On Marine Fish Diseases

CHAPTER 15

On Marine Fish Diseases CARL H. OPPENHEIMER Institute of Marine Science, Port Aransas, Texas1

I. II. III.

IV. V. VI.

VII.

VIII. IX. X.

I.

Introduction Marine Environment Methods of Study Bacterial Diseases A. Vibrio and Pseudomonas B. Furunculosis C. Tau Rot D. Tuberculosis E. Eye Infections F. Kidney Disease G. Miscellaneous Bacteria Virus Diseases Fungi Protozoans A. Mastigophora B. Sporozoa C. Ciliata Larger Parasites A. Trematodes B. Cestodes C. Nematodes D. Copepods Atypical Cell Growth Therapy of Marine Fish Diseases General Relationships A. Disease versus Catastrophes B. Effect of Disease on Fish Abundance C. Man as Host to Fish Diseases D. Disease in the Natural Environment E. Detection of Diseased Fish F. Fish Slime G. Control References

541 543 544 545 545 547 547 548 548 549 549 550 550 551 552 553 555 556 556 557 558 559 560 561 562 562 562 563 563 564 566 566 566

Introduction

The diseases of marine fish may be classified as: (1) dietary; (2) produced by organisms; or (3) from atypical cell growths. This chapter 1 Present Florida.

address:

The

Marine

Laboratory, 541

University

of

Miami,

Miami,

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consists mainly of a review of those diseases resulting from the activities of organisms. Microorganisms responsible for marine fish diseases may be bacteria, fungi, viruses, or protozoans. Larger parasites such as the trematodes, nematodes, cestodes, or Crustacea are commonly found associated with fish, but little is known of their pathogenicity. However, a few of the trematodes have been reported to cause death of fish; e.g., Benedenia melleni, a monogenetic trematode, lives in the skin and eyes of fish, and Cryptocotyle lingua cercariae larvae localize and encyst in the skin, causing black spot disease. The larger external parasites, such as the copepod, the brachiurian Argulus or so-called "fish louse," have one characteristic in common, that is, they irritate or break the integument or epithelial layers, providing entrance for the smaller unicellular pathogens, and they may also be quite destructive. While this latter situation may be common in nature, it is quite difficult to document. It is possible that the larger parasites may shorten the life span of fishes. There is some difficulty in distinguishing between marine and freshwater diseases in fish like salmon or eels which have their habitat in both spheres. This is also true of fish found "in brackish waters. The study of fish diseases as a whole has been a seriously neglected field. Very little effort has gone into the study of marine fish diseases. Most investigations have been undertaken sporadically, and Ross Nigrelli at the New York Aquarium and Reichenbach-Klinke from the Technical Institute at Braunschweig, Germany, have regularly made important contributions. Several books containing references to marine fish have been written on fish diseases: "Handbuch der Fischkrankheiten" by Hof er (1904), "Praktikum der Fischkrankheiten" by Plehn (1924), "Fischkrankheiten" by Schäperclaus (1941, 1954), "Culture and Diseases of Game Fishes" by Davis (1956), but these books have been devoted primarily to fresh-water fishes, or to exotic aquarium fishes, as in the book of this title by Innes (1952). Most of the bacteria listed in this chapter are described in the sixth or seventh edition of "Bergey's Manual" (Breed et ah, 1948, 1957). The protozoans are described in "Protozoology" by Kudo (1954). The author makes no claim as to completeness of known or reported marine fish diseases. The proceedings of a special conference on fish diseases held in Leningrad, March, 1957, covers both marine and fresh-water fishes and surveys in a comprehensive way Soviet investigations in this field (Pavlovskij, 1959). A special review is devoted to Soviet studies of the parasitology of marine fishes (Poljanskij and Byochovskij, 1959).

