mucus excess and hemorrhaging, edema, breathing; labored balance; of loss tion in the gill lamellae; and respiratory arrest. Fish died in 35 to 49 min. Reaction of fish to the hemagglutinin fraction was similar (20 fraction hemagglutinin to the with i.v. injected were mice When fraction. that of fish to the hemolytic died from respiratory paralysis. The lethal dose was 2 to 4 mg/kg. Further studies of the neurotoxins of Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS moribund in 1.5, 3, and 9 h when immersed water:fresh in seawater the in mixtures ratios of of filtrate 1:0, of 1:1, pond and 1:3, became moribund 1.5 respectively. hours after immersion Black in the filtrate. Fish sea lost equilibrium at bream died. eventually and breathing, arrhythmic shallow with immobile became then first, Moribund fish recovered if removed and (Su placed into strongly aerated seawater when exposed to cell-free filtrates of filtrates cell-free to exposed when of filtrate cell-free a and cells A

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 is [as was Boops cells/ 3 Mactra may be ), Pacific 10 × species was (Horiguchi, Venerupis species. Bloom Braarud). There is There Braarud). corsicum nagasakiense , and bogue, . Karlodinium micrum Karlodinium cf. G and surfclams ( . (1995), was associated Gyrodinium corscium

Gyrodinium ), , to neurotoxic fractions of sp. appears to be morpho- et al Gyrodinium cells/ml were lethal to juvenile to lethal were cells/ml 4 Mytilus galloprovincialis 10 × Gymnodinium Solen strictus Gyrodinium (

Heterocapsa circularisquama Heterocapsa Dicentrarchus labrax cells/ml. Economic losses were estimated Cyprinus Cyprinus carpio ., 1999) (Table 2). , which was described from the same 4 264 , after 1 h of exposure and 1.09 10 et al Gymnodinium galatheanum Gymnodinium × , that came in contact with the toxic seawater. toxic the with contact in came that , sp.) significantly reduced the fishes’ heart rates. ., 1995). , it was not definitively identified as such (Arzul et al ), razor clams . (2000) (= (2000) . ., 1995). If this is the case, then ., 1992). recently described by Paulmier

, ), Mediterranean mussels ( ., 1995; Garcés et al et et al ., 1995; Table 2). This et al Gymnodinium nagasakiense Pecten maximus Pecten sp. Gyrodinium corsicum et al Paralichthys olivaceus cf. et al

. (as G in Fukuoka Bay (Yamamoto and Tanaka, 1990). Subsequently, this

cells/ml was observed. Histopathological examination of gills from ) 3 Crassostrea gigas philippinarum , in the coastal lagoon of Diane, Corsica (Table 2). A maximum density of ] 10 ., 1994). Therefore, it is unclear whether this mortality could have been × In September 1993, a bloom reported as reported bloom a 1993, September In pulchellum . another species that produces hemolytic fatty acids (C18:5n3) similar to those known in other microalgae (Table 12). location 1 year later and has also been involved in mortalities of the same fish species (Paulmier e. Heterocapsa circularisquama of bloom harmful a 1988, September In Filtered seawater contained elevated levels of the toxic PUFA octadecapentaenoic PUFA toxic the of levels elevated contained seawater Filtered acid (= C18:5n3). Although reported as the species associated with et this al mortality was attributable to d. Gyrodinium In early August 1992, a Sea bloom South the in finfish of cultured of mortalities mass a for responsible be to novel considered ichthyotoxic of Korea (Kim logically different from other described ichthyotoxic concentrations measured up to 3.3 bastard halibut, ml after 2 h exposure (Kim 1995) was first recorded in Uranouchi Bay, associated with bivalve mortalities (Table 2). Kochi The following year this species was Prefecture, Japan, and was responsible for the mass mortality of Tapes Japanese littleneck clams ( moribund fish showed congestion, hemorrhage, and slight subepithelial edema. moribund fish showed congestion, larvae 9-day-old the of 40% of deaths the caused cells/ml 4 containing Seawater scallops, great of 1.448 of densities Experimental million. $12.5 at G hypertrophy mild very showed lamellae gill secondary the and erratically, swam Fish and edema (Endo c. Gyrodinium corsicum Gyrodinium corsicum with fish and shellfish mortalities near both Corsica and the Mediterranean coast of Spain (Paulmier to similar morphologically is that dinoflagellate naked small, a Daugbjerg in Larsen little information on its toxins, mode of action, or exact effect on aquatic animals. associated with mortalities of sea bass, boops 2.7 oysters ( chinensis LANDSBERGis available. Exposure of common carp, REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., × . No et al et clearance

cells results in circularisquama cell/ml, . 1 H ., 1996, 1997b). Low closed their valves, circularisquama . 10 × H et al circularisquama . . (2001) suggest that the hemolytic H blooms were estimated to amount to et al cells/ml, 50% died within 48 h (Nagai h 48 within died 50% cells/ml, , that were naturally exposed to low circularisquama . H 265 ., 1997b). When Mediterranean mussels were mussels Mediterranean When 1997b). ., at densities exceeding 5 et al et cells/ml, most oysters died within a few days. Dead circularisquama 4 fucata . cells/ml. circularisquama circularisquama (whose cells are comparable in size and shape) at a 3 . H ., 1996). No marked change in behavior was observed 10 ., 1997a). When juvenile oysters were exposed to H × 10 × . et al et al triquetra circularisquama . . cultured H 4 H cell/ml) (Matsuyama (Matsuyama cell/ml) 10 3 × 10 × ., 1996; Matsuyama, 1999). Similar harmful effects were observed on Pacific ., 1996; Matsuyama, 1999). Similar ., 1996), which is comparable to the length of time oysters took to die in the in die to took oysters time of length the to comparable is which 1996), ., Laboratory exposure studies have demonstrated that in addition to bivalves, Caged pearl oysters, ., 1996, 1997b). Purification and characterization of toxic fractions have not been not have fractions toxic of characterization and Purification 1997b). 1996, ., cell/ml. Juvenile mussels showed inhibition of byssus production and sus- 4 circularisquama . rates in the mussels were noticeably edges, showed intermittent shrinkage reduced. of the exhalent siphon, and Mussels were unable retracted the mantle to completely close the valves during the percent mortality (N = 20-min 20) was noted after experiment. 5 days of One exposure to 9.8 to hundred 12.3 10 (Matsuyama experiment the of start the after h 17 to 5 closure valve tained organisms such as gastropods, solitary ascidians, jellyfish, and protists dinoflagellates) (including (Tables 10 and 11) are also affected by 5 to 10 to 5 concentration of 8 species has also been implicated in other mass mortalities of bivalves (Table 2). Economic losses to the shellfish aquaculture industry from the marketable products direct by killing of of decrease mantle, the of shrinkage marked a by characterized were individuals the glycogen lobe attached to the mantle, and et al gut discoloration Hiroshima in (Matsuyama occurred that tide red the during mussels Mediterranean and oysters Bay (Matsuyama (3.7 exposed to 1998b). By comparison, there was no effect on clearance when mussels were exposed to demonstrable effect on mice, finfish, crabs, lobsters, shrimp, starfish, copepods, or diatoms could be found (Matsuyama, 1999). Although current information suggests that there is a direct cytotoxic effect, the Toxicity appears to toxin be closely has related to the still outer components of not the cells, been and a characterized. toxic, proteinaceous substance is probably present on the cell surface (Matsuyama al et 1999). (Matsuyama, conditions neutral under lability high their of because successful Recent studies demonstrate that bioactivity of toxin may be one H of the causes of mollusc mortalities associated with Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS (Matsuyama, decade last the in million) $1 approx. (= yen billion 10 approximately or crustaceans or fish of populations cultured or wild to effects Detrimental 1999). to public health have not been observed (Matsuyama al et field (Matsuyama when oysters and mussels were exposed to filtrate of oxygen concentrations were observed the in during occurred the event mortality main bottom the but Bay, water Ago in layer collapsed had bloom only after the middle period of the bloom when dissolved oxygen levels were still near satura- tion. (50 to 200 densities cell/ml) cell When concentrations mantle. the of contracted and valves, the “snapped” periodically increased to 5 to 10 hemolysis of mammalian erythrocytes. Oda

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . ., et al et al cells/ 4 . (2000). exhibited 10 × Gyrodinium et al Macrocystis pyrifera Macrocystis (Japanese strain) within ., 1996; Mackenzie Porphyra tenera Porphyra Braarud (Kite and Dodge) kill a range of marine fauna et al Chang), was associated with the mikimotoi ) (European strain) within 20 min Larsen in Daugbjerg . K Hansen and Moestrup in Daugbjerg brevisulcata . 266 K mikimotoi was toxic to brine shrimp, abalone larvae, cultures, the soft, leafy part of the seaweed brevisulcatum

has been associated with a series of fish kills in . cf. species (cell concentrations up to 3.33 G species. This species has been classified as classified been has species This species. ., 1997; Dickman and Tang, 1999; Yang and Hodgkiss, Gymnodinium galatheanum (as digitata Karlodinium micrum sp. producing gymnodimine (see section on gymnodimine) on section (see gymnodimine producing sp. brevisulcata et al . . K K is considered to be ichthyotoxic (Steeman Nielsen, 1953, cited 1953, Nielsen, (Steeman ichthyotoxic be to considered is Karenia brevisulcata Gymnodinium Karenia brevisulcat Gymnodinium ., 2000). Karenia mikimotoi are more nonpolar than those of gymnodimine and, unlike cultures demonstrated lysis of Gymnodinium . (Braarud) or K et al (up to 75% of the seaweed cover) during the bloom in the harbor. A dominant Toxins (as yet uncharacterized) of pyrifera brevisulcata brevisulcata . . . Karlodinium micrum Karlodinium but is now known as galatheanum aquaculture facilities in western Japan, southern killed China, and and Hong farms Kong fish (Table coastal 2). 26 Kong’s Hong of 22 affected bloom the 1998, March In 90% of the cultured fish at an estimated loss of $32 million. The fish skin and fish gills of range wide the to addition In patches. reddened with damaged severely were cultured that noted also was it affected, species h. Karlodinium micrum (= Gyrodinium galatheanum) Mass fish mortalities in the region of Walvis Bay, South Africa, in the a 1940s to were attributed (2000) (reported as observed health effects (Chang, 1999a, 1999b). This species is apparently different the of that from found in the same area of New Zealand (Haywood (Chang, 1999b). was introduced into g. Karenia digitata Since the mid-1990s, irregular growth (Baba 1999; Yang 2 h and of tintinnids, amphipods, barnacle nauplii, polychaetes, seaweed brown common the When turbot. lobster and III), and I stage larvae (phyllosoma dead of observations earlier the confirmed This h. 48 to 24 in apart fell and collapsed M ml), now named as as well as other K phytoplankton and macroalgae. Preliminary tests using LANDSBERGf. Karenia brevisculcata In New Zealand, a sporadic massive mortality of fish and other marine fauna was Major 1998. summer to 1997 mid-January from coast east central the along reported kills of eels and flounders were people first 200 noticed than in More Wellington 2). (Table Harbor; invertebrates the marine mortalities and fish pelagic to spread then suffered from respiratory distress; swimmers, surfers, and beach-goers complained of a dry cough, sore throat, running nose, and 1999a, 1999b). eye and skin irritiations (Chang, REVIEWS IN FISHERIES SCIENCE gymnodimine, were extractable with ether (Chang, 1999a, 1999b). 1996a; Chang, 1999a, 1999b). Shellfish bloom collected tested during the negative Wellington for Harbor brevetoxins, presence of but a “quick action” toxin mouse that is typical of bioassays gymnodimine. Mice died demonstrated very the rapidly, within 4 to 6 min. Preliminary tests showed that the toxins extracted from K

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , . has , the et al . The Pecten (30 min); cells/ml. 8 brittlestars

Hemimysis , the crabs Amphioxus micrum 10 . (18 min); the × K , were exposed , the ) within 30 min. 30 within ) Lasaea rubra Calliactis Pleuronectes platessa ) with natural mortalities and ), great scallops ( Eusepia officinalis Trachinus vipera ) Larsen in Daugbjerg , the mysid shrimp mysid the , Gadus morhua has been demonstrated experi- has been demonstrated

Asterias rubens , the bivalve Palaemon serratus Gadus pollachius micrum Rana temporaria Rana . K (15 to 20 min). All fish showed similar showed fish All min). 20 to (15 , and the cephalochordate cells/ml lesser weaver, (5 to 15 min); plaice,

m. Anemonia sulcata Anemonia ), and the 267 , the starfish , the shrimp Tigriopus fulvus (3 h); pollack, micrum galloprovincialis . . and Gymnodinium veneficum K 8 M virescens . (< 45 min); (= K. veneficu 10 G and Gymnodinium veneficum × (= was affected similarly. Other organisms that were slowly and Ctenolabrus rupestris Ctenolabrus Ophiocomina nigra Aplysia punctata Cancer pagarus and and Scyllium canicula Macromysis flexuosus , the fish stayed close to the surface, often with their heads out of the (Ballantine, 1956; Abbott and Ballantine, 1957). Mice that were injected Mytilus edulis and Gobius niger ), seahares ( aurata Blennius gattorugine micrum A concentration of 1.2 A concentration of . K the mussels blenny the gobies jellyfish anemones cnidarian the included affected in South Africa ( and Van der Post, 1967; Pitcher, 1998), and Pitcher, 1998), Van der Post, 1967; Africa (Pieterse and in South been implicated recently in a series of fish kills in aquaculture ponds in the (D. USA Terlizzi, personal communication; Landsberg and Steidinger, unpubl. data). Dead fish had distended gills, and their osmolality blood mouths mean higher were significantly a had wide fish dead open. recently or Blood lethargic from samples had and lethargic were that Fish mOsm). (336 did fish control the than mOsm) (444 been exposed for less than 2 days showed extensive separation of the respiratory gill Increased 1993). (Nielsen, gills the of cells pillar underlying the from epithelium osmoregu- impaired and edema, osmolality, plasma increased in results permeability lation. (2000) was described from Devonport, England (Ballantine, 1956), and when fish were experimentally immersed in cell cultures of the species, it was found lethal to to be dogfish, wrasse the and min); (30 maximus copepods i. Karlodinium veneficum Karlodinium veneficum responses when either immersed in seawater cultures, or exposed swimming violent gobies, supernatant In to away. swim fluid to resuspended attempt violent from a cells was reaction cultures, immediate in or exposed to toxic extracts. either forward or backward continued for The about 2 min and then subsided. Intense vasodilation and expansion of the skin chromatophores down. produced upside a or side marked their color on floated fish the and upset was control Balance pattern. violent spasmodic, a had fish the immersion, after Soon down. slowed rate Breathing extended, gills the with paralyzed, are gobies the while occurs Death reaction. cough and is apparently due to respiratory failure. Rapid death following was also molluscs: noted waved for the ( in Nielsen, 1993). A few reports have associated lamornae Ophiothrix fragilis lanceolatus ( frogs and min 4 to 2 within dead were toxin with mortality documented no been have there toxicity, experimental proven the Despite events in the wild involving Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS to Carcinus maenus mentally to be toxic to and to result in a decrease in the (Nielsen and Strømgren, growth 1991). When rate juvenile cod, of mussels water. Fish died within 2 days of exposure to cell densities of 1.15

