[Advances in Marine Biology] Advances in Marine Biology Volume 3 Volume 3 || Marine Toxins and Venomous and Poisonous Marine Animals

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<ul><li><p>A&amp; . mar . Bid., Vol . 3. 1966. pp . 265-384 </p><p>MARINE TOXINS AND VENOMOUS AND POISONOUS MARINE ANIMALS </p><p>FINDLAY E . RUSSELL Laboratory of Neurologiml Research. Loma Linda University. Loe Angela County Hospital. Lo8 Angela. California. U.S.A. and Depart- </p><p>ment of Zoology. University of Cambridge. England </p><p>I . Introduction . . . . . . . . . . . . . . . . A . Definitions . . . . . . . . . . . . . . C . B . History and Folklore . . . . . . . . . . ctenerel Chemistry and Zootoxioology of Marine Poisons </p><p>11 . </p><p>rn . </p><p>Iv . </p><p>V . </p><p>VI . </p><p>M . </p><p>MI . </p><p>Ix . X . </p><p>Protist8 . . . . . . . . A . Paralytio S h e W Poisoning B . Chemistry . . . . . . C . Toxioology . . . . . . </p><p>Porifera . . . . . . . . A . Poiaoning . . . . . . B . Chemistry and Toxioology . . </p><p>#lidaria . . . . . . . . A . VenornApparatus . . . . B . Chemistry and Toxioology . . C . ClinioalProblem . . . . </p><p>EOhhOdeRll8b . . . . . . A . VenomApparatus . . . . B . ChemistryandToxioology .. C . ClinioalProblem . . . . A . Venomous .. . . . . B . P d y t i o Shellfbh Poisoning c . VenomAppar8tut3 . . . . D . Chemistry and Toxioology . . E . ClinioalProblem . . . . </p><p>PohnousFishes . . . . . . A . Iohthyoeareotoxio Fishes . . B . Iohthyootoxio Fishes .. C . Iohthyohemotoxio Fishes . . </p><p>VenomousFishes . . . . . . A . Stingray . . . . . . B . W e e v e m . . . . . . C . Soorpionfbh . . . . . . D . Summary. Phyeiophermeoology </p><p>Aoknowledgmenta . . . . . . Referenoes . . . . . . . . </p><p>M O k l 8 0 8 . . . . . . . . </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>.. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. </p><p>266 266 267 269 262 263 263 208 271 271 272 273 274 278 281 283 284 287 290 290 291 293 296 297 302 304 306 329 329 330 338 346 357 368 369 369 </p><p>266 </p></li><li><p>256 FINDLAY E. RUSSELL </p><p>I. INTRODUCTION This review treats of the toxins of some of the more venomous and </p><p>poisonous marine animals of the world. For the most part, it is con- cerned with the chemical, zootoxicological and immunological proper- ties of the toxins, the animals venom apparatus, and the mechanism of envenomation. Some attention has been given to the general biology of the animals, and to the problem of the poisoning in man by the various forms. A second purpose of the review is to present an account of several of the more interesting problems in the emerging field of marine toxinology. </p><p>Approximately 1000 species of marine organisms are known to be venomous or poisonous. For the most part, these species are widely distributed throughout the marine fauna from the unicellular protistan, Gonyauh , to certain of the chordates. They are found in almost all of the seas and oceans of the world, and while their numbers may sometimes be quite large, they do not produce major ecological effects by virtue of their toxicity alone, nor are they other than a local danger to mans health and economy. It is generally believed that most of the venomous marine animals have been identified; although it must be conceded that a number of forms have not yet been adequakly des- cribed, and certainly our knowledge of the potentially dangerous deep- sea organisms is meager indeed. </p><p>A. Definitions It might be wise to consider a few words and terms that are peculiar </p><p>to toxinology. The term venomus animals is usually applied to those creatures which are capable of producing a poison in a highly developed secretory organ or group of cells, and which can deliver this toxin during a biting or stinging act. Poisonous animals are generally regarded to be those whose tissues, either in part or in their entirety, are toxic. In reality, all venomous animals are poisonous but not all poisonous animals are venomous. Animals in which a definite venom apparatus is present are sometimes called phnerotoxic (Gk. plivqxk, evident + T O ~ L K ~ Y , poison), while animals whose body tissues are toxic me called cryptotoxic (Gk. ~ p v m d S , hidden). The rattlesnake, stingray and black widow spider are venomous or phanerotoxic animals, while the blister beetles, certain puffer fishes and toads are said to be poisonous or cryptotoxic. </p><p>Although the terms venomozcs and poisonous are often used synony- mously, most investigators have tried to conhe the use of the term venomous animals to those creatures having a gland or group of highly </p></li><li><p>MARINE TOXINS AND VENOMOUS AND POISONOUS MARINE m s 257 </p><p>specialized secretory cells, a venom duct (although this is not a constant finding), and a structure for delivering the venom. While there haa been a tendency to employ the term venom apparatus to denote only the sting, spine, jaw, tooth or fang used by the animal to inject or deliver its venom, most biologists now use the term in its broader