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MARINE ENVIRONMENT

Because fish are found in both fresh water and salt water, and some species, such as the salmon, inhabit both environments at different stages of their life, it is important to describe the biologically important aspects of the environment. The following points must be considered in any discussion of fish diseases because the activities of microorganisms are directly influenced by the presence of salts, temperature, and pH. Basically, the marine environment is distinguished from fresh-water environments by the presence of an average of 3.5% of salts in the former. It is quite difficult for the biologist interested in estuarine environments to draw a line between fresh-water and sea-water environments. The proportion of salts in marine brackish water remains relatively constant, being influenced primarily by the types and quantities of salts in the diluting fresh water. Marine parasites may be killed in fresh water and vice versa. A knowledge of the effects of temperature on the activities of bacteria is important to the understanding of bacterial diseases of fish. Usually, bacterial activities are enhanced by increasing temperatures. Thus, many bacterial diseases are more prevalent during warm summer months. The temperature in the marine environment will fluctuate during the day and season, and the degree of temperature change is proportional to latitude and depth of water. For example, in a temperate environment where the water is shallow or protected, near-shore areas may have a day temperature of 30°C. during the summer and be frozen during the winter. In the open sea the temperature is much more constant. During adverse temperature conditions in the natural environment, fish may move to deeper water where the conditions are more stable and where surface temperatures do not penetrate rapidly. Sea water is normally considered buffered at approximately pH 8.2 by the carbonate system, although the activities of photosynthesis and respiration may change the pH as much as from 6.5 to 8.5 during a 24-hr. period due to the uptake and production of carbon dioxide. Again, the degree of change is directly related to the activities and abundance of organisms and the mixing of the water. Fish are sensitive to changes in pH (McFarland and Norris, 1958), and death may be correlated with acidosis or alkalosis. It is a matter of opinion whether we use the term marine fish disease for the lesion disease of pike caused by Vibrio anguillarum in the Baltic Sea at a salinity of 0.6%. Some microorganisms such as Bacillus sal· monicida which cause fish furunculosis may survive up to three days in sea water, but the disease only occurs in fresh-water salmonid fishes,

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whereas an organism such as Ichthyosporidium hoferi can cause internal and skin lesions in both fresh-water and sea-water fishes, although in the latter case fuhire investigations might establish that the fresh-water and sea-water species of Ichthyosporidium are different (see further Chapter 16). II. Methods of Study

Methods for the detection or study of microbial diseases are similar to those which are used in the medical laboratory. In accordance with Koch's postulates, a disease-causing organism can be defined, first, through isolating in pure culture, second, by producing the same disease in a healthy fish, and third, when reisolated from a second diseased fish. Nutrient medium with either distilled water or 75% sea water and var­ ious sources of amino acids or extracts from fish tissues may be used to isolate the microorganisms. Usually each microorganism requires a special medium for cultivation (Liston, 1957). It is imperative that either very recently dead or, preferably, live fish be used for microbial examination for diseases. Normal fish have many bacteria living in the intestinal tract and on the surfaces of the integu­ ment which can overgrow the disease microorganisms within a short time after the death of the fish. The causative organism can be obtained with less time and effort if a live fish is examined. Usually the bacteria causing disease are difficult to cultivate and may require special growth factors. If an inspection of a fish reveals some commonly known disease or­ ganism 2>uch as Pseudomonas ichthyodermis, one may turn to the liter­ ature to find the proper medium, which for this particular organism is sea-water peptone. Rare disease organisms require experimentation in order to find the proper medium. Various staining techniques and mi­ croscopic examination may be used for the partial identification of bac­ teria. The acid-fast stain may be used to identify the Mycobacterium which produces tuberculosis. The gram stain may be used to study the gram-negative Diplobacillus which causes kidney disease. Dead fish and even canned fish can be inspected for atypical cell growths or organisms which cause internal lesions. Infected tissues are sectioned, stained, and examined for the identification of organisms such as Ichthyosporidium hoferi or Oodinium ocellatum which can be iden­ tified by morphology. Usually those stains which are positive for nuclear material are used. Virus diseases are difficult to study because of the minute structure of the causative organism. The presence of a virus disease is usually sug­ gested when a specific disease-causing organism is not detected. The presence of a virus is proven when material from the diseased area re-