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 was ., 2000; ., ., 1992; Pfiesteria Pfiesteria ., 1989b). Callinectes et al et et al ) had been ) (Burkholder ) ., 1992, 1995a, 1992, ., et al , has also been also has , ., 1999, 2000). sp. killing tilapia ., 1988). Although et al et et al ., 1998). Aspects of et al ., 2001b; Morris 2001; et al ), pediveliger bay scallops bay pediveliger ), shumwayae ., 1992, 1995, 1998, 1999, ., 2000; Oldach 2000; ., et al . ., 1997, 1999, 2000; Rublee P , et al et et al ., 1968; Oshima et al Brevoortia tyrannus Gymnodinium ., 1995; Rublee ., 2001a). Unlike more typical HABs, typical more Unlike 2001a). ., ., in press). et al Mercenaria mercenaria Mercenaria Pfiesteria et al et et al et al 268 ., 2001; Glasgow Crassostrea virginica Crassostrea ., 1995; Levin ., 1989b). ., 2000; Marshall 2000; ., ., 1996a) and has been documented to occur along occur to documented been has and 1996a) ., et al et al et al et et al et al et are generalist predatory dinoflagellates that target fish ., 2001; Glasgow 2001; ., ., 2000; Springer ., 1993, 1996; Burkholder and Glasgow, 1995, 1997; Glasgow 1997; 1995, Glasgow, and Burkholder 1996; 1993, ., was first reported as a was implicated in a large freshwater fish kill in Japan et al et was implicated in these mortalities (Burkholder et al ), and northern quahogs ( quahogs northern and ), et al et has also been reported to kill shellfish, blue crabs ( ., 1968) (Table 2), and glenodinine and polonicumtoxins were shumwayae g/ml (Oshima ., 2000; Backer . (1968) suggested that under alkaline conditions glenodinine activity glenodinine conditions alkaline under that suggested (1968) . . µ ., 2001); only a brief summary is provided here. P spp. et al sp. in a North Carolina, USA, aquarium (Smith piscicida . et al et et al et al piscicida ., 1993, 1996) (Table 2). As well as being responsible for mass mortalities P and . spp. may represent a minor component of the plankton community during community plankton the of component minor a represent may spp. P ), pediveliger eastern oysters ( oysters eastern pediveliger ), et al ., 1995; Springer Pfiesteria piscicida ., 1999, 2000, 2001; Bowers 2001; 2000, 1999, ., ., 1998; Cancellieri 1998; ., Argopecten irradians Argopecten Hashimoto the life cycle, detection, distribution, ecology, taxonomy, and toxicity of spp. and their effects on aquatic organisms and threats to public health have been well reviewed in recent publications (Burkholder Noga 2001b; 2001a, and Burkholder, 1995; Lewitus was more lethal than under neutral conditions. During a heavy bloom, pH levels were observed to increase to 8.7 to 9.2 and were accompanied by fish mortalities. When killifish were exposed fins, pectoral to the of stiffness by i.p. followed excitement slight injections were observed symptoms of polunicumtoxins A, B, and C, decrease of response to stimuli, loss of balance, before and death. Fish occasional died 30 to abrupt 40 min jumping after injection with a minimal lethal concen- tration of 0.13 presence the highlighted have threats health public and kills fish of series a Recently, from different cycles life with dinoflagellates armored lightly heterotrophic, small, of those of the more typical planktonic photosynthetic blooming forms. al et isolated from this dinoflagellate (Hashimoto k. Pfiesteria (Hashimoto piscicida Silbergeld Steidinger particularly prevalent in North Carolina’s estuaries for many years, it was not until 1991 that Noga of fish, sapidus ( (Steidinger 1996 in described LANDSBERGj. Peridinium polonicum toxins. produce to documented been have dinoflagellates freshwater of species Few Peridinium polonicum REVIEWS IN FISHERIES SCIENCE and shellfish and Burkholder 1996; are 1995, Glasgow, capable and (Burkholder of microfauna and microalgae, consuming bacteria, a diverse range of prey, including al et Pfiesteria fish kill events and are not typically associated with the production of discolored water so common to other pigmented species (Burkholder (Burkholder Mexico of Gulf and USA the of seaboard eastern the Oreochromis massive fish kills (mostly of Atlantic menhaden, et al 1995b, 1998, 1999, 2001a, 2001b; Lewitus toxic described recently of species second A

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., and et al et ., 2001; ., ., 2000; ., 2000; Pfiesteria ., 2001b). ) in March et al et Landsberg et al et al piscicida ., 1992, 1995; Prorocentrum et al . P ). Further fatalities Further ). et al ., 2001; Moe Moe 2001; ., ., 1999, 2001a, 2001b). 2001a, 1999, ., ., 1999). Although the et al et ., 2001a; Kimm-Brinson 2001a; ., et al et et al Crassostrea gigas et al et of active toxicity where repeated where toxicity active of

bioactive compound mimics the ., 2001a). in fish disease events is still being Amyloodinium ocellatum et al Hudnell 1998; ., Venerupis semidecussata Venerupis Pfiesteria et al et 269 species has not yet been found in the wild receptors on immune cells such as activated ., 1995, 1997, 2001b; Glasgow 7 Pfiesteria ., 2001), areas 2001), ., ., 2000; Burkholder 2000; ., spp. along the eastern seaboard of the USA and USA the of seaboard eastern the along spp. et al et al et et al et or cell-free filtrate taken from aquarium-cultured are still being characterized (Ramsdell Pfiesteria Pfiesteria has a widespread distribution, and most strains are consid- are strains most and distribution, widespread a has ., 2001). Pfiesteria Pfiesteria -like species have been reported from the eastern USA and other and USA eastern the from reported been have species -like et al ., 1995), with significant losses to that state’s economy. Outbreaks ., 1995) or was it proven to be ichthyotoxic. ., 1995). This ., 2001a; Rublee 2001a; ., ., 2001). The role of ., 1995; Ramsdell 1995; ., spp. continue to threaten natural resources, have been implicated in et al and fish kills in Chesapeake Bay in 1997 brought national and ., 2001), it is known that there are different strains of et al et al et et al et al Pfiesteria et al et et al ., 2001; Melo ., 1994) (see section on parasites), was characterized from a Florida aquarium Pfiesteria Mice exposed to Pfiesteria et al macrophages and microglial cells in the brain. Such a mode of action may lead to the neurological effects and inflammatory responses observed in fish and humans (Glasgow l. Prorocentrum minimum minimum Prorocentrum ered to be nontoxic, at 114 least which to in occurred humans. [VSP]) In Poisoning Shellfish Japan Venerupin in (termed event March poisoning 1942, a rare shellfish ( asari toxic consuming after died people resulted after the consumption of toxic Pacific oysters ( 1943 and asari in March 1949. This event was poisoning distinguished events from by other shellfish the symptomatology of the poisoning Symptoms outbreak included hemorrhagic diathesis, centrilobular in necrosis, and fatty humans. degen- eration of the liver, frenzy, unconsciousness, and coma (Akiba and Hattori, 1949). Although toxic shellfish from the same region were associated with implicated implicated in fish kills along the mid-Atlantic coast and has been demonstrated to be ichthyotoxic in fish bioassays (Glasgow several public health incidents in the eastern USA (Burkholder Grattan 1997; Glasgow, and Burkholder (Burkholder toxic highly to nontoxic from range they that Recent findings suggest that a putative neurotransmitter ATP and targets P2X Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 2001; Morris, 2001; Schmechel and Koltai, 2001; Shoemaker and Hudnell, 2001), and 2001), Hudnell, and Shoemaker 2001; Koltai, and Schmechel 2001; Morris, 2001; have been responsible for the mortality of millions of fish in North Carolina alone (Burkholder of international recognition to this genus of distribution (Burkholder, widespread 1998). Although there is a (thus far) to a lesser extent in the northern Gulf of Burkholder Mexico (Bowers et al (Landsberg toxins produced by Moeller (Landsberg water showed signs of decreased learning ability (Levin fish kill events occur are in North Carolina and to a lesser extent the eastern shore of Chesapeake Bay (Burkholder species, suspected of killing fish in pathogenic tropical parasitic fish tanks dinoflagellate but of co-occurring fish with a ( investigated (see section on HABs as stressors in disease). An undescribed An disease). in stressors as HABs on section (see investigated Additional Burkholder 1997; Glasgow, and (Burkholder Mexico of Gulf the in areas Steidinger

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 to 2.5 sp. at 3 appear ., 1999). Exuviella or lysate minimum 10 . × [= et al P has not been ; early larval Mytilus edulis was involved at 33% of bloom of 33% at minimum . minimum strains to mice was Prorocentrum P . Venerupis decussatus P toxins. These results minimum minimum sp. has been implicated . . P minimum P and . Prorocentrum cultures were toxic to mice P mariae-lebouriae minimum ) in South China. Twenty-five ., 1993; Wikfors and Smolowitz, and Wikfors 1993; ., . minimum P . var. P et al et Prorocentrum minimum . P Cardium edule 270 either at a bloom density of 8.9 ., 1997; Denardou-Queneherve minimum . to human health has yet to be confirmed, there confirmed, be to yet has health human to Ostrea rivularis cells/ml. Excess mucus was noted in the gills, P 2 et al 10 ., 1995). Tank-conditioned adult oysters frequently ., 1997). Toxic components of toxic extracts, symptoms in mice (convulsions and × ., 1999) or that this species was not responsible for the for responsible not was species this that or 1999) ., sp. or minimum et al . ., 1993; Luckenbach Luckenbach 1993; ., et al minimum P could be responsible for shellfish toxicity in the natural . ., 1993). Recently, a et al et P et al et fed

minimum , et al (Nakazima, (Nakazima, 1965a, 1965b, 1965c, 1968; Okaichi and Imatomi, .

P ]) ., 1995; Grzebyk minimum Prorocentrum . (Bardouil P were documented in the Netherlands in blue mussels, showed no cytotoxicity on hepatocytes in culture, suggesting that this showed no cytotoxicity on hepatocytes et al (as cell/ml died within 14 days, and 43% of those fed those of 43% and days, 14 within died cell/ml 5 More recently, the potential toxicity of some Although the risk of risk the Although minimum minimum minimum . . . 10 and histopathology revealed intense hemocytosis, tissue edema, and atrophy of the and histopathology revealed intense hemocytosis, tissue edema, and atrophy digestive tubules (Yomgjia × do not in the presence of bloom densities of development is impaired and oysters die at high rates when from ruptured cells is present in culture tanks. All of the juvenile eastern oysters, Crassostrea virginica LANDSBERGminimum REVIEWS IN FISHERIES SCIENCE mariae-lebouriae (Kat, 1979); in Portugal in the cockles (Silva, 1985); in Norway (Tangen, 1980); and in China (Chen and Gu, 1993). In the Netherlands, the poisoning event was probably DSP (Quilliam and Wright, 1995), but it is unclear what toxins these other events were attributable to. 1979) and partially purified toxins from (Okaichi and Imatomi, 1979), conclusive evidence that with VSP has still not been obtained. Other shellfish toxicity events associated with P confirmed, and natural shellfish toxicity was demonstrated in laboratory exposures (Denardou After injection of to block calcium channels, and in laboratory exposures toxins could accumulate in The mussels. cultured of meat and hepatopancreas the in amounts equivalent nearly a during collected mussels wild of samples in found was toxicity same spasms with pronounced palpitations) were suggestive of neurotoxic activity. When activity. neurotoxic of suggestive were palpitations) pronounced with spasms died mice and rapidly appeared symptoms neurotoxic injected, was dose sufficient a within minutes (Grezebyk hepatotoxic effects noted at that time. The exact role of bloom. In 1993, consumption of mussels from the Salses-Leucate Lagoon French on Mediterranean the coast produced neurological symptoms in mice those similar observed to when mice were exposed to suggest that environment and thus present a risk for human P health. Mediterranean strains of 1942 in event poisoning Japanese the in involved strain the from different was strain (Denardou-Queneherve confirmed since the earlier poisoning reports from Japan, and venerupin toxins have toxins venerupin and Japan, from reports poisoning earlier the since confirmed not yet been characterized (see section on microcystins and hepatotoxic shellfish poisoning). is increasing evidence that this effects, species inhibition affects of aquatic feeding, organisms. P Pathological and mortality occurred in shellfish exposed to concentrations of 2.01 to 6.77 1993, 1995; Wikfors in mass mortalities of flat oysters ( hectares of oyster fields worth more than two million yen were lost between late April and late May 1994. Water samples were dominated by