context, that is, to denote the gland and duct in addition to the sting or fang. Poisonous animals, as distinguished from venomous animals, have no such apparatus ; poisoning by these forms usually takes place through ingestion. </p><p>It does not include that type of poisoning which may occur following ingestion of fish contaminated by bacterial pathogens. Halstead (1964) haa divided ichthyotoxic fishes into three subdivisions : (a) Ichthyosarwtoxk &amp;hes-those fishes which contain a toxin within their musculature, viscera or skin, which when ingested produce deleterious effects. This type of poisoning is generally identified with the kind of fish involved : elasmobranch, chimaeroid, ciguatera, tetraodon, scombroid, etc. ; it also includes hallucinatory fish poisoning. (b) lchthyootozic J2he.s- those fishes which produce a toxin that is generally confined to the gonads. In these fishes there is a relation between gonadal activity and the production of toxin. Most members of this subdivision are fresh- water species. This group would include those fishes whose roe is poisonous. (c) lchthyohemotoxic $shes-those fishes which have a toxin in their blood. Some fresh-water eels and several marine fishes make up this group. </p><p>Fish poisoning is synonymous with ichthyotoxism. </p><p>B. History and folklore Few areas in biology have had their beginnings as steeped in </p><p>superstition and myth as have toxinology and the poisonous animals. The investigation of such complex substances as toxins, often capable of destroying life by complicated and sometimes undeterminable means, has by its very nature invited exaggeration, and sometimes pure fantasy. In early times the consequences of the bites or stings of venomous animals were often attributed to forces beyond nature, sometimes to vengeful deities thought to be embodied in the animals. To these early peoples the effects of venoms were so surprising, varied and violent that venomous animals and the injuries they inflicted were always shrouded with much myth and superstition. Even today considerable folklore about venoms still exists, and this is particularly conspicuous with respect to the methods of treatment for the injuries inflicted by venomous animals (Russell, 1961). The task of separating fact from </p><p>A.Y.B.-3 N </p></li><li><p>258 FINDLAY E. RUSSELL </p><p>fiction is often a formidable one, and one not always lightened by the passage of time. </p><p>Egyptian medical records, possibly dating from 1600 B.c., contain some advice on the treatment of venom poisoning, and some des- criptions of venomous animals. Perhaps one of the earliest references to toxic marine organisms is found in Exodus 7 : 20-21 : " . . . and all </p><p>FIG. 1. The stingray, Dasyatispmtinaca (L.), and the greater weever, Trachinus draco L. From Grevin (1571). </p><p>the waters that were in the river turned to blood. And the fish that was in the river died; and the river stank, and the Egyptians could not drink of the water of the river . . . " (Moses, c. 1491 B.u.). This description is thought to refer to a " bloom " of toxic dinoflagellates. </p><p>But the most exhaustive and credulous early writer of natural and </p></li><li><p>MARINE TOXINS AND VENOMOUS AND POISONOUS MARINE m s 259 </p><p>medical history fact and fiction was Pliny, whose voluminous work Historia Naturalis contains numerous fascinating accounts of venomous and poisonous animals. Describing the stingray, Pliny writes : </p><p> So venimous it is, that if it be struchen into the root of a tree, it killeth it : it is able to pierce a good cuirace or jacke of buffe, or such like, as if it were an arrow shot or a dart launched: but besides the force and power that it hath that way answerable to iron and steele, the wound that it maketh, it is therewith poisoned. So frequently quoted were Plinys works that Hulme (1895) com- </p><p> Several writers of antiquity influenced the mediaeval authors, but it is scarcely necessary to detail their labours at any length, since if they lived before Pliny he borrowed from them, and if they lived afterward they borrowed from him, so that we practically in Pliny get the pith and cream of all. As Klauber (1956) has so aptly concluded: Plinys Historia </p><p>Naturalis was the funnel through which we can watch the ancient folklore pouring down into the mediaeval and modern worlds. </p><p>Between the days of Pliny and the present a number of works on venomous and poisonous animals, including the marine forms, and their toxins have been published. These include the fine contributions by Grevin (1571), Autenrieth (1833), Bottard (1889), Phisalix (1922), Pawlowsky (1927), Evans (1943), Phillips and Brady (1953), Buckley and Porges (1956), Kaiser and Michl(1958), Nigrelli (1960), Keegan and Macfarlane (1963), Halstead and Russell (1964). By far the most comprehensive work on toxic marine animals is that being prepared by B. W. Halstead (1965-66). The author recently had the pleasure of browsing through Dr. Halsteads manuscript. The two volumes are the most complete and