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mains infective after being passed through a bacterial filter. Special pro­ cedures are necessary for the cultivation and examination of viruses in the laboratory. Protozoan diseases may be identified by their various forms during life cycles. Often the disease is recognized by the presence of a specific intermediate host such as the snail Littorina litorea, which is an inter­ mediate host for the trematode, Cryptocotyle linguae, causing black-spot disease in herring, or the presence of the cysts of the dinoflagellate Oodinium ocellatum which are found on the aquarium bottom. Char­ acteristics will be further explained in the section on protozoan diseases. III.

Bacterial Diseases

Although "red disease" in the common eel has been known since 1718, one of the first reports of a causative agent of a marine fish disease was that of Hof er (1904), who showed in 1877 that the "Lachspest" (ulcer) disease of salmon was caused by the organism Bacillus salmonis pestis, a short, thick, motile rod which grew in sea-water medium. Soon after this discovery, reports on the cause and diseases of fish in fresh­ water culture ponds and streams became more numerous. Bacteria are ubiquitous, microscopic, unicellular rods, spheres, or spirals which are found living in sea water and marine sediments. Water acts as a perfect vector for the bacteria that may exist either attached to larger particles or floating free in the water. Because they do not pass through an unfavorable atmospheric environment, as human pathogens usually do, it is easy to see how a fish pathogen can pass from one fish to another, except for the viruses, which are believed to pass directly from host to host. Many of the bacterial fish pathogens live as saprophytes in sea water and sediments. Most marine bacteria contain or liberate proteolytic enzymes which actively decompose the fish proteins after the death of the fish. During life, however, the mucus, scales, and skin are an effective barrier to the action of destructive bacteria. Bacteria may gain entrance if the layers of the integument are damaged or removed, by punctures by fins of other fishes, by the abrasive effect of the sea bottom or aquarium, or by the action of parasites living on the fish surfaces. A. Vibrio AND Pseudomonas The "red disease" of the common eel, caused by Vibrio anguillarum (Brunn and Heiberg, 1932), has been known for more than 200 years. At times 30% of the inspected eels were infected. The disease is systemic, and the bacteria are thought to gain entrance through the gills or in­ testine. The disease is first noted by the occurrence of a red coloration

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at the tail or fin base which later spreads over the entire body. Eels nor­ mally die before the red coloration spreads over the entire fish. Mi­ croscopic examination reveals that tissues of the fish are congested with blood and contain the Vibrio. This disease occurs during the warm months and is primarily reported from Danish waters and surrounding areas, the Baltic Sea, and the Mediterranean. During the summer, the eel fishery suffers considerably because the European housewife purchases living eels and will reject those appearing abnormal. Vibrio anguillarum is also infective to young plaice and mature codfish (Dannevig and Hansen, 1952) and may cause extensive mortality in the aquarium. Its taxonomical characteristics have been studied by Nybelin (1935). As a whole, it is considered halophilic and generally specific to eels. A recent epidemic (1959) was reported from the Baltic coast of Germany (Mattheis, 1960) after an interval of no less than 27 years with relatively low rates of incidence. High temperatures of surface waters are required to allow this agent to develop in great numbers. Warm weather con­ sequently is a prerequisite. A toxin is produced with multiple effects on the eel, generally resulting in death. There are two types of this Vibrio: A and B, with slightly different physiological characteristics. An eel epi­ demic is generally followed in the next year by a pike epidemic (Mat­ theis, 1960). The motile bacterium Pseudomonas ichthyodermis, which causes ex­ ternal lesions and death in many fish (Wells and ZoBell, 1934), has been found in both the Atlantic and Pacific Ocean, where it apparently is a part of the normal flora. For a limited time in the warm summer months the microorganism may cause severe damage to aquarium fish and then apparently loses its pathogenicity. Laboratory experiments by the author during one summer outbreak of the disease in the Scripps aquarium and in the natural environment revealed that the organism, isolated from fresh skin lesions, remained pathogenic for approximately two weeks. It lost its pathogenicity at about the same time that the lesions in aquarium fishes began to heal. During this time, free divers noted the rise and decrease in the incidence of fish with lesions in the vicinity of the aquar­ ium. The latter example of the loss and gain of pathogenicity of a marine organism may be only one example of a common occurrence. Hemorrhagic septicemia appears to be a common disease in both aquaria and natural habitat. It is easily noted by the presence of lethargic fish with external lesions or by surface bleeding. At times, freshly netted fish such as sardines will demonstrate external bleeding caused by me­ chanical abrasion from the net. This surface bleeding disappears after a short time in the aquarium. The disease, caused by bacterial invasion of tissues, is characterized by the bleeding and edema of internal tissues,