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., and , but et al 2 mixed Mercenaria ., 1995; Shaw Prorocentrum , were fed the minimum . may produce an et al ., 1999), the likely the 1999), ., minimum P . had an abnormal ., 1997) (Table 10), P et al et et al , (4) was reported in Gwadar Bay, supported 50% survival, and minimum proved to be an unsatisfactory an be to proved . P does pose a threat to both public both to threat a pose does Isochrysis Argopecten irradians ., 1993). micans . caused 100% mortality in one week in , and (2) and (3) two concentrations of concentrations two (3) and (2) and , ., 1995, 1996; Miralto P minimum minimum et al

. Prorocentrum minimum P minimum et al 271 . and P ., 1996; Uye, 1996; Ban minimum alone. Another group was not fed. Clams survived Clams fed. not was group Another alone. . et al sp. (60%). The mortality event did not appear to be P Isochrysis Prorocentrum and was considered to be sufficiently toxic to affect aquatic ., 1994; Ianora Isochrysis Prorocentrum micans Prorocentrum ., 1990). ., 1998) or due to diatom toxicity (Ianora toxicity diatom to due or 1998) ., et al Congresox et al et al et Isochrysis minimum , and (5) and ,

cells/ml coincided with mortalities of nine species of fish (Table 2), , held in grew poorly during a bloom of of bloom a during poorly grew Sound Island Long in held , 4 mixed with the standard bivalve food, 10 × Isochrysis ., 1995a, 1995b; Chaudron These few examples indicate that indicate examples few These Following observations that experimentally caged northern quahogs, northern caged experimentally that observations Following An unusual fish kill associated with micans . all of which could have harmful consequences for marine food webs (Poulet 1994). Although researchers have debated whether the observed effects on feeding are due to a nutritional insufficiency in the copepod diet (Jónasdóttir and Kiorboe, Jónasdóttir 1996; species (Rabbani health and to natural resources, yet the exact mechanism toxicity and is not possible yet clear. mode of et al apo- involvement of bioactive compounds has been demonstrated recently. Several related to low dissolved Prorocentrum oxygen. When it appeared at high concentrations, Several genera of diatoms have been shown feeding recently deterrents, to or cause mortality, affect act the as reproductive Poulet, 1993; Poulet success of copepods (Ianora and produced discolored brown water for about 3 days. Maximum cell concentrations of 4.5 mostly pike conger 2. Diatoms density died within 22 days. 22 within died density food when present at high concentrations, reducing filtration rates and mortality in elevating juvenile oysters (Luckenbach mercenaria spp., controlled laboratory feeding Hard clams and bay scallops, Smolowitz, 1993). exposures were undertaken (Wikfors unialgal (1) following: and P Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS with well in all experiments, but in none of the diets supported adequate growth in bay scallops. Unfed bay scallops showed incremental mortality over a 5-week period, mixed or individual diets of a mixed diet of one trial and in 4 weeks in the second. Bay histopathological scallops observations showed ingested poorly developed digestive diverticula, at- thrombi Large necrosis. and vacuolation, abnormal with epithelium the of tenuation diverticula, digestive the mantle, the of system vascular open the in noted also were and tissues of the gill, heart, and kidney. thrombi persistent produces and cells absorptive affects systemically that enterotoxin throughout the vascular system (Wikfors and Smolowitz, 1993). Additional studies showed that spat of eastern oysters exposed accumulation to of lipid autolysosomal in dinoflagellate contained diverticulum digestive the of cells the absorptive stomach epithelium. Accumulation bodies bodies, indicating nutritional within interference (Wikfors and Smolowitz, 1995). southwestern Pakistan, in November 1987. The bloom extended over 7 km

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 by , that . The (Parry ., 1995a, Thalassiosira et al Mytilus edulis ) concurrently . Additionally, was fed a diet californicus . and T cells/ml was totally 7 demonstrated a dose- a demonstrated 10 Rhizosolenia chunii Rhizosolenia × Paracentrotus lividus tricornutum Tigriopus Tigriopus californicus , total egg production and . P ., 1995). It was suggested that suggested was It 1995). ., ., 1995a). When adult female ., 1995). Whether these effects these Whether 1995). ., . (1989) discussed the possibility the discussed (1989) . to 5 6 et al et et al et et al produced eggs for only 3 to 4 days, 4 to 3 only for eggs produced et al et , the egg quality was poor and the . Researchers considered that these 10 × anomalies of the eggs included a dark a included eggs the of anomalies

., 1999). In the presence of the diatom (Shaw Thalassiosira rotula Thalassiosira 272 et al Phaeodactylum tricornutum tricornutum Phaeodactylum Thalassiosira rotula ), and flat oysters ( californicus tricornutum were more susceptible to by the protist the by parasitism to susceptible more were . . P T Temora stylifera Temora . Embryonic development was arrested when eggs or tricornutum chunii . . P R were fed Pecten alba , female , are feeding deterrents to the copepod ), scallops ( Prorocentrum minimum ., 1989). A bloom of this species occurred in Port Phillip Bay, southeastern ., 1994). Bioassays of embryos of the , Chemically mediated inhibitory effects of diatoms on egg viability have also Chaetoceros curvisetum blocked at the one-cell stage (Buttino a. Rhizosolenia One of the few documented cases demonstrating a chronic effect to and subsequent exposure with associated that was shellfish of mortality bitter The bloom. the during reported mortality no was there taste, bitter a developed taste was concentrated in the digestive gland, which also showed extensive inflam- mation and degeneration. The bitter taste retained in the mussels rendered unmarketable them for 7 months. Digestive gland lesions September were 1987; these lesions evident progressively became more severe, in and after 3 mussels to 8 in months, mortalities began. No other pathogens or pollutants were demonstrated to Parry mortalities. chronic the for responsible be to exposed scallops that amounts of apo-fucoxanthinoids necessary to reduce feeding in costatum Skeletonema were due to bioactive compounds or some known. other mechanism is currently un- et al Australia, from late August to mid October 1987. Although mussels ( planulatus LANDSBERGby produced pigments fucoxanthoid REVIEWS IN FISHERIES SCIENCE after which they became sterile or died (Ianora died or sterile became they which after pseudonana pseudonana lower times 1000 approximately was range This ppm. 20.2 to 2.22 from ranged 50% than the total apo-fucoxanthinoid concentration in concentrations of apo-fucoxanthinoids ranging from 36.8 to 76.7 ppm caused 50% mortality in a population of compounds may be ecologically significant in the control of bloom formation and in reducing the amount of copepod grazing on the diatoms (Shaw (Ianora 20% as low as was rate success hatching 1995b). been considered (Uye, 1996). When the copepod of the reduced egg production and pigments viability produced by were due to the apo-fucoxanthoid hatching success were considerably lower than when they were fed the dinoflagel- the fed were they when than lower considerably were success hatching late were exposed to diatom extracts. Structural extracts. diatom to exposed were brown, and opaque outer membrane, globular cytoplasm, blockage of pronuclei, and dispersed chromatin scattered in the egg matrix of unhatched eggs (Poulet et al diatom the of extracts to exposed were dependent effect on cell division. The assembly of tubulin necessary for microtu- incubated embryos and eggs of development The inhibited. was development bule in a water-soluble extract equivalent to 5

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 blooms are blooms . Fibrocapsin was Fibrocapsin . Phaeocystis japonica . F ., in ICES, 1998). ., 1998) and, in general, has not been et al et al was present along the coast of Fujian and Fujian of coast the along present was ., 1987). When colonies break up and the 273 was observed in the , two adult were reported in the German Wadden Sea in ., 1987). High concentrations of mucilaginous et al ., in ICES 1998). et al et al japonica is not usually associated with aquatic mortalities, a fish a mortalities, aquatic with associated usually not is . F reduce seawater quality (Rogers and Lockwood, 1990) and Phaeocystis globosa Phaeocystis is a regular component of phytoplankton communities in Fibrocapsa japonica pouchetii . P Phaeocystis globosa Phaeocystis sp., which although known to be pathogenic to shellfish (, 1993), At the same time that Huang, 1999). kill involving it was reported in southeastern of bloom a months, 2 China in 1997. For approximately detected in both the water samples and from blood and organ samples from seals. Whether fibrocapsin played a role in the seal the mortality is still unknown, or is it clear whether the animals might have been exposed through through direct the diet contact (Rademaker or a. Phaeocystis globosa Although Guangdong provinces. The discoloration caused by the bloom was intense enough to be detected by SeaWIFs imagery. The bloom caused a severe mortality of caged fish — an estimated 60,000 tons of fish, a value of $8 million, were killed (Lu and b. Phaeocystis pouchetii Phaeocystis pouchetii boreal and arctic environments (Lancelot reports early Nevertheless, 1989). Moestrup, and (Larsen species toxic a as regarded food pelagic the affected negatively have may species this of blooms that suggested 1930). (Savage, herring of migration the influenced and web 4. Prymnesiophytes Perkinsus was not implicated in these mortalities. 3. Raphidophytes a. Fibrocapsa japonica Low densities of and three newborn seals kept in a rehabilitation for positive center were center adjacent the in collected to samples Water the sea died. now considered to be nuisance blooms that have increased in some areas because of coastal eutrophication (Lancelot Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS and fish killed species this by produced toxin studies, experimental In mid-1990s. the was determined by mass spectrometry to be of a slightly different structure to that experiments Preliminary fibrocapsin. named tentatively was toxin The brevetoxin. of concentra- at conductivity neural blocks fibrocapsin that showed nerves isolated on tions as low as 0.01 ng/ml. After mice developed were given tonic-clonic subcutaneous seisures, injections they indicating that the through the blood-brain toxin barrier (Nannen is capable of passing bloom collapses, characteristic layers of foam may form long bands along the open bloom collapses, characteristic layers of foam may form long bands along coast (Rogers and Lockwood, 1990) unsightly or foam banks (Lancelot beach and ultimately produce massive colonies of can lead to reduced bivalve spawning success and poor viability of larvae (Walne,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., had et al Gadus spp. that pouchetii pouchetii . . Phaeocystis but actively P P pouchetii . in north Wales during feeding P Acartia

is an inadequate , m diameter) or solitary Phaeocystis µ pouchetii pouchetii . . P P colonies adhered to the gills. The gills. the to adhered colonies ., 1980). Oyster larvae affected by affected larvae Oyster 1980). ., was extracted from cultures by Arenicola marina et al et is toxic, unpalatable, or at least of low of least at or unpalatable, toxic, is resulted in a mass mortality of numerous of mortality mass a in resulted ., 1990). This would seem to indicate that 274 pouchetii . Phaeocystis P et al sp. toxins. The presence of active material was material active of presence The toxins. sp. pouchetii . P pouchetii . ., 1990). Egg production of female P produces chemical compounds that are lethal to cod to lethal are that compounds chemical produces et al Prymnesium blooms. A light-induced toxic effect on cod larvae was found , either as gelatinous colonies (>200 and to collect food, because food, collect to m), was not significantly different from that of starved females (Verity different from that of starved m), was not significantly µ blooms grew 26 to 83% slower than oyster larvae in control groups did groups control in larvae oyster than slower 83% to 26 grew blooms pouchetii pouchetii . . ., 1990; Nielsen P P Phaeocystis pouchetii Phaeocystis , larvae reported in aquaculture facilities in Norway coincided with the end , larvae reported in aquaculture facilities et al ., 1998). The toxic effects of filtered seawater on cod larvae during In 1988, a heavy bloom of bloom heavy a 1988, In There is mixed evidence that evidence mixed is There It was observed that increased mortalities and the poorest growth of cod, cod, of growth poorest the and mortalities increased that observed was It The assumed toxic principle of pouchetii . copepod feeding behavior is not altered by the presence of toxins (Stabell grazed on senescent colonies (Estep 1999). benthic invertebrates, particularly lugworms, cells (3 to 5 and Smayda, 1989). Copepods avoided healthy colonies of (Rogers and Lockwood, 1990). Mortalities were attributed to bloom; the anoxic conditions collapse developed when dying of colonies formed the a mucilaginous could fish mobile whereas suffocated, were fauna Immobile bed. sea the over layer leave the area (Rogers and Lockwood, 1990). An enormous bloom of LANDSBERG1974, cited in Rogers and Lockwood, 1990). compounds such as acrylic Concentrations acid (Sieburth, 1960), methyl (Armstrong of bromide (Saemundsdottir dimethyl potentially as such toxic compounds sulfur volatile and 1998), Matrai, and mortalities animal marine cause to sufficient considered not were 1960) Boalch, and (Rogers and Lockwood, 1990). In New Zealand, blooms of the “Tasman Bay slime” fishing choke and fish destroy to said were and 1860s the as early as reported were nets (Hurley, 1982, cited in Moestrup, 1994). REVIEWS IN FISHERIES SCIENCE larvae, and the production of these compounds appears to be dependent on level the of irradiance. When fish larvae were offered (Walne, 1970, cited in Verity and Smayda, 1989), and recently. nutritional value to copepods (Tande and Båmsted, 1987; Verity and Smayda, 1989; 1987; copepods (Tande and Båmsted, nutritional value to Estep were fed in the western Dutch Wadden Sea in May 1978 harmed the ability of blue mussels, edulis Mytilus reduced food intake caused by clogged gills and feeding inhibition contributed to starvation, to reduction in lipid and protein content, to resorption of ripe gametes, ( failure reproductive to ultimately and P Smayda, and Verity in cited 1971, Walker, and (Gabbott oysters adult for source food 1989). morhua phase of experiments, acute mortalities were more often observed when also found in filtered seawater collected during a bloom, confirming that confirming bloom, a during collected seawater filtered in found also been exposed to high irradiance levels in spring (Eilertsen and Raa, 1995; Aanesen et al blooms indicate that the toxins are released into spring the with water. coincided levels It toxin high was that postulated and water that intake the with entered toxins bloom characteristics (Eilertsen and Raa, 1995). filtering and solid-phase sorbent techniques. The active material from the cultures was found within the chemical fraction previously established for the separation of Chrysochromulina