fascinating work yet prepared on this subject. They will greatly enhance our knowledge of the venomous and poisonous marine organisms. </p><p>mented : </p><p>C. General chemistry and zootoxicology of rnarine poisons Marine toxins vary considerably in their chemical and zootoxi- </p><p>cological properties. Some are proteins of low molecular weight, while others are proteins of obviously high molecular weight. Some of the toxic fractions appear to be amines or quaternary ammonium com- pounds or polypeptides or mucopolysaccharides, while the structure of still others is unknown. Some marine toxins contain enzymes but these substances are not nearly as common as they are in the reptilian venoms, nor are they to be found in such large quantities within a </p><p>x 2 </p></li><li><p>260 FINDLAY E. RUSSELL </p><p>single toxin. Another characteristic of many marine venoms is their relative instability. Some are very labile, even at temperatures down to 0C. While the marine toxins as a whole are far more varied in their chemical composition than the venoms of terrestrial snakes or arthro- pods, there is some degree of consistency (or lack of consistency) within a particular phylum, which is not unlike that seen in the terrestrial venomous animals. The more simple marine forms have poisons composed of one or several components having deleterious biological effects ; the higher forms have poisons containing more components, and in general these fractions appear to be more complex in structure and function. </p><p>The zootoxicological properties of marine toxins vary as remarkably as do their chemical properties. Some marine venoms provoke rather simple effects, such as transient vasoconstriction or vasodilatation, while others provoke more complex responses, such as parasympathetic dysfunction or multiple concomitant changes in the blood-vascular dynamics. The effects of the separate and combined activities of the fractions of these poisons, and of the metabolites formed by their interactions, is further complicated by the response of the envenomated organism. The organism may produce and release several autopharma- cologic substances which may not only complicate the poisoning but which may in themselves produce more serious consequences than the venom. </p><p>The zootoxicological study of marine poisons is further complicated by the fact that qualitative as well as quantitative differences in the chemical composition of these toxins may exist, not only from species to species within the same genus, but from individual to individual within the same species. A venom may even vary within the individual animal at different times of the year or under different environmental conditions. Thus, discrepancies in the proposed mode of action of a toxin are likely to occur until our knowledge of the individual fractions of these complex substances is more complete. In addition, obvious difficulties in determining the chemical and zootoxicological properties have arisen because of the differences in the methods of extracting venom, the methods of storage (fresh, lyophilized or crystallized), and the problems inherent in the methods of bioassay. </p><p>Most of our information on the zootoxicological properties of marine toxins is based on studies with mammds, which, of course, somewhat limits its application as far as understanding the design of the toxin in the animals armament. The venom of the black widow spider, for instance, did not evolve and adapt to the problems existing between that spider and mammals. Thus, it is not surprising to fhd </p></li><li><p>W I N E TOXINS AND VENOMOUS AND POISONOUS MARINE ANIMALS 261 </p><p>that its venom is twenty times less lethal to some arthropods than it is to the mouse, while on the other hand is is also ten times more lethal to certain other arthropods which have not adapted in the same manner. Some sharks appear to be relatively immune to stingray venom while others from completely different habitats are very sensitive to this toxin. Some non-venomous reptiles are not only immune to the venom of certain of the snakes of their area but actually feed upon these snakes. These various studies indicate that care must be exercised in applying data derived from studies in one group of animals to conclusions about the biological effects of a toxin in another group of animals, or to data on the design, use and adaptation of a toxin. </p><p>No comprehensive classification for toxins now exists. Our know- ledge of the chemical and zootoxicological properties of these complex substances is not broad enough or consistent enough, at the present time, to permit the adoption of a single working classification. It would seem, in the absence of a useful and reliable method for classifying toxins that it might be wisest to develop a system based on the tax- onomy of the animal. While such a classification would be somewhat bulky, and would need to be altered periodically with changes in taxonomy, it might serve our purpo...</p></li></ul>


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