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although it is possible that an external bleeding of the skin resulting from the advanced systemic infections by Pseudomonas, Vibrio, and other undescribed organisms also may be classified under this disease. Several Vibrio and Pseudomonas species shown to be pathogens may form lesions, produce a general infection of other tissues, and finally cause a general septicemia just prior to death (Hodgkiss and Shewan, 1950). Several outbreaks of hemorrhagic septicemia accompanied by heavy mortality for salmon, herring, and sardines have been reported in the vicinity of the Washington State Fisheries Station (Rucker et al., 1953). Vibrio species were found to be responsible for these diseases. Other Vibrio may be the cause of widespread diseases (Rucker, 1959). Mass mortalities have been reported in the herring and pilchard population in Alaska and Vancouver Island waters, and in herring held in live bait tanks (Rucker et al, 1953). The disease is similar to furunculosis and is spread by contaminated water or from fish to fish by predation. About 10% of the codfish caught off the south coast of Sweden between Ystad and Trälleborg had a Vibrio infection of the eyes (Rucker, 1959). Aquaria-held plaice had Vibrio infections involving 49% of the fish (Rucker, 1959). B.

FURUNCULOSIS

McGraw (1952) indicated that furunculosis, a common disease of fresh-water fish caused by Aeromonas salmonicida, is not found in the marine environment. The disease has been reported to be present in adult Pacific salmon (Wood, 1959). However, it is problematical whether the disease was caught in fresh water and then survived the exposure to salt water. The bacteria were isolated from lesions found in the kidney and soft, blister-like areas filled with blood just under the skin. It is quite possible that the disease was acquired after the fish entered the fresh water. This systemic disease has been extensively studied because of its prominence in hatcheries—see further Chapter 17. C.

TAIL ROT

Tail rot commonly occurs in fish kept in marine aquaria. It is also reported from natural habitats. Only two specific examples are given here. However, it is the author's opinion that this is a very common disease of marine fishes. Herring in tanks and sea water at Boothbay Harbor, Maine, repeatedly have had tail rot (Sinderman and Rosenfield, 1954). The first sign of the disease is a hemorrhagic area under the scales near the peduncle and a progressive growth of a lesion accompanied by the sloughing of scales. Most often this occurs at the base of the tail, although it may appear at other places on the body. The early stages of tail rot can easily be seen in the water but not as easily as when the fish

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are in air. As the disease progresses, there is a general sloughing-off of the tail region, and death occurs soon after. A gram-negative rod was isolated from the lesions and other external parts of the fish. Death is believed to be caused by a systemic reaction to the disease organism. Tail rot is also quite prevalent in cod holding cages in Norway (Oppenheimer, 1958). The symptoms are identical to those described above, and the causative agent was found to be an organism similar to Pseudomonas. The disease causes some concern to fish handlers since the aver­ age Norwegian prefers to purchase living codfish for his table, and the diseased-fish are not eye-appealing, even though the causative organism is not pathogenic to humans. D.