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . was , and ., 1999). cells and cells thiebautii is common . et al T Trichodesmium Trichodesmium Trichodesmium Phaeocystis Hilsa kanagurta filling 80 to 90% of the of 90% to 80 filling Trichodesmium bloom dominated by dominated bloom did not create a problem for farmed Trichodesmium value ranging from 5- to 85-mg bloom , fed on sardines, 50 275 , with , erythraeum appear to be anesthetizing and may temporarily may and anesthetizing be to appear . thiebautii . T - T , occasionally have been implicated in mortality inci- bloom in Brazil was even held responsible for “Tamandare for responsible held even was Brazil in bloom pouchetii . P thiebautii is different from that described for other prymnesiophytes. cells had been removed would also indicate other mecha- . Larus brunicephalus . (1998) proposed that ingestion of toxic of ingestion that proposed (1998) . aggregations provide an extremely important habitat for a T spp. thiebautii et al et ., 1963). . Rastrelliger kanagurta Rastrelliger T blooms are ubiquitous in tropical, subtropical, and temperate seas et al pouchetii . Phaeocystis P . (1999) suggested that the fact that larvae also died in filtered seawater and Trichodesmium releases the active material into the natural environment. Hemolytic et al Trichodesmium Generally, coastal blooms of 1 h. However, an unexpected response was observed: some of the flies that were presumed dead regained motility within 4 hours, and the proportion of recovered flies was inversely proportional to the dose injected. Regained found when the motility injected material was obtained was from filtered natural seawater. The also by released compounds narcotize fish larvae; in excess doses, they may be toxic (Stabell Although Aanesen Although intestinal absorption was the most probable route of Stabell intoxication of cod larvae, from which nisms: (1) cod larvae may ingest particles that are coated by partly precipitated hydrophobic, material that contains “toxic” compounds, or (2) (during a ionic toxin absorption regulation) resulting from harmful the material. swallowing of seawater containing began to decay, and oysters subsequently died (Chellam and Alagarswami, 1981). Recently, a highly potent neurotoxic mixed a compound from isolated that resembles anatoxin-a was were Caribbean the in Islands, Virgin US Thomas, St. from collected samples Bloom highly neurotoxic to mice with an i.p. LD pearl oysters and there were no reported mortalities. However, when bloom water was used for experiments on oysters isolated in the laboratory, the Indian mackerel, Indian 5. Cyanobacteria a. Trichodesmium Trichodesmium and form some of the largest phytoplankton aggregations ever observed (Sellner, 1997). dents and negatively affecting marine life (Table whether these are due to poor water 2), quality or to toxicity. A public health incident but it is still unclear a as involving to fever,” but the exact etiology of this disease and the role of pouchetii activity was almost absent in principle in the material tested, demonstrating that By the the fly-bioassay toxic method, a rapid response to injected resulting material in was a obtained, high proportion of apparently “dead” flies being registered within diverse microheterotrophic community, including cyanobacteria, bacteria, dinoflagellates, diatoms, copepods, and protists, hydroids. Metazoan herbivory by fungi, and fish on harpacticoid copepods, salps, zooplankton, adverse apparent no were there and 1997), Sellner, 1994; 1992, Roman, and (O’Neil effects when seagulls, species, marine dominant Two 1970). (Ramamurthy, volume gut fishes’ erythraeum unclear (Satô Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 extracts Oscillatoria ., 1981), and Scomberomorus et al from the Caribbean the from erythraeum . T was nontoxic (Hawser or erythraeum blooms can apparently feed on . T erythraeum . thiebautii . T T (Suvapepun 1989). spp. (Table 10). However, harpacticoid 276 . Trichodesmium ., 1992; O’Neil and Roman, 1994). By producing ., 1992). These authors further recommended that Daphnia is toxic to some cyclopoid and calanoid copepods, are not usually considered toxic (Hawser and Codd, Trichodesmium is toxic, whereas et al et al thiebautii . may avoid predation by dominant copepod groups (Haw- may avoid predation ), and T with no adverse effects and appear to have some undefined erythraeum thiebautii . . Artemia accumulated into reddish-brown foam patches in the surf. Decom- T T bloom and associated poor water quality (Chacko, 1942; Table 7). In 7). Table 1942; (Chacko, quality water poor associated and bloom ., 1991; Hawser and Codd, 1992). No significant inhibition of cholinest- of inhibition significant No 1992). Codd, and Hawser 1991; ., , which was implicated in ciguatera poisoning. Extracts of the water- et al et Trichodesmium thiebautii were used as a positive control. ., 1992). Based on the various assays and toxicity tests evaluated, it was . T Although there was no evidence for toxicity in toxicity for evidence no was there Although Blooms of Trichodesmium thiebautii et al erythraeum . possibly having a role in coral mortality event bleaching involving numerous species (Coles, of animals was 1994). caused by In a decaying India, T a wide-scale June 1983, when a bloom was driven to shore along Trichodesmium the east coast of Thailand, position of the bloom along a 20-km stretch of beach was associated with respiratory with associated was beach of stretch 20-km a along bloom the of position However, noted. were life marine wild to effects harmful no but humans, in irritation an was there and coast, the along farms fish to damage extensive caused bloom the conditions anoxic to attributed was mortality fish The million. $1.16 of loss estimated generated by decomposing and Codd, 1992; Hawser with associated bacteria to due toxicity possible of because tested be cultures axenic natural populations of associated with ciguatera-like outbreaks in 1991; Sellner, 1997), but they have been Australia (Hahn and Capra, 1992), avoidance mortalities behavior by of fish coral (Nagadhushanam 1967; polyps Eleuterius (Endean, 1977), (Hawser and Codd, water-soluble 1992), by produced those a to similar strain mice in from symptoms produced Australia that material contained toxic, water-soluble extracts of the flesh commersoni of the narrow-barred Spanish soluble material mackerel, from the cyanobacterium and the fish were chromatographically indistinguishable. The water-soluble material was lethal to mice within 30 included injected i.p. at a concentration of 0.25 g/kg mouse. Symptoms in the mice min if ataxia, piloerection, quiescence, loss of balance, exophthalmus, labored breathing, and frequently, convulsive spasms. also Necropsy was component lipid-soluble A sinuses. engorged and revealed vessels, blood distended markedly swollen livers, identified that resembled the chromatographic and toxic properties of a scaritoxin- like substance isolated from narrow-barred soluble Spanish component mackerel. was When injected the lipid- into produced by classic ciguatoxin, but mice the ciguatoxin-like material was different from it produced signs classic ciguatoxin. No epibiotic organisms such as dinoflagellates similar were observed to to those LANDSBERGdry weight/kg body weight. Toxicity in mice was characterized by severe convul- sions and then death within 2 to 20 lacrimation, min or from signs respiratory typical failure. of No hepatotoxic microcystin salivation, poisoning were observed (Hawser erase was obtained when high levels of the REVIEWS IN FISHERIES SCIENCE adaptive strategy (Hawser toxin(s), ser suggested that were used, in contrast to the inhibition noted when neurotoxic extracts of extracts neurotoxic when noted inhibition the to contrast in used, were agardhii brine shrimp ( copepods that are associated with toxic

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 4 ; and ; did. (Hahn in situ in in situ blooms ingested diet, and diet, Trichodesmium C, pH 8.3, NH 8.3, pH C, Trichodesmium blooms in the have a possible a have nomades o Acartia

. Coincident with Coincident . Trachinotus blochi Trachinotus Ostrea Trichodesmium Trichodesmium Tetraselmis suecica erythraeum . , and T ., 1984, cited in Suvapepun . Levels of zinc, cadmium, and cadmium, zinc, of Levels . Trichodesmium Trichodesmium et al trichomes/ml at 29 at trichomes/ml 2 , larvae were exposed to water from (Lam and Hai, 1996). Poor survival of were exposed to intact 10 both in natural blooms and laboratory

, × filaments, filaments, but heterotrophic bacteria may sp. developed beyond the first protozoea first the beyond developed sp. 277 Lopha cristagalli , was of no nutritional value to the larvae. The Saccostrea amassa Saccostrea ., 1993); snubnose pompano, snubnose 1993); ., Lates calarifer Acartia tonsa et al et Trichodesmium bloom, 30% were killed within 24 h when the density bloom, 30% were larvae was documented during a bloom of bloom a during documented was larvae Trichodesmium Penaeus monodon Trichodesmium Trichodesmium 1.46 ppm (Suvapepun bloom in pond culture in Vietnam has been associated with 3 Trichodesmium filaments were starving. Swollen mitochondria and vesiculated ., 1998). erythraeum merguiensis Pinctada margaritifera . . P T et al erythraeum . survivors were weak, less responsive to probing, and unable to move. T A When white sea bass, When adult copepods, A recent report of an indirect effect of the presence of presence the of effect indirect an of report recent A healthy cells, egg production could not be supported on a on supported be not could production egg cells, healthy poor assimilation of food was indicated. After Acartia 17 h immersed in aged cultures, Affected copepods also had distended possible toxic effect and (Guo and Tester, everted 1994). intestines, suggestive of a mortality of the shrimp the prawn the sp., there were no apparent ill effects, but a high mortality rate was noted when adult when noted was rate mortality high a but effects, ill apparent no were there sp., copepods were exposed to aged or homogenized cells. This suggests that intracel- lular toxins were released after cell lysis or disruption. Although spp. in Albatross Bay, Gulf of Carpentaria, Australia. Prawn larvae reared These stage. protozoea first the beyond develop to failed bloom the during results were confirmed by the results of laboratory experiments in which none of fed were that larvae the role in ciguatera outbreaks in Australia and can be accumulated in narrow-barred (Endean mackerel Spanish the oysters and Capra, 1992). It is unclear, however, what potential effects these toxins have on may the health of these species. a natural stage, whereas 94% of those fed Although the larvae ingested green have been present. Ciguatoxin-like compounds from compounds Ciguatoxin-like present. been have 6.915 exceeded cyanobacterium the of 1992). Mortalities in this case, however, may have been quality. due to poor water experiments, the be associated with the 1.54 ppm, and NO ultrastructure of larval gut Trichodesmium cells indicated that the endoplasmic larvae reticulum that were had observed in ingested responses the to starvation gut that cells; have been these documented in are (Preston other the crustacean larvae typical Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS may be very significant to molluscan and public health. water Significant quality were changes documented in during intensive central Great Barrier Reef, near Townsville, Australia. Chemical speciation mea- surements have established that of presence cadmium, the during lead, bioavailable more become and copper ions in seawater the increased bioavailability of trace elements during the blooms, significant and sustained increases in total iron, zinc, cadmium, copper, silver, and manganese oyster, lip black the in occurred copper in these oysters exceeded health guidelines by 8000%, 4000%, and 3000%, and 4000%, 8000%, by guidelines health exceeded oysters these in copper respectively (Jones, 1992).

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . , ., were (and , and et al

rotula . T Chaetoceros Salmo salar Salmo wighami

, and . (1989) suggested bloom, pathological ., 1989). et al that had a granulomatous a had that et al

, reared in a seawater net Chaetoceros , ., 1989). Under laboratory spp. damaged the physical et al on sockeye and coho salmon ., 1993; Rensel 1993). Short-term Oncorhynchus tshawytscha Chaetoceros et al Salmo salar Thalassiosira aestivalis caused mortality at concentrations as , Chaetoceros convolutus was implicated in a mortality of red king ., 1995). 278 . C et al [see under silicoflagellates] in one incident) , in Alaska. Although low levels of dissolved ., 1974, in Rensel have been implicated in salmonid mortalities in the in mortalities salmonid in implicated been have Catostomus commersonii Catostomus concavicornis . et al C , to Skeletonema costatum convolutus . C cells/ml seawater (Yang and Albright, 1994). Although several kisutch spp. Chaetoceros convolutus 1 . and O Dictyocha speculum 10 × and Paralithodes camtschatica An unusual case of mechanical damage associated with several diatom species debile . 1. Diatoms A histopathological examination of the gills in moribund fish showed congestion of congestion showed fish moribund in gills the of examination histopathological A the central venous sinus at the base sloughed, of the and primary fragments lamellae diatom and diffuse Mucus-containing edema lamellae. at secondary the of base the necrotic epithelial cells were observed between the lamellae. No other pathogens were observed in the gills, and lesions were most severe when directly associated with accumulations of algae (Kent sucker, white a in reported was that the mortalities resulted from direct clogging and abrasion of the delicate structures by the silicified diatom frustules. Market value of the Atlantic salmon gill lost was approximately $1.6 million (Johnson, 1988; Bruno concavicornis Pacific Northwest, where fish are cultured in net pens (Taylor, 1988; Rensel theories suggested that the mortalities were due to penetration of the fish gills by the diatom spines, which resulted in excess mucus production, hemorrhaging damaged of tissue, and blood hypoxia or subsequent secondary bacterial infections (Bell, 1961), other mechanisms may also have been operating (Yang and Albright, 1992; Rensel, 1993). Barbed spines of pen in British Columbia were associated with a dominated dense mostly by bloom of mixed diatoms enteritis. Wolke and Trainor (1971) suggested that the granulomatous response to the of dioxide silicon the to either response cellular a by generated was diatoms the diatom cell wall or to the mineral exoskeleton (= ). changes in the gills associated with the penetration of diatom spicules were also a 1995). Mahoney, and (Tester mortality the in factor major C smolts in two separate incidents extensive in tissue necrosis and sloughing, with separation Scotland. of the secondary lamellae Histological gill sections to noted were cells Plankton filaments. the of base the at hyperplasia moderate and showed be in close association with the secondary lamellae. Bruno 1989; Yang and Albright, 1992, 1994; Albright laboratory studies on the effects of smolts indicated that exposure to several million cells/l caused over total a mortality (Bell 1- to 7-h period a. Chaetoceros In August 1992, low as 1.5 Mortality and gill lesions in Atlantic salmon, crabs, oxygen were reported at the same time as the salmon, Atlantic 550,000 to up of deaths the for responsible considered conditions, exposure of juvenile chinook salmon, coho salmon, LANDSBERGU. HARMFUL SPECIES THAT CAUSE MECHANICAL DAMAGE REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , which led to cells/ml immedi- , began dying within dying began , ., 1989). mykiss . did not penetrate host et al O ). There was no evidence 2 kisutch , like other diatom species, . O concavicornis . Corethron cells/ml — mortalities can occur 4 C

