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 Fish and Wildlife Conservation Commission, 100 Eighth Avenue Southeast, St. Petersburg, FL 33701. 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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 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS Reviews in Fisheries Science, 10(2): 113–390 (2002) The Effects of Harmful Algal Blooms on Aquatic Organisms Jan H. Landsberg Florida Marine Research Institute, Florida Fish and Wildlife Conservation Commission, 100 Eighth Avenue Southeast, St. Petersburg, FL 33701 TABLE OF CONTENTS Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 Abstract ...................................................................................... 114 I. Introduction ............................................................................... 115 II. Harmful Algal Blooms .............................................................. 117 III. Routes of Exposure................................................................... 117 A. Direct Exposure .................................................................. 171 1. Intact Cells.................................................................... 171 a. Intracellular Toxins ............................................... 171 b. Extracellular Toxins (Exotoxins) or Exudates .... 173 c. Cell Surface Contact ............................................. 173 2. Lysed Cells ................................................................... 174 B. Indirect Exposure ............................................................... 174 1. Trophic Toxin Transfer, Bioaccumulation, Biomagnification .......................................................... 174 IV. Impacts ...................................................................................... 175 A. Acute ................................................................................... 175 B. Chronic ................................................................................ 176 C. Organisms and Habitats Affected ..................................... 176 V. Toxins and Harmful Mechanisms ............................................ 189 A. Saxitoxins ............................................................................ 191 B. Tetrodotoxin ....................................................................... 204 C. Spirolides............................................................................. 205 D. Brevetoxins ......................................................................... 205 E. Gymnodimine ..................................................................... 213 F. Okadaic acid and Derivatives (Dinophysistoxins) .......... 213 G. Pectenotoxins...................................................................... 216 H. Ciguatoxins ......................................................................... 217 I. Yessotoxins ......................................................................... 223 J. Azaspiracid .......................................................................... 224 K. Hemolysins.......................................................................... 225 1. Dinoflagellates.............................................................. 226 2. Prymnesiophytes .......................................................... 232 L. Domoic Acid ....................................................................... 237 M. Anatoxins ............................................................................ 241 1064-1262 /02/$.50 ©2002 by CRC Press LLC 113 LANDSBERG REVIEWS IN FISHERIES SCIENCE 1. Anatoxin-a .................................................................... 241 2. Anatoxin-a(s) ................................................................ 242 3. Homoanatoxin-a .......................................................... 243 N. Mueggelone ........................................................................ 243 O. Cylindrospermopsin ........................................................... 244 P. Microcystins ........................................................................ 245 Q. Nodularins ........................................................................... 253 R. Reactive Oxygen Species ................................................... 255 1. Chattonella spp............................................................ 256 2. Heterosigma akashiwo................................................. 256 3. Olisthodiscus luteus ..................................................... 257 S. Species with Multiple Toxins ............................................ 257 1. Dinoflagellates.............................................................. 258 Downloaded By: [California Digital Library -CDL (CRC journals only) Consortium] At: 21:00 21 February 2007 2. Cyanobacteria............................................................... 260 T. Species with Uncharacterized Toxins or Bioactive Compounds ......................................................................... 261 1. Dinoflagellates.............................................................. 261 2. Diatoms ........................................................................ 271 3. Raphidophytes ............................................................. 273 4. Prymnesiophytes .......................................................... 273 5. Cyanobacteria............................................................... 275 U. Harmful Species that Cause Mechanical Damage ............. 278 1. Diatoms ........................................................................ 278 2. Silicoflagellates ............................................................. 280 3. Pelagophytes ................................................................ 280 V. Harmful Species and Water Quality ................................... 282 1. Dinoflagellates.............................................................. 282 2. Diatoms ........................................................................ 285 W. Suspected Species with Unidentified Mechanisms........... 285 1. Dinoflagellates.............................................................. 285 2. Ciliates .......................................................................... 287 VI. HABs as Potential Vectors for Pathogens and Stressors in Disease ...................................................................................... 288 A. HABs as Potential Vectors ................................................. 288 B. HABs as Stressors in Disease ............................................ 289 VII. Parasites and Pathogens ........................................................... 292 VIII. Conclusions ............................................................................... 294 Acknowledgments ..................................................................... 295 References.................................................................................. 295 ABSTRACT: This review provides an in-depth survey of the recorded incidences in aquatic organisms of mortality and disease events suspected or known to be caused by microalgal or ciliate blooms, their biotoxins, or their harmful mechanisms. Some 200 species of dinoflagellates, diatoms, raphidophytes, prymnesiophytes, silicoflagellates, cilliates, and cyanobacteria are currently known to be, suspected to be, or have the potential to be toxic or harmful to a wide spectrum of organisms. This review summarizes the current information on toxic or harmful microalgal species that affect aquatic organisms (and, when relevant, those that affect terrestrial organisms, including humans), provides an updated list of such species, cites pertinent case 114 Volume 10 (Issue #2) 2002 EFFECTS OF ALGAL BLOOMS ON AQUATIC ORGANISMS histories, and includes relevant information on harmful or toxic species from freshwater, brackish, and marine ecosystems. It is hoped that this review will provide documentation and reference material suitable