TUBERCULOSIS

Earlier incidences of tuberculosis in fishes were reviewed by Griffith (1930) and Aronson (1938). The first described case refers to carp (Ba­ taillon et al., 1897), and the first report of this disease from marine fish (cod) was provided by Johnstone (1913). A comprehensive recent review of this field of research has been presented by Parisot (1958). The disease is characterized by internal lesions or dull white tu­ bercles in the kidney, liver, spleen, and intestine. Occasional external lesions have been found. The acid-fast bacilli Mycobacterium marinum may be found in the lesions. The infected fish may appear normal and are not suspected of being diseased until tissue smears of bacteria are shown to be positive. Death is probably indirect, resulting from weak­ ening of the fish, which easily become prey. The disease has been re­ ported for the common eel, cod, sergeant major, croakers, black sea bass (Parisot, 1958), chinook salmon (Earp et al., 1953), and several other species of marine fish. The organisms reported have not been pathogenic to man. The disease is probably transmitted by an infected fish when eaten by another fish. In a study of salmonid fishes returning from the ocean for spawning in 1952-1956, a high incidence of tuberculosis was established. As the disease was confined to fish known to be of hatchery origin—in some cases 100%—the feeding of untreated carcasses and viscera from origi­ nally tuberculous fish was considered a prime factor (Wood and Ordal, 1958). Such a direct transfer of this disease from feed to young fish was reported by Baker and Hagen (1942). E.

EYE INFECTIONS

Eye infections in marine fish in aquaria and in the natural habitat are common. Very few reports have been published on the cause of the eye disease except for fresh-water fishes; the causative agent is thought to be

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Ichthyophthirius multifiliis, Dermocystidium sp., Ichthyosporidium hoferi, and several bacteria, including P. punctata (Reichenbach-Klinke, 1954b). Fish death statistics from the New York Aquarium indicate that 0.32% of the fish died of exophthalmos or protrusion of the eyeball (Nigrelli, 1940). The disease in sea-water fish is characterized by a protrusion of the eyeball and subsequent loss of the eye or the growth of an opaque covering over the eye. F.

KIDNEY DISEASE

Kidney disease is common in salmon during hatchery procedures (Earp et ah, 1953). It is characterized by lesions of the kidney, liver, and spleen in which a small, gram-negative Diplobacillus is always found in large numbers. The disease is usually confined to fresh-water aquaria, although it will continue when the fish are placed in sea water. It may also be pertinent that the Diplobacillus will grow in sea-water medium. The Diplobacillus is always present, but investigators are not certain that the microorganism is the primary disease agent. G.

MISCELLANEOUS BACTERIA

Several other bacteria have been reported to cause fish death but have been little studied. Treponema cotti, T. pavonis, T. triglae, and T. perexile have been found in the intestine and blood of the marine bullhead, blenny, and other marine fishes (Breed et al.9 1948). Spirochaeta aeglefini and S. gadi have been found in the blood of haddock and cod. The descriptions of these organisms in the sixth edition of Bergey's Manual were not sufficient to be included in the seventh edition. It is problematical whether the microorganism caused death of the fish, but it is interesting to note that such microorganisms have been found. Recent studies have shown that Flavobacterium piscicida isolated during a red tide outbreak along the Florida Coast produces a compound, possibly a neuromuscular toxin, which is toxic to marine fish (Meyers et al, 1959). It is not unreasonable to assume that the byproducts of some bacteria may be toxic to marine fish in unusually large plankton blooms where the toxic material is concentrated. However, large growths of bacteria are only found in the sea where planktonic blooms are present. In occurrences of red tide where organisms such as Gonyaulax or Gymnodinium have produced materials toxic to fishes (Galtsoff, 1954), it is quite difficult to distinguish other toxins produced by bacteria. By-products from toxic bacteria may also cause death of fish as reported in a preceding paragraph. Inclusion bodies resembling Rickettsia have been found in the leucocytes or distributed in the plasma of the blood and kidney of Tetrodon