5 minimus 10 . × L 10 × 279 were detected, and unusual behavior and mortalities increases their susceptibility to bacterial disease (Albright -like diatoms. The setae of minimus . L sp. Chaetoceros experimentally exposed to 4

Corethron , ., 1993) (see section on disease). at higher concentrations (Clement, 1994). c. Leptocylindrus minimus In November 1989, in the center and south of the Chiloe Archipelago, Chile, high concentrations of were observed in trout and salmon (Clement and Lembeye, 1993). A similar event occurred in 1993 in the same area. Although it is not clear what mechanisms involved are in the mortality, it is possible that b. Corethron salmon, coho g 40 60,000 to up 1987, October early In their of corners the to retired salmon bloom, the During damage. mechanical causes gills The water. the of out fins dorsal their with surface the at swam trout and cages, loss a was There secretion. mucus excess from pale slightly were fish the of some of of appetite, especially in trout, but also in coho salmon and Atlantic salmon. These effects were seen at concentrations below 1 integrity of the respiratory epithelium of rainbow trout, increased mucus secretion. Accumulation of mucus on and between the secondary lamellae inhibited the oxygen uptake by concen- the lactate and gill, glucose resulting blood in increased by hypoxic (characterized conditions, metabolism anaerobic trations as well as haematocrit values), leucopenia count), and subsequent mortality (Yang and Albright, 1992, 1994). (depressed Atlantic salmon, white blood cell Salmo salar ately displayed a cough response and began to die within 3 h. Blood from dying fish was severely hypoxic and hypercapnic (elevated CO that the diatom spines had penetrated the gills. The primary mechanism inducing hypoxia and hypercapnia appears to be oxygen minimal mucus in production resulted production resulting mucus Excess from epithelium. diatom gill the of irritation pressure partial blood-oxygen the in change a to led which gills, the across transfer (Rensel, 1993). There is salmonids also to compelling evidence that et sublethal al exposure of Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS one week after their transfer to saltwater net pens located along the coast of British Columbia. Clinically, the fish hung listlessly near the top and sides of Rapid, the labored respiration with constant net opercular flaring was typical. pen. Some fish had petechial hemorrhages midway dying stock remaining the of along 10% approximately with December, early and November the fin rays. Mortality each week. continued Surviving fish had extremely poor growth rates, and through many subsequently and Histological disease). on section (see Disease Kidney Bacterial by affected became ultrastructural examination of sequential samples of dramatic This lamellae. gill gill of fusion tissue extensive by accompanied branchitis revealed suppurative a dramatic host response and subsequent high rate of mortality appeared to be in response to a bloom of epithelial tissues, except in occasional nonrespiratory areas on the arch. Instead, the impacted diatom apparently became enveloped by the process of Extensive lamellar lamellar fusion. fusion reduced the surface area for change, resulting oxygen in reduced and respiratory efficiency electrolyte (Speare ex-

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., and et al densities Aureoumbra ., 1993; Buskey , selective avoid- selective , and ., 1998). et al et al cells/ml were recorded, were cells/ml 2 irritated the gills, which ., 1993). Cell counts of up 10 × Aureococcus Dictyocha speculum Dictyocha Aureococcus et al Aureococcus anophagefferens Dictyocha (K. Tangen, personal communication, in 280 ., 1987; Buskey and Stockwell, 1993) have cells/ml. Gills were clogged with mucus and 3 blooms (Table 6). In April 1987, at a rainbow et al 10 ., 1996; Bricelj and Lonsdale, 1997). Only a brief × bloom, but with the naked phase of the life cycle. speculum ., 1997). . , died, and it was suggested that the high density of et al D et al to 1.3 2 (Cosper 10 speculum . × D Salmo salar Dictyocha speculum cells/ml were reported (Thomsen and Moestrup, 1985). An organism 4 10 × ., 1996, 1997; Street Although some protists are capable of consuming of capable are protists some Although In Denmark in 1983, a wide-scale mortality of cultured rainbow trout was also ance by most protists has been demonstrated both experimentally and in the field, brown to contribute may microzooplankton by grazing of inhibition that suggesting Gowen, 1987; Ø. Moestrup, personal communication, in Erard-Le Denn, 1991). 3. Pelagophytes Extensive blooms of brown tide caused by are considered to be nontoxic, the sheer biomass and persistence of monospecific blooms that can last for several months are sufficient to result in numerous direct been have tides Brown habitats. their and organisms aquatic on effects indirect and responsible for dramatic losses in species abundance and diversity (Cosper et al referred to as “flagellate X” associated with cultured fish mortalities in Ireland and the as same the be to considered was 1991) Denn, Erard-Le 1987; (Gowen, Scotland naked flagellated form of trout farm in Douarnenez, France, an overnight bloom produced turbid water, and asphyxiation. and distress of signs showed fish caged a. Dictyocha speculum mechanical with associated been have Europe in fish cultured of mortalities Several damage due to varied between 1.3 of the lamellae. It revealed edema and hyperplasia many algal cells; histopathology was suspected that the siliceous skeleton This potential. exchange gill of reduced a and mucus more produce to fish the caused effect, coupled with possible low levels of dissolved 1990). Ryckaert, oxygen and overnight Denn due (Erard-Le to asphyxiation the from die to fish the caused bloom, A similar scenario 2 to occurred up of densities Cell recently high. were there levels in oxygen Galicia, cultured northwest 5000 Almost production. Spain, mucus increased and but irritation gill showed dissolved fish and Atlantic salmon, associated with a Aureoumbra lagunensis species. individual on and ecosystems aquatic on both effects significant in resulted Recent reviews have summarized the major aspects of the ecology and effects of these species (see Buskey Although here. provided is impacts their of summary 1987; Shumway 1990; Buskey and Stockwell, 1993; Montagna LANDSBERG2. Silicoflagellates REVIEWS IN FISHERIES SCIENCE siliceous skeletons was sufficient to cause mortality (Prego The absence of a siliceous skeleton in this phase therefore was not responsible for causing mechanical damage. Toxicological assays was oxygen dissolved of levels low that considered was on it and toxicity, demonstrate the naked stage failed to the most likely factor in the mortalities (Henriksen to 2.5

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 ., 1997). , was the Aureococcus Aureoumbra et al , several studies ., 1989; Gallager ., 1988; Bricelj and , whereas the mean (Street 2 2 et al et al (C. Onuf, submitt. in cells contains a bioactive a contains cells ., 1989). Because eelgrass Mulinia lateralis Aureococcus ., 1989; Gainey and Shumway, et al studies demonstrated that the was responsible for the demise et al ., 1990; Gainey and Shumway, 1991; Aureococcus In vitro Halodule wrightii fishery on Long Island, New York, USA, et al

(Dennison , 281 in 1994 (K. Dunton, unpublished data cited cells at high densities, rather than indigest- 2 was related to mussel mortalities attributed to attributed mortalities mussel to related was anophagefferens . A digitata . L Zostera marina and anophagefferens . Argopecten irradians A to less than 4000/m

., 1996). Light attenuation due to the brown tide bloom was also diet to support gametogenesis (Bricelj and Lonsdale, 1997). et al has been occurring since January 1990. Zooplankton populations de- in 1992 2 ., 1990). In 1985 and 1986, ., 1996). In Laguna Madre, seagrass-shoot density declined from 8000 to 10,000 Persistent brown tides in Long Island reduced light availability, which resulted Based on experimental evidence (Tracey, 1988; Bricelj of bloom monospecific nearly persistent, a Texas, Madre, Laguna In ., 1989; Bricelj and Kuenstner, 1989), Bricelj and Kuenstner (1989) suggested that suggested (1989) Kuenstner and Bricelj 1989), Kuenstner, and Bricelj 1989; ., in Buskey responsible for declines of the seagrass 1991) have demonstrated that inhibitory effects on bivalve feeding require direct cell direct require feeding bivalve on effects inhibitory that demonstrated have 1991) metabolites in cell-free filtrates of intact or contact and are not elicited by dissolved lysed cells. in a decline in eelgrass, (Tracey, 1988; Ward and Targett, 1989; Gallager provides shelter and substrate for numerous estuarine benthic eelgrass species, habitat may have contributed a to poor recruitment loss of bay scallops in and the slow recovery of the stocks (Tettelbach and Wenczel, 1993). A decline of the saccharina Laminaria lagunensis clined following the outbreak of the bloom (Buskey and Hyatt, 1995). Prior to the onset of the Texas brown tide, dominant benthic organism, with the densities as high dwarf as 6000/m surfclam, abundance (Buskey of invertebrates the adult to species toxic is after tide the brown Texas bloom the declined that evidence to no 24/m is There et al shoots/m of the bay scallop, Bricelj and Lonsdale, 1997). As discussed by Bricelj and Lonsdale (1997), although specific cell toxins have not been identified from tide initiation and negatively were bivalves, and polychaetes maintenance of those including larvae, meroplanktonic (Bricelj and Lonsdale, 1989). Fofonoff, and (Smayda concentration tide 1997). brown with correlated Abundance of reduced the feeding abundances activity the lowered of and filter-feeding drum, black shellfish and red by in success inhibiting egg-hatching depressed ciliary action, of zooplankton, shellfish, and other benthic filter et feeders al (Tracey, 1988; Draper Tettelbach 1988; (Tracey, failure recruitment subsequent and starvation inducing by and Wenczel, 1993). Economic losses to the bay scallop fishery of million New York $2 State at estimated were tide brown to attributed landings reduced the by caused per year (Kahn and Rockel, 1988). Recruitment failure may have been caused by gamete resorption in reproductive adults Aureococcus (Tracey, 1988) or the inability of an al et chronic toxicity of quality may be responsible for the detrimental ibility, small size, or poor nutritional effects and mortalities observed in bivalves. of layer polysaccharide diffuse, extracellular, compound, which is released by amylase digestion (Tracey frequency beat ciliary lateral the in reduction for responsible is that 1989), Kuenstner, of isolated gills of some bivalves (Draper brown tide because of the loss of suitable molluscan substrate for kelp attachment (Smayda and Fofonoff, 1989) Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , was , , eggs (Cho, (Buskey and bloom. Low

cells/ml), and Ceratium 2 . Microzooplankton 10 × /ml after the onset of 1 Spisula solidissima Spisula ., 1996). 10 Cynoscion nebulosus (1.36 (1.36 × Ceratium tripos et al cells/ml), but the bloom also 3 Acartia tonsa tripos . 10 C ., 1980; Onoue, 1990) (and see below see (and 1990) Onoue, 1980; ., × et al et (2.8 282 cells/ml), 2 /ml to less than 3.0 furca . 2 C 10 10 ., 1996). Spotted seatrout, × × et al to 2.0 (2.72 ., 1996). No adverse effects of brown tide on adult fish or 1 10 et al × spp. cells/ml showed significantly reduced hatching rates and 2- to 3-day ., 1996). Because seagrasses stabilize sediments, loss of seagrass 6 lineatum . . An estimated loss of 143,000 metric tons of surfclams was attributed to 2 C 10 et al × In August 1987, reports of discolored brown water in Norway were confirmed included the bloom. Although some mortalities of benthic fish were reported, the principal effect was to alter established migration and distribution patterns, with recovery of and (Mahoney bloom the of decline the after months some occurring stocks fish the Steimle, 1979). to be a bloom dominated by a. Ceratium Mass mortalities of of species several aquatic of blooms of result a organisms as developed that conditions (Table 2) have been blamed on anoxic 1. Dinoflagellates V. HARMFUL Ultimately, SPECIESquality. water reduce to potential the has bloom concentrated highly Any ANDand conditions anoxic to contribute can combined cells respiring and decomposing WATER Although the discussion of environmen- the subsequent formation of toxic sulfides. QUALITYtally associated effects of HABs is beyond the scope of this communication, some examples of large-scale mortality events involving nontoxic species are provided below. under mixed species). In summer 1976 in the New York Bight, USA, a large-scale surfclams, Atlantic of especially animals, marine of mortality (ciliates, copepod nauplii, and rotifers) population abundance changed from proximately ap- 2.0 the bloom. Microzooplankton grazing rates on phytoplankton standing stock were reduced from approximately 95% to less than 5% during the bloom Stockwell, 1993; Buskey rates Feeding hatching. after day third the to survived fish the of 20% Only survival. of larval spotted seatrout on rotifers in brown tide water reduced were when also compared considerably to those in nonbloom water (J. Holt, cited unpublished data in Buskey Nicholls 1979; Steimle, and Mahoney 1979; associated with the decline and ultimate collapse of a levels of dissolved oxygen and the contributed subsequent to production the of mortality. hydrogen Oxygen sulfide depletion extended 13,000 km over an area covering LANDSBERGBuskey REVIEWS IN FISHERIES SCIENCE macrobenthic communities were apparent (Buskey contributes contributes to resuspension of sediment particles, leading to further light reduction and subsequent further losses of seagrass. There were also noticeable declines in mesozooplankton populations, largely due to reduced feeding, growth, and fecun- dity of dominant plankters such as the copepod from laboratory spawns that were placed in brown tide having concentrations of 1 to 1.6

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , to 3 , on (see 10 × furca . polygramma C . G Jasus lalandii cells/ml. The mass 4 . 10 Seriola quinqueradiata Seriola × in the plankton samples Ceratium cells/ml (depth 0 to 15 m), 2 sp., were also present in the 10 × Ceratium furca fraction isolated from the fire sponge, cells/ml, red seabream were extremely β 3 ., 1985) indicated the possible role of an 283 Rhizosolenia 10 × . In March 1982, 200 tons of ichthyoplanktonic et al exceeded 6 C, salinity 32 ppt), but tilapia survived for more o was considered to have produced the toxin found toxin the produced have to considered was and cells/ml). There were some reports that farmed 2 , died in Carenero, Venezuela. Analysis of the fish fusus -associated aquatic mortalities are attributed to poor 10 furca . × Ceratium . C C ., 1985). However, other potentially toxic or harmful species, (3.4 Ceratium et al cells/ml, red seabream died within 56 to 58 min (dissolved oxygen (dissolved min 58 to 56 within died seabream red cells/ml, in False Bay and Walker Bay, South Africa, in April-May 1962. More than More 1962. April-May in Africa, South Bay, Walker and Bay False in

. Based on the dominance of , and excess mucus and marked edema were observed in the secondary the in observed were edema marked and mucus excess and , red tide. Thirty-nine thousand juvenile yellowtail, juvenile thousand Thirty-nine tide. red fusus . Cetengraulis edentulus C Pseudo-nitzschia seriata

4 fusus fusus . . C 10 C Although most In April 1997, a large stranding (1500 tons) of rock lobster, During March 3 to 6, 1990, fish farms in Kagoshima Bay, Japan, were affected × the South African west coast followed the decay of a massive bloom of bloom massive a of decay the followed coast west African South the index, feasibility a on and reported that was 100 tons of marine organisms, including more than 100 species (Table 2), were killed as a result of the bloom. Cell concentrations were estimated at 1 sardines, Tedania ignis in the fish (Mijares such as kill. the for responsible been have may species what unclear is it so samples, plankton Because toxins were only verified from field-tested mixed clonal plankton cultures, this and report should not be considered from inconclusive. also section on mixed species). Exhaustion of nutrients and eventual decay of the bloom inshore led to oxygen depletion throughout the water column. As a conse- quence, animals concentrated in the shallow in resulted strandings further weeks, surf few next the During conditions. zone bottom anoxic in an attempt to escape lobster rock Because million. $50 at valued lobsters rock of tons 2000 of loss total a of the the regional fishery is likely to be are slow-growing and long-lived, recovery slow (Pitcher, 1998; Pitcher and Cockcroft, 1998). water quality, one report (Mijares uncharacterized toxin from indicated the presence of a high concentration of isolated paralytic was toxin absorbance that UV killed and mice weight by molecular same the with toxin A injection. i.p. both from extracts toxic The kill. fish the of site the at collected samples plankton from fish and plankton were identical to the mortality was attributed to the large biomass of the bloom, subsequent death and decay b. Gonyaulax polygramma with associated be to documented ever mortality largest-scale the Probably died when the density of 3.2 for a longer period, there were reports of mortality of both large (6 to 8 kg) and small salmonids stopped eating, or had a reduced and appetite, and when fish were exposed kg) 8 to (6 large both of mortality of reports were there period, longer a for fish. Slight lesions were present in the gills. Symptoms were attributed to the (Tangen, 1988) (Table 2). a by than 3.5 h. At cell densities of 3.6 lamellae. Onoue (1990) suggested that mortalities occurred because of suffocation induced by excess mucus production in response to Prorocentrum Prorocentrum micans resulting in an economic loss of $462,000. In laboratory exposures to 9.8 5 to 6 ppm, temperature 18 to 19 prostrated, but no death ensued after 1.5 h of exposure. Fish gills became clogged with Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 6 10 × ., 1996). ., cells die et al et Noctiluca red tide in Uwajima in tide red developed along the blooms (Table 7), and cells/ml to 0.1 cells/ml ., 1988, cited in Elbrächter in cited 1988, ., 4 10 et al et × cells/ml. A substantial shellfish polygramma 3 . Noctiluca G 10 bloom can create a large quantity of × at maximum cell densities of 5.3 blooms can affect the concentrations of Gonyaulax spinifera 284 Noctiluca Noctiluca Noctiluca bloom. The most serious fish kill resulted in a loss of 35 is heterotrophic, large blooms can consume significant quan- Noctiluca cells/ml. During the red tide, oxygen-deficient water — and ultimately, and — water oxygen-deficient tide, red the During cells/ml. 4 10 × . High concentrations of ammonia, nitrogen, and phosphate were measured just measured were phosphate and nitrogen, ammonia, of concentrations High . 3 From late August until mid-November 1994, a 1994, mid-November until August late From In May 1988, large-scale fish kills and two incidents of shellfish mortality in Tolo in mortality shellfish of incidents two and kills fish large-scale 1988, May In and Qi, 1998) and fish kills occur (Okaichi and mortality Nishio, of demersal 1976). fish In and benthic July organisms 1986, along the a east mass coast of Jakarta Bay, Indonesia, coincided with a bloom of and lyse, accumulated ammonia is released (Schaumann (Schaumann released is ammonia accumulated lyse, and levels of dissolved oxygen (Subramanian, 1985; Ho and production Hodgkiss, has been 1992). associated with Ammonia older blooms, because when cells/m tities of oxygen, or their subsequent decay and bacterial production can result in severe in result can production bacterial and decay subsequent their or oxygen, of tities low to attributed been have kills fish Several 1998). Qi, and (Elbrächter depletion oxygen west coast of Vancouver km, 100 as far as offshore stretched Island, and km 400 than more for alongshore extended British Columbia, Canada. Recorded The Columbia. British in record on tide red extensive most bloom, the time the at was which surface cell concentrations were as high as 9 mortality believed to be due to hypoxia occurred in Barkley Sound (Forbes, 1990). d. Noctiluca scintillans Several mortality events have been attributed to mucus, which mechanically damages (Subramanian, 1985). In other cases, fish gills or interferes dissolved oxygen with or ammonia (Suvapepun, fish 1989), which indirectly respiration contribute to fish kills. Because anoxic conditions — and the high sulfide and ammonia (Koizumi mortalities large-scale concentrations the to contributed decomposition bloom caused by c. Gonyaulax spinifera of During August 1990, a massive bloom water adverse to contribute significantly can blooms these toxic, considered not although quality. In some cases, such conditions may not result in mortality but in avoidance by fish, leading to localised reductions in fisheries (Bhimachar and George, 1950; Ogawa and Nakahara, 1979; Devassy, 1989). A subsequent to the fish kills. The high water pH (8.85) caused by the photosynthetic the by caused (8.85) pH water high The kills. fish the to subsequent which ammonia, un-ionized in increase an to contributed likely bloom the of activity is also known to stress fish (Lam, 1988; Lam and Yip, 1990). Bay, Japan, caused recorded densities mass cell Maximum mortalities yen. million 800 of than more cultured worth were and that 2) (Table wild fish and shellfish stocks 6.8 were LANDSBERGof the plankton, and the severe depletion of oxygen in the water (Grindley and Taylor, 1962, 1964). Harbor, Hong Kong, China, were associated with anoxic conditions caused Gonyaulax by polygramma a REVIEWS IN FISHERIES SCIENCE tonnes of fish, estimated at $Hong Kong 7 million. A thick mucus layer was noted on the gills of dead fish. The collapse of the bloom was coupled with the onset of the summer destratification and depletion of oxygen in the bottom waters. Before the water column below levels were virtually zero throughout the fish kills, oxygen a depth of two meters. The bloom declined from 5

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 and reached 2 Dinophysis ., 1991). and cells/ml, although et al 3 Ceratium furca 10 × , eggs laid on gravel and marl ) Hansen and Moestrup (Daugjberg Moestrup and Hansen ) Alexandrium catenella Alexandrium 285 , that were living in water 0 to 3 m deep. Mouse deep. m 3 to 0 water in living were that , cells/ml. The presence of the bloom resulted in Clupea harengus 2 onto the egg layer. The oxygen level of 2.0 ppm 10 × mol/l. A complete absence of rock lobsters and other µ cells/ml were also recorded. A 30-km stretch of St. Helena Perna 3 bloom that covered an area of approximately 3 km 3 approximately of area an covered that bloom , which is capable of switching to anerobic metabolism under 10 Gymnodinium sanguineum Gymnodinium × (= , with lower concentrations of concentrations lower with , scintillans . , made up the bulk of the dead fish (1250 tons), with the remainder being remainder the with tons), (1250 fish dead the of bulk the up made , N Skeletonema costatum , and 1500 tons of fish, comprising about 50 species, washed ashore. Mullet, ashore. washed species, 50 about comprising fish, of tons 1500 and , . Average cell concentration was approximately 1 Siphonaria capensis ., 2000) occasionally has been associated with fish and shellfish mortalities . The mortality was associated with a heavy precipitation of organic material largely material organic of precipitation heavy a with associated was mortality The . 2 1. Dinoflagellates ridges at a depth of 18 m was noted along the south coast of the Isle of Arran, Scotland, Arran, of Isle the of coast south the along noted was m 18 of depth a at ridges 163,000 approximately be to estimated was patch spawn the of area The 1990. April in m W. SUSPECTED SPECIES WITH UNIDENTIFIEDa. Akashiwo sanguinea (=Gymnodinium MECHANISMSsanguineum) sanguinea Akashiwo acuminata 2. Diatoms a. Skeletonema costatum A mass mortality of Atlantic herring, composed of measured in water taken from under the egg mortality indicated mat anoxia as midway the likely through cause of the death (Morrison period of et al after the mortality (Adnan, 1989). From July to September 1988 in an Venezuela, Punta Patilla Bay, population densities as high as 5 of the gills showed that the mussels were not toxic. An examination bioassays revealed suffocation to contributed likely that substance mucilaginous a by covered were they that of the animals (La Barbera Sanchez, 1991). e. Mixed species — Ceratium furca/Prorocentrum micans In March 1994, the worst marine mortality ever subsequent recorded and in South entrapment Africa the occurred by along caused was event The Bay. Helena St. in coast west the decay of a mixed phytoplankton bloom dominated micans Prorocentrum by nontoxic concentrations up to 7 anoxic conditions. The mortality was caused by suffocation due to oxygen depletion and toxicity due to hydrogen sulfide poisoning. Oxygen concentrations were maintained at anerobic by generated sulphide hydrogen and bay, the of waters bottom the in ppm <0.5 bacteria was recorded at lobsters juvenile 50 of recruitment and event, the after month 1 recorded was life invertebrate did not occur until seven months later (Matthews and Pitcher, 1996). Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS mussels, of mortality 100% a lobster, rock of tons 60 Approximately fauna. marine dying and dead with littered was Bay lalandii Jasus richardsoni Liza fish. Surveys of the rocky shores revealed that dominated by sharks and bottom-dwelling most mussels, , sea urchins, and other intertidal life had died, except for the false

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . A and (79%), ., 1996). Akashiwo Akashiwo appeared sanguinea sanguinea Callinectes Crassostrea . . ., 1996), and 1996), ., et al A A (reported as ., 1999a) may cells/ml) of 2 et al et was significantly were significantly were ., 1996). et al 10 × et al sanguinea . A sp. densities in juvenile oysters juvenile in densities sp. Acartia tonsa from the lagoon at Obidos, on Obidos, at lagoon the from Polydactylus octonemus Polydactylus

, ., 1992; Lee Calanus pacificus Calanus ., 1979, cited in Shumway, 1990). A Vibrio , and et al and et al 286 layer (24 m depth) than in water from 10 m. ) in coastal waters off southern California was pacificus Cardium edule Cardium . C ., 1996). The role of microalgae as potential vectors ., 1996). The role of microalgae as potential , et al splendens sanguinea . . parvus A G has occasionally been associated with human shellfish . Paracalanus parvus Paracalanus has been considered to have either no effects on oysters P ) mortalities and disease outbreaks coinciding with coinciding outbreaks disease and mortalities ) ., 1992) was partially attributed to vibriosis (Lee vibriosis to attributed partially was 1992) ., bloom, it was noted that high that noted was it bloom, micans et al et can produce low concentrations of ROS (Kim

did not appear to be a primary etiological factor (Lee ), in Puget Sound (Cardwell may therefore influence bacterial loading and contribute to an increase (reported as was reported to be highly toxic to larvae of the Pacific oysters, Pacific the of larvae to toxic highly be to reported was . The kill was considered to be caused by hypoxic conditions in combi- , and Japanese littleneck clams, sanguinea . On several occasions there have been reports of oyster ( In June 1984, low-salinity water and a bloom of japonica A sanguinea sanguinea . . . gigas V with coincided sometimes that disease) oyster (juvenile syndrome recent (Bricelj blooms A suddenly off Galveston, Texas, Atchafalaya USA. rivers High between discharge March and from downcoast late current the preceded May the Mississippi- and appearance of a the low-salinity demersal bloom, strong, the of appearance water first the of week a Within and bloom. associated wind-driven, its threadfin, Atlantic by dominated numerically fish, began dying early in the morning and washing ashore. Mortalities included an crabs, blue and species, fish 16 comprising fish, million 13 estimated sapidus nation with the hydrogen sulfide produced by the nocturnal metabolism of the Guillen, and (Harper cells dinoflagellate dead of decay anaerobic by and bloom 1989). lower in individuals collected in the bloom layer than in those examined from three from examined those in than layer bloom the in collected individuals in lower other depths (Fiedler, 1982). virginica Crassostrea blooms (Table 2) (Nightingale, 1936; Bricelj sanguinea Akashiwo sanguinea (Wikfors and Smolowitz, 1994) or only indirect effects. In field observations during an suggest that a high biomass of the algae may initiate animals harmful more susceptible effects to and disease. render and in water samples from nursery floats coincided with the bloom. lower in water from the Mean gut fullness of sanguinea in bacterial pathogens (Lee for aquatic animal pathogens is discussed an below (see section important on HABs as aspect vectors). The recent in demonstration that HAB A dynamics and is poisonings or with aquatic mortality events, but potential toxins have not been cockles, toxic 1955, In identified. b. Prorocentrum micans Prorocentrum sanguinea actively avoided by macrozooplankton, whose vertical distribution patterns cated indi- several different patterns of active avoidance of the migrating dinoflagellate of rates Filtration layer. LANDSBERG(Table 2) or behavioral effects, but the definitive mechanism by which this species may be harmful is unknown. A dense, subsurface bloom (5.1 REVIEWS IN FISHERIES SCIENCE