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fahaka (Mohamed, 1939). The fish had died for some unknown reason. On either side of the head and body there appeared several necrotic ulcers which resembled an ulcerative dermatitis. Cultures made from the liver and blood were negative for bacteria. No conclusive evidence for a Rickettsial disease was found other than the presence of the inclusion bodies, which were not normal for other similar fish. Chondrococcus columnaris is causing concern to the canners on the Columbia River. This bacterial disease has become prevalent among the blueback salmon migrating up this river for spawning. It is known that this disease is contracted in warm waters (Anonymous, 1960). IV. Virus Diseases

In general, there have been very few reports of viruses in the marine environment (Watson, 1953). The most noteworthy disease caused by a virus is Lymphocystis found in flounder, and many other marine fish (Nigrelli, 1940; Weissenberg, 1949). The disease is widespread, found in both fresh-water and marine fishes, and is characterized by the appear­ ance of nodules on the body or fins, which in severe situations may cover most of the fish. The virus, which can be seen as inclusion bodies in the nodules, stimulates growth of fibroblast cells in the connective tissue which enlarge sometimes up to 2000 microns during a year. The en­ larged cells surrounded by a thick transparent membrane are called lymphocystis cells. Desiccated pieces of the nodule placed in the tissue of other fish will experimentally cause infection. Bruised fishes will be infected if placed in water containing an emulsion of lymphocystis mate­ rial. The virus organisms pass through a Chamberlain filter. High mor­ talities have not been common, and it is assumed that most detrimental aspects of the disease arise from predation of the diseased fish. V.

Fungi

The most common fungus disease of marine fish so far known is caused by systemic infection of Ichthyosporidium hoferi, reported in 1893 for fresh-water fish and later found in such marine fishes as flounder, haddock, mackerel, and herring (Sindermann and Scattergood, 1954; Gustafson and Rucker, 1956). Its occurrence in the Mediterranean has been subject to a special investigation by Reichenbach-Klinke (1957). He also discussed trends as to its distribution in this important basin. Two epidemics (in herring) have been reported (Sihdermann and Rosenfield, 1954). The greatest infections occur near shore in late winter and early spring. The disease produces marked changes in the tissue as well as changes in behavior; e.g., erratic swimming motion. A sandpaper ef­ fect—roughening of the skin—usually in the lateral regions, is the first

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external evidence of disease. The effect is due to the presence of small, raised papules caused by the rapid growth of the fungus under the epidermis. When the papules perforate the surface of the epidermis, small black areas are revealed. The small black areas should not be confused with the similar effect caused by trematode metacercariae which encyst under the epidermis (Sindermann and Rosenfield, 1954). In later stages the epidermis sloughs off as necrosis becomes more extensive and lesions are formed. In addition to the external effects, gross infections occur in muscle and internal organs. The internal lesions have the appearance of white nodules as large as a centimeter in diameter. The heart is usually the first organ to be affected, and the small nodules may grow until the heart is enlarged several times and is completely covered by a white fungus mat. The external and muscular infections are more common in the herring and often absent in the mackerel and flounder. At times the myxosporidian Chloromyxum clupeidae will produce pus pockets in the muscle of herring which are similar to fungus nodules. The fungus may be found during the microscopic examination of stained sections of the fish. Both round, sporelike bodies (10 to 250 microns) and fungal elements may be demonstrated. Single spores are indicative of an early infective stage, and the presence of filaments indicates advanced infection. The infection is spread through the fish by the distribution of the spores through the blood. Evidence of infection was noted within 30 days in fish inoculated with the fungus. Spores attached to copepods eaten by the fish were also a source of infection. Infective spores were liberated to the water from skin lesions or after decay of infected fishes. Reichenbach-Klinke (1954a) reported the presence of a fungus in gill tumors of codfish. It was not possible to determine whether the fungus was the primary or secondary cause of the tumor. Saprolegnia is a common fungus parasite of fresh-water fish which grows as a white, cottony mat on the surface of the fish and infiltrates into the integument and muscle. The parasite may be found in brackishwater fishes (Vishniac and Nigrelli, 1957). Earp et al. (1953) found that Saprolegnia disappeared when infected chinook salmon were placed in sea water. VI.