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., ., For

sp.), et al et et al bloom micans . P was associ- ) (Pillet and in bioassays micans ., 1989; Martin

Palaemon Shumway, 1990). Tapes decussatus Tapes

et al micans sp.) caught in the . Artemia P ., 1990a, 1990b; Crawford Morone Prorocentrum , which caused slight PSP in has been implicated in several bloom (Grindley and Sapeika, bloom (Grindley et al ., 1956; Kat, 1984; Carver 1984; Kat, 1956; ., , and palourdes, palourdes, and , gave no evidence of toxicity or et al et Prorocentrum minimum micans micans . . P P micans 287 Donax serra . blooms on aquatic organisms have been sp.), and bass ( P Mytilus edulis Mytilus is a cosmopolitan species that occasionally spp., spp. and mixed species sections under Harmful Mugil have suggested that this species is nontoxic. blooms (Romalde was implicated when 40 to 50% of blue mussels were Mesodinium Ceratium Ceratium micans . P micans ., 1994). Although there is no evidence for toxin production,

Mesodinium sp.), mullet ( cells/ml) was coincidentally present in the lagoon and therefore and lagoon the in present coincidentally was cells/ml) ., 1993). 3 ., 1995). et al Mesodinium rubrum 10 et al × et al Portunus ., 1993) (see under section on blooms as vectors), poor water quality and ., 2000), and discoloration in molluscan digestive glands (due to phycoeryth- to (due glands digestive molluscan in discoloration and 2000), ., Prorocentrum In conjunction with (1.5 to 2.7 to (1.5 sampled from the same area were also toxic to mice. Prawns ( crabs ( time were nontoxic to mice. A lagoon at the same reaches bloom densities Crawford, 1985; Lindholm, 1981; Horstman, and (e.g., estuaries and zones, upwelling forms red tides in 1989; Cloern temperate coastal waters, the possible effects of discussed in some instances. These effects include the association of bacterial pathogens with 2. Ciliates a. Mesodinium rubrum The ciliate its adverse effects on marine fauna (Horstman, 1981; Hayes al et Pomeroy 1935; (Clemens, accumulation) rin et al the west the coast in sensitivity of loss including of symptoms, Neurological poisonings. numerous Portugal, were associated with of the legs, tremors, in the arms and hands, paraplegia lips and chin, numbness one human fatality lethal were extracts reported. and were feeling floating a and walking, ataxic mussels, blue addition, In mice. to October In 1956). Silva, and (Pinto producer toxin major the be to suspected was 1968, low-level toxicity in mussels and oysters in reported following a South Africa, was Algoa Bay, Port Elizabeth, 1969). In November 1973 in Dwarskersbos, South Africa, ated with toxicity in white mussels, to the symptoms. molluscs in stressors as blooms the of role potential a indicate may which 1996), (Crawford Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS killed in northern Brittany, France, but this mortality event was probably due to low levels of dissolved oxygen (Lassus and Berthome, 1988 in human consumers (Horstman, 1981). In these unclear what toxins were involved or what was the relationship with human poisoning cases, it is Most studies with example, in feeding experiments, sublethal effects in bivalves (see section on Houvenhaghel, 1995; Wikfors and Smolowitz, 1995) or to marine mortalities (see Species and Water Quality). Evidence for minimal ROS production (Kim (Demaret 1999a) may suggest that high cell concentrations could have significant effects on aquatic organisms, but further studies are required.

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , Karenia , Alexandrium ., 2001). Bacteria 2001). ., ., 1988, 1989), the et al et ., 1995; Doucette and Mesodinium rubrum et al , (Buck and Pierce, 1989; et al (Epstein, 1995). However, Gambierdiscus toxicus Gambierdiscus , ) to salmonids depended on the ., 1990a; Landsberg, 1997). There is a carterae et al Vibrio cholerae . 288 H ., 2001). Under experimental conditions, the Heterosigma akashiwo ., 1990a, 1990b; Bates , (as et al et al Pseudo-nitzschia multiseries Amphidinium carterae Amphidinium , ., 1997; Carrasquero-Verde, 1999; Hold 1999; Carrasquero-Verde, 1997; ., , and et al et ., 1995; Hold tamarense . ., 1998). In many cases, these bacteria are intracellular symbionts; in ., 1989; Romalde et al A , Heterosigma akashiwo et al et al Gymnodinium catenatum AND STRESSORS IN DISEASE , The presence of HAB-associated bacteria has been documented in several Aside from the possible role of HAB-associated bacteria in toxin production, strong possibility that HABs act as reservoirs and transmission pathogens in aquatic systems. In certain cases, agents there is a possibility that in addition for animal to stressing aquatic animals through physically harmful compromised to transfer can that pathogens bacterial mechanisms vector species HAB exposure, or by toxin be may HABs with associated bacteria pathogenic of biomass increased An animals. able to more readily infect (e.g., external through breached damage skin) HAB a caused to stressed subsequent or during by disease animals bacterial to organisms with HABs. aquatic of suceptibility There are documented cases for event (see below), but no increased study has been done to determine if the same bacterial other cases, they are extracellular but are strongly associated populations. with dinoflagellate microalgal species known to affect aquatic organisms, including real role of bacteria in the becoming production evident of (Rausch some microalgal de toxins Traubenberg Doucette is and only Lassus, now 1991; Doucette, 1995; lusitanicum Nayak 1995; Trick, are important determinants of bloom population dynamics and toxin has indicated (1) a strong role for interactions (Doucette, 1995). Recent information production metabolites bacterial to related is toxicity that and species HAB and bacteria between (Doucette, 1995) and (2) species a HAB the in synergism occurs than production between toxin higher much HAB in results that species bacteria and their symbiotic alone (Bates toxicity of are bacteria HAB-associated some that possibility the to paid been has attention little also aquatic-animal pathogens (Romalde presence of bacteria in cultures — axenic cultures were nontoxic to fish (Carrasquero- fish to nontoxic were cultures axenic — cultures in bacteria of presence Verde, 1999). The role of bacteria in toxin production will not be discussed further here, but the importance of HAB-associated bacteria should always be considered with respect to toxic effects on aquatic organisms. (Doucette, pathogens human for vectors as act to HABs for potential the also is there vibrios, for vectors as HABs of role potential the to drawn been has Attention 1995). especially for human pathogens such as A. HABS AS POTENTIALThe association of bacteria and other symbionts VECTORSwith HABs has been many years, and known several components of HAB-microbial interactions for play an impor- tant role in HAB ecology. Although such a role was postulated in the 1960s (Silva, 1962) and later demonstrated by a few researchers (Kodama Ostreopsis lenticularis LANDSBERGVI. HABS AS POTENTIAL VECTORS FOR PATHOGENS REVIEWS IN FISHERIES SCIENCE Tosteson brevis

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., ., ., et al et et al et et al et ., 2001). spp. in the et al anguillarum . V , Pfiesteria ., 2001a; Glasgow 2001a; ., ., 2000) and is responsible is and 2000) ., et al et et al et ., 1999; Vogelbein (Buck and Pierce, 1989; Romalde Vibrio alginolyticus Vibrio et al , and amoebae) and bacterial pathogens 289 (Blazer causes mechanical damage to fish skin during ., 1996) and have been involved in several HAB- cells or to water filtrate (and presumptive toxin) that et al Aeromonas hydrophila Pfiesteria is a primary pathogen (Kiryu pathogen primary a is ., 2001). Found predominantly in the Atlantic menhaden, showed behavioral changes and sloughing of the fish Uronema marinum and , , along the eastern seaboard of the United States, UM is caused et al P. shumwayae Aphanomyces invadans and other microalgal species that produce bioactive compounds may piscicida . or ., 1993; Riquelme ., 1996). That P et al et al ., 1990a), are known fish and shellfish pathogens (see Austin and Austin, 1987; Heavy mortalities of more than 20 species of tropical reef fish were considered Pfiesteria parahaemolyticus, piscicida . . Brooklynella hostilis There are several documented examples in which either acute or chronic exposure to microalgal toxins or harmful mechanisms associated with HABs were sufficient to increase the susceptibility of aquatic organisms to disease. to be induced by the chronic dinoflagellates inhabiting exposure coral reef areas (see section of on Caribbean ciguatera). the In in 1980, fish 1993 widespread were fish to tropical of mortalities toxins mass 1997, and 1994, produced to by benthic 1993 the in months several of period a Over 1995). (Landsberg, USA Florida, in and to 1994 event in Florida, fish were affected extensively by a number ( of parasites species involved in the example, disease for HABs, were from isolated species those bacterial vectored by the HAB. Some of V the B. HABS AS STRESSORS IN DISEASE that are known to induce mortalities in fish. pathogens There were was the no evidence principal that cause these invaders of of fish whose the health had already disease been compromised. The fish but displayed rather numerous were behavioral signs secondary (see ciguatoxins) before dying. The precursor disease to was postulated this to be chronic exposure to microalgal toxins that acted immunosuppressants as and increased the susceptibility of fish to disease. At certain times, when fish are exposed to benthic toxins through dietary consumption, they become weak and susceptible to certain pathogens. In this scenario, it was consid- dinoflagellates, benthic from toxins to exposed likely than more were fish that ered some of which are involved in ciguatera (Landsberg, 1995) (see ciguatoxins). diseases. Fish exposed in aquaria directly to have a potential role in aquatic animal P Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS et al Thune related disease interactions (see below). Some aquatic mortality and disease events therefore may be strongly interrelated with pathogenic microbe and HAB associa- tions, instead of separately pathogens, triggered the currently accepted model. by either toxic microalgae or infectious contained epithelium, which led to invasion by numerous secondary, opportunistic pathogens opportunistic secondary, numerous by invasion to led which epithelium, (Noga pedunculate feeding, which (Vogelbein could development lesion of mode allow possible a is for pathogens, opportunistic subsequent invasion by secondary 2001). There has been considerable debate about the role of (Burkholder (UM) mycosis ulcerative as described lesions fish of development 1998; Dykstra and Kane, 2000; Noga, 2000; Burkholder 2000; Noga, 2000; Kane, and Dykstra 1998; 2001b; Vogelbein Brevoortia tyrannus by a fungus, Aphanomyces invadans Aphanomyces

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ) et sp., spp. ., 2000; kisutch ., 1998). . Pfiesteria have been O et al et al Spirulina concavicornis Zoothamnium, . , the cumulative the , , C Chaetoceros is complicated by Penaeus orientalis to the water greatly water the to convolutus ., 2001b), the extent to . C ., 2000, 2001b; Burkholder 2001b; 2000, ., et al cells/ml of cells/ml increased the mortality rates mortality the increased and coho salmon, coho and 1

concavicornis and , et al et . Aphanomyces 10 C × Leucothrix mucor for 7 days died. Of the 37 coho and the presence of presence the and bloom in the Pei Hai Sea (Northern Sea Hai Pei the in bloom ., 1993). At sublethal concentrations and added on day three had died by day Vibrio anguillarum Vibrio sp. in the water column, benthic layers, benthic column, water the in sp. et al convolutus . C 290 cells/ml of cells/ml 1 blooms also harmed the commercial farming ., 1998; Glasgow and Pfiesteria anguillarum 10 ., 1996; Glasgow Glasgow 1996; ., . × Nodularia V anguillarum et al . et al et V ., 2001), and is suspected in the development of fish Noctiluca Noctiluca scintillans Noctiluca ., 1989; Albright Chaetoceros concavicornis Oncorhynchus tshawytscha Oncorhynchus Oscillatoria corakiana et al sp., and sp., et al concavicornis . cells/ml of in Australia (Smith, 1996). Prawns started to show symptoms ., 1993). Although the mechanism(s) by which by mechanism(s) the Although 1993). ., 6 C has been shown to initiate lesions in fish exposed under

, cells/ml of 6 10 et al et × 10 spp. None of the 37 untreated control coho salmon or of the 36 coho 36 the of or salmon coho control untreated 37 the of None spp. by decreasing frond production and causing the macroalgae to putrefy to macroalgae the causing and production frond decreasing by × Oscillatoria ., 2000). Pfiesteria et al sp., ., 2001a; Vogelbein Planktonic blooms of blooms Planktonic The harmful diatoms During 1989 to 1990, a a 1990, to 1989 During Chaetoceros Laminaria ., 2001). The relationship between ., 2001). The relationship mortality was 38% during the same 7-day treatment period. However, 100% of the 3.7 to 1.4 with water in maintained 37) = (N coho (Albright seven increased the susceptibility of salmonids to bacterial diseases was not determined, this critical role as a stressor is clearly another important factor whereby HABs can significantly influence the economic success of aquaculture operations. Lyngbya or surface mats were coincident with four mortality episodes of cultured prawns, Penaeus monodon 4 months after stocking. Initially, consumption decreased, growth slowed, and they had difficulty prawns in molting. By the were lethargic when filaments time the ponds were dominated by benthic or floating mats or planktonic handled, food of Oscillatoriales blooms, prawns were found Visually, dying sick prawns had at dark body color, the limbs that were edge often red, muscle of that the was ponds. white, gills that were sometimes fouled (by an almost coincident geographical distribution of these genera and of fish with UM along the eastern coastal United States. However, there are areas where (Landsberg present, and fish with UM are still found is nontoxic, or not mariculture operations. Poor water quality caused an increase in the susceptibility of prawns to parasitic infections, which led to disease, mortalities, and additional losses valued at $1 million. of and spoil (Chen and Gu, 1993). 3.7 to 1.4 with water in maintained and with 1.7 et al lesions in the field (Burkholder Although Although (Noga conditions experimental which these lesions are the same as those caused by UM is minimal (Vogelbein al Lewitus China Sea) resulted in losses of about $100 million to shrimp ( directly implicated in mortalities of mechanisms). In aquatic addition, cultured salmonids became noticeably more susceptibile organisms (see section on to harmful two major bacterial diseases when they were exposed to low concentrations of these diatoms (Speare cells/ml, 5 to 0.4 of cultured in seawater in net pens die from vibriosis and/or bacterial kidney disease adding conditions, laboratory Under (BKD). LANDSBERGfor UM in other fish species and in other geographical regions (Lilley REVIEWS IN FISHERIES SCIENCE at which chinook salmon, chinook which at accelerated the mortality rate of coho salmon exposed to sublethal concentrations of treated with 2

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . , . nauplii harveyi . V , II, and other was associated

Artemia , Cyprinus carpio Spirulina subsalsa alginolyticus in extracts from the same anguillarum . . V V , I, Artemia Penaeus stylirostris 291 proteolyticus ). The hepatopancreas was small and pale, . spp. anguillarum V . V , . Immersion of Vibrio spp. ( Nitzschia , and the fact that water was toxic to were considered to be the major contributing factor to Vibrio japonicus ), and shells that were thin and covered by fouling organisms fouling by covered and thin were that shells and ), . Artemia P parahaemolyticus and . V , monodon . An unusual case of disease in blue shrimp, In August 1976, in a fish farm in southern France, extensive algal blooms of P vulnificus . Nitzschia, Oscillatoria Nitzschia, (i.e., barnacles, bacteria, and with a cyanobacterial bloom. Epizootics occurred in on six blue different shrimp occasions between in 1975 Sonora, tanks and Mexico, and 1977 or resulted in in high raceways Puerto rates Peñasco, of mortality (up to mortalities 85%). was Each preceded of the by, or concurrent with, a bloom of Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS and the hind guts were empty or contained watery feces. Affected ponds had higher had ponds Affected feces. watery contained or empty were guts hind the and farm. the near water estuarine did than ammonia and phosphorus dissolved of levels Levels of presumptive V prawns sick of tissues muscle and gills, hepatopancreas, the in higher were vibrios) with ponds in prawns affecting was vibriosis that indicating prawns, healthy in than blooms of Oscillatoriales. The indicated levels that although of mortalities were Vibrionaceae caused by secondary found bacterial subacute infection, in toxicity was water due pond prawn the to that showed the bioassays tissues mouse Oscillatoriales episodes, bloom three first was the In the disease. primary cause of the pond Boiled symptoms. neurotoxic displayed mice the although nontoxic, clinically water and extracts from a tank culture of benthic Oscillatoriales caused mortalities of After the sudden collapse of the bloom, activity, decomposing and there cells was stimulated a marked bacterial decrease in dissolved bloom oxygen increase the in after and ammonia. days About Ten a a surface. significant day water’s after the the at bloom gasping collapsed, and carp pond were the gathered to inflow the at on mucus excessive distention, abdominal had pond the from netted carp collapsed, the gills and skin, damaged and clumped gill and fins, and associated internal pathologies in petechial hemorrhages on the body lamellae, vascular congestion and most of the visceral organs. The quantities some intestine of in which undigested was lesions, skin food, had soft gas carp the bubbles, of and and 20% to yellow transparent, 10 mucus. approximately bloom, Twenty-one with the after days large cases penetrated deeply into the skin and muscles and occasionally body reached cavity. the multifactored At a has the that Europe time, in carp the of syndrome disease disease a was septicemia, hemorrhagic considered to be similar to infectious etiology. The disease was considered to be triggered ionized by ammonia sublethal associated levels with of bloom un- decomposition and creased susceptibility the of carp subsequent to disease in- (Seymour, 1980). tank culture also caused mortalities. Smith (1996) demonstrated that the Oscillatoriales the that demonstrated (1996) Smith mortalities. caused also culture tank blooms produced a water-soluble, heat-labile toxin and that mortalities were pos- sibly caused by a neurotoxin. Supernatant culture tank water, both unfiltered and filtered, was toxic to before a digestive tract was developed indicated that the toxin was water soluble. Smith (1996) proposed that sublethal possibly levels via of a toxin(s) neurotoxic weakened effect the that immune system, prawns, and that the prawns thus were susceptible reduces to secondary infections feeding behavior and impairs the of pathogenic bacteria such as Aphanizomenon flos-aquae the development of a disease syndrome of farmed common carp,

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 . has , were , have a alginolyticus . V blooms revealed and also include the include also and Vibrio alginolyticus subsalsa . S was blooming but not in tanks Hematodinium ., 1987; Wilhelm and Mialhe, 1996; filaments, it was suggested that their , and , subsalsa . et al S that are associated with coral diseases. 292 subsalsa . S Piscinoodinium , produces a weak toxin that is only mildly toxic to shrimp, bloom, were occurring. Although there was no discernible was absent. Stomach contents of both apparently healthy and species are dinoflagellate endoparasites of commerically impor- commerically of endoparasites dinoflagellate are species Phormidium/Oscillatoria ., 2000). Affected Alaskan Tanner crabs, subsalsa subsalsa . . S S as a dietary supplement for 3 weeks, 2 out of 19 shrimp examined had subsalsa were often found attached as epibionts on the gill lamellae, accessory gill accessory lamellae, gill the on epibionts as attached found often were . et al S Hematodinium subsalsa subsalsa . . VII. PARASITESdefinitions most from excluded are pathogens ANDand parasites microalgal Traditionally, PATHOGENSof HABs. For the purposes of this review, they are discussed, but only to refer to those species that have substantial impacts aquatic (either resources. ecological These or species economic) affect on operations marine and include fauna, fish fisheries, and or shellfish ocellatum Amyloodinium aquaculture pathogens such as the dinoflagellates cyanobacteria in included not are events mortality aquatic with association their and genera These Table 2. 1994). (Shields, crabs and lobsters as such macrocrustaceans tant been associated with meats the of spoilage to mass due products fisheries of marketability mortalities, the in loss or mance, pathology, decreased swimming perfor- (Newman and Johnson, 1975; Meyers Stentiford in which green algae were dominant and protected from (2) potential the toxins by absence a chitinous of cuticle, lesions and for necrosis in resulted that midgut example, the in in Lesions rectum. and hindgut, the esophagus, tissues stomach, interruption of the lining epthelium allowed a route of entry for Because feeds medicated with antibiotics (normally effective were against vibrios vibrios) the were that concluded was it mortality, stopping in effective completely not secondary invaders and not the agents that digestive tract caused (Lightner, 1978). the lesions observed in the LANDSBERG Epizootics of the disease syndrome were not observed in other tanks or raceways in which REVIEWS IN FISHERIES SCIENCE hemocytic lesions in the anterior conditions, midgut. It was concluded that under certain moribund shrimp from shrimp populations affected by presence reduced respiratory efficiency. S When healthy shrimp were fed even when relatively large quantities of the hemocytic alga the are of consumed. epizootics Observations of that occurrence the (1) included hypothesis this supported enteritis syndrome in systems in which histopathological effect from the that the shrimp were ingesting large septicemic bacterial infections, amounts principally by a strain of of this alga. Shrimp documented dying in every epizootic. from Affected shrimp with the disease syndrome in its final stages had whitish, opaque abdominal musculature, a slightly pale cuticular coloration, and often an empty midgut. Many had early or sublethal lesions in the digestive organs. In the most common lesion, there was necrosis of epithelium the mucosal of the midgut and a S consequent hemocytic infiltration. Filaments of losses, presumed to be pleopods of shrimp from tanks in which processes, and the due to the

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 , ., et al partially

Aristhichthys Amyloodinium Hematodinium Piscinoodinium consists of three ., 1991; Landsberg infection ranging 0 to et al ), bighead carp ( carp bighead ), ., 1998). Amyloodinium sp. was considered to be et al sp. An examination of dying crabs ) were infested by infested were ) , in southern Brittany, France, were Hematodinium ., 1994, 1995) and, rarely, in the wild ., 1993). In June 1986, unusual mortalities 293 can build up in several days. Stressed fish . This bitter flavor results in an unmarketable an in results flavor bitter This . fishery in France (Wilhelm and Mialhe, 1996). Mialhe, and (Wilhelm France in fishery et al et al Leptobarbus hoevenii Leptobarbus Hematodinium Hematodinium ruber Necora ruber . N is a nonspecific, marine fish ectoparasite (Brown, is known from freshwater systems, where occasionally where systems, freshwater from known is is a nonspecific dinoflagellate ectoparasite and has a Punitius gonionotus Punitius ), jelawat ( jelawat ), Amyloodinium Hematodinium was highly susceptible and suffered mass mortalities. Clinical mortalities. mass suffered and susceptible highly was fishery in Scotland (Field

, ., 1987, 1990; Love ., 1991; Landsberg et al et al Piscinoodinium gonionotus . , P ), and lampam jawa ( jawa lampam and ), Amyloodinium ocellatum ., 1994, 1995, Kuperman and Matey, 1999). Although it has been postulated that postulated been has it Although 1999). Matey, and Kuperman 1995, 1994, ., pillulare Piscinoodinium Ctenopharyngodon idella Ctenopharyngodon pink carapace, chalky-textured meat containing hemolymph with a distinctly bitter product, flavor, which, for example, caused and an economic loss of approximately milky $176,000 to the Alaskan fishery in 1986. The bitter crab disease caused by affected as many as 95% of the Tanner crabs in southeastern Alaska (Eaton Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS 1991; Meyers of the swimming crab, attributed to heavy infections of revealed parasites in the hepatopancreas, 1100 gonads, from fell and catches crab muscles, 1988, and as 1984 Between well haemolymph. the as in infestations heavy tons to 48 tons, a decrease of 96%. Samples of crabs examined between June 1986 and Feburary 1992 showed prevalences of 87%. Based on this information, the of declines for responsible Similar conclusions were made with respect to Nephrops norvegicus declines in the Norway lobster, 1934; Brown and Hovasse, 1946) that can have devastating effects in aquaculture facilities. There are numerous reports of its incidence and pathogenicity to fish in mariculture and aquarium systems Laurencin, (Lawler, 1979; Paperna, 1980; 1977, Baticados and Quinitio, 1980; 1984; Aiello and Paperna D’Alba, 1986; and Noga Baudin- (Kuperman and Matey, 1999). The life cycle of stages: a trophont that feeds while attached motile and fish, to the from detaches trophont the after develops that skin tomont encysted and gill surfaces of fish, an flagellated dinospores that are released after the tomont divides. In closed systems, high concentrations of congregate on the surface, gasp rapidly for air, lose their appetite rapidly, and can fish on found are trophonts Typically, infected. heavily when days several within die after heavy mortalities (Lawler, 1980; Paperna, 1980, Noga et al et Lawler, and (Lom stages trophont feeding the by produced are compounds cytolytic the by caused is damage localized Extensive confirmed. been not still has this 1973), attachment of the trophont to the gills, rhizoid penetration, and feeding on the fish cytoplasm. In heavy infestations with epithelium becomes hyperplastic, trophonts and there is attached additional fusion of to the secondary the lamellae. Cellular damage and necrosis of the gills, gill tissue affect osmoregulation and the gill respiration. With respiratory function considerably impaired, fish die from asphyxi- trophonts the by induced damage mechanical the to response pathological The ation. ultimately leads to death (Paperna, 1980). it has been associated with mass mortalities of cultured fish. Like ocellatum fish of susceptibility the in differences be to appear there However, cycle. life similar species to infestation. In ( freshwater pond culture in Malaysia, grass carp nobilis only but

Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 is a ., 1983; et al corallyticum . P ., 1990). In other infestations of et al is toxic (Rutzler and Santavy, 1983), this trophonts induced epithelial hypertrophy, 294 Piscinoodinium Phormidium corallyticum water (e.g., molecular probes) and other technologies that will track HABs and provide capabilities for predicting their spatial and ances. temporal appear- Such and fisheries, information wildlife, health, public will endanger that situations to enable appropriately managers to respond quickly aquaculture and resource managers, and better integration among of all investigating animal HAB mortalities and disease knowledge and data in animal health and may be responsible for many currently “unidentified diseases” or “unexplained mortalities.” For example, mortalities enhanced been have may expression viral fact in when virus, a are to attributed often by chronic biotoxin exposure. Incidents to consider include morbillivirus Microalgae are also causing a series of diseases that negatively affect coral reefs. coral affect negatively that diseases of series a causing also are Microalgae 2. Increased interdisciplinary activity between scientists, diagnosticians, and 3.4. Strategies that are better integrated and geared toward “managing HABs” Recognition that chronic effects of biotoxins are likely to be very significant VIII. CONCLUSIONSOther than the classical “red tide” fish kills long chronicled by coastal dwellers, the begun just only has events disease and mortality animal in species HAB of role wider organisms these of presence the of implications broader The recognized. fully be to in aquatic systems around the world needs to years few be next the In elucidated. attitudes. and approaches new For of integration an be this must there to happen, it is hoped that the following will exist routinely: 1. Rapid screening techniques that will identify toxins in animal tissues and in aquarium fish, attached focal and diffuse hyperplasia, edema of the respiratory epithelium, lamellar fusion, and reduced respiratory efficiency (Ferraz and Sommerville, 1998). Numerous disease syndromes affecting several coral reef species throughout world are often named by the gross macroscopic description of the affected corals, the for example, black band, white band, red band, and white pox ( Rutzler and Santavy, 1983; Richardson, 1992). Recent studies suggested that several species of cyanobacteria (Table 1) may be diseases responsible (Rutzler for and black Santavy, and 1983; red band Richardson, suggested 1992). that Although it was earlier LANDSBERGsigns included a rust-colored appearance of the skin, lamellae. gill a the dense of fusion covering and of body, the mucus, on petechiae regions, dorsal and head dark If dissolved oxygen reached low levels (2.6 to occurred in 5.5 the ppm), ponds then (Shaharom-Harrison 100% mortalities REVIEWS IN FISHERIES SCIENCE has still not been proven, and it is still unknown whether these coral “pathogens” are toxic or produce bioactive compounds. Recent studies have suggested that the chemical environment of the microbial consortium, of which major component, contributes to anoxic and sulfidic conditions along the length of the infected coral where the black band occurs (Richardson, 1993).

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