Protozoans

The author is not cognizant of any protozoan diseases of fish which have been reported to have man as a host. Most of the protozoan parasites are host-specific, although some are not; Eimeria clupearum lives in the liver of herring, mackerel, and sprat, and Ichthyophthirius multifiliis

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and Oodinium ocellatum are parasitic to many species of marine fish. Eimeria may be found in the feces of human beings who have eaten in­ fected fish. However, the spores of the Eimeria, common to commercial marine fishes, may pass through the digestive tract of man and appear in the feces (Kudo, 1954). It is quite possible that protozoans such as the myxosporidian Henneguya may incite the formation of tumor growths such as have been reported for the papillary cystic disease of catfish. The parasite causes the formation of fibroblasts which replace tissue. Protozoans are currently classified into Mastigophora, Sporozoa, Sarcodina, Ciliata, and Suctoria. Most families of the protozoans have been found indigenous to the marine environment, and it is therefore not un­ reasonable to anticipate that most forms parasitic on fresh-water fish will have marine counterparts. The Sporozoa are, without exception, para­ sites. Protozoa representing all classes except Suctoria have been re­ ported to be parasitic to marine fish. Although many of the parasites do not appear to cause death directly, they may act as loci for infections by bacteria or fungi. Epidemics have been caused by the sporozoans (Myxosporidia and Microsporidia) endoparasites of several marine fishes. Most of the protozoan parasites are hyperinfective in marine aquaria where they are in constant contact with the fish during the life cycle of the microorganisms. In the environment, reinfection of the host by its para­ sites may be rare because the fish will move away during the various metamorphic changes of most of the protozoans. Perhaps for this reason most of the parasites do not cause extensive damage in nature unless crowded conditions exist, as in schooling behavior. The endoparasitic forms are often overlooked because microscopic observations are needed to detect the causative microorganisms. The exoparasites appear as growths, open sores, or pustules which can usually be seen without mi­ croscopic examination. A.

MASTIGOPHORA

Three species of the class Mastigophora are known to parasitize marine fish. Amoeba, although found in fresh-water salmonid fishes, has not been found in marine fish. The dinoflagellate Oodinium ocellatum is commonly found in marine fish in tropical waters, during temperate summer temperatures, and in aquaria (Nigrelli, 1936, 1940; Kudo, 1954; Dempster, 1955; Reichenbach-Klinke, 1956). Oodinium is a free-swim­ ming dinoflagellate which attaches to the gills of the host fish by cytoplasmic processes. Occasionally the organisms attach to other parts of the integument. The parasite can cause extensive damage in closed aquaria because the mature organism falls off the gills to the sea bottom

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or aquarium floor where it goes through its life cycle and is infective. The organism enlarges, and the cyst contents divide into 128 flagellated cells, which further divide into 256 free-swimming flagellates capable of reinfecting the host. The infection impairs the normal gill function and when severe may cause death. Trypanosomes have been found in the peripheral blood and Protrichomonas in the esophagus of marine fish, but little is known about the toxicity of these organisms to the host. Trypanosomes have been reported for many years from marine fish of European waters (Lebailly, 1904; Neumann, 1909), and a number have been described from similar hosts in Africa (Fantham, 1919; 1930), Australia (Mackerras and Mackerras, 1925) and New Zealand (Laird, 1951). Only a few, however, have ever been encountered in marine fishes of the United States (Saunders, 1959). Such trypanosomes were found in a winter skate, Raja ocellata Mitchill, from Woods Hole, Massachusetts, and were identified as T. rajae (Laveran and Mesnil, 1902). Bullock (1958) also reported it from the common skate Raja erinacea Mitchill, from southern New England. Trypanosoma rajae and T. giganteum are found in the ray Raja oxyrhynchus. Trypanosoma granulosum has been reported in the eel (Kudo, 1954). B.

SPOROZOA

Sporozoan protozoans may cause extensive damage to natural fish populations. Walford (1958) lists some of the Myxosporidia and Microsporidia which have been reported to be parasitic to marine fishes. These organisms are endoparasites which invade the tissues of the host fish. Because the life cycle varies, the free-existing mature cysts in general are taken into the gut of the host with food or in connection with other activities. The cysts germinate into sporozoites which penetrate the epithelial cell or other host tissues, depending on species. Each organism then matures through several stages and eventually ruptures the host cell to liberate oocysts or spores. The spores are quite resistant and may exist in rather unfavorable environments and still be viable. The spores of the sporozoans are quite typical, having the appearance of symmetrical asterisks or petal arrangements of 4 to 6 segments separated by a suture. The coccidian Eimeria has been found in the testes of sardines, the liver of herring, mackerel, and sprat, and the swim bladder of cod and the testes of menhaden. The presence of haemogregarines in the blood of fishes was first reported by Laveran and Mesnil (1901) from France. Since that time, other occurrences have been reported spasmodically from a few other countries. There is an uncertain record of its possible observance in east-

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ern Canada (Fantham et al., 1942). Laird (1951), who made the first systematic study of this group, found a considerable number in the fishes of New Zealand and the Fiji Islands. Kudo (1954) states that it occa­ sionally occurs in the blood of soles. The first recorded finding from the United States dates from 1955 and only in low frequency (Saunders, 1955). Some sporozoans are capable of excreting a powerful proteolytic enzyme which digests muscle. Soft, mushy white areas in muscle, caused by Chloromyxum and Hexacapsula, have been reported in hake, swordfish, Japanese sea bass, and yellowfin tuna (Fletcher et al., 1951; Matsumoto, 1954). In one instance in North Africa, 40% of the hake were infected. The jellied condition of swordfish was originally attributed to Chloromyxum sp. (M'Conigle and Leim, 1937). Subsequently, Arai and Matsumoto (1953) discovered that in yellowfin tuna the active agent was Hexacapsula neothunni. Such jelly-like meat has been observed in a great many food fishes besides the two mentioned, such as halibut, turbot, and other tuna-like fishes. The causative microorganism has in general not been identified, frequently under the supposition that this radical change in texture was due to some unexplained biochemical activity, presumably of an enzymatic nature and constituting part of the autolytic breakdown of the tissue (Tsuchiya and Tatsukawa, 1954; Tsuchiya and Kudo, 1957). The myxosporidian Chloromyxum thyrsites has been reported from the following South African fishes: snoek (barracuda), hake, and John Dory. This organism also is found in the pilchard and the maasbanker (horse mackerel) (Rowan, 1956). Pilchard tissue with more than two million spores per gram collapses and turns mushy when canned or otherwise processed (Rowan, 1956). This tissue breakdown through these sporozoans is generally attributed to proteolytic enzymes acting extremely rapidly. Whole fish can be decomposed within 24 to 70 hr. at normal temperature. Chloromyxum clupeidae causes ulcer disease of herring (Sindermann and Rosenfield, 1954). Only circular exterior ulcers are present, and internal lesions have not been noted. The common haddock parasite—a myxosporidian, Myxobolus aeglefnv—occasionally is encountered in the cranial tissue of such other fish as the plaice, thus emphasizing the opinion upheld by several fish pathologists that sporozoan parasites tend to be tissue- rather than host-specific (Kabata, 1957). Certain boreo-arctic microsporidians have been studied by Kabata (1959) and reported from such far-separated regions as the White SeaIceland, and the Gulf of St. Lawrence.

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The microsporidia are widely distributed as parasites of fish. They invade and undergo asexual division and sporogony within the host cell. The host's invaded cells become hypertrophied. The organisms are small spores, 3 to 6 microns in diameter. The spores are easily identified by the presence of an outline or ridge which separates the spore into two halves. The spores, taken into the intestinal tract, pass through the epi­ thelium, enter the blood stream and reach the specific site of infection. Glugea has been reported in the mucosa of the gut of smelt, stickleback, and plaice (Kudo, 1954 and Bückmann, 1952). Certain myxosporidian sporozoans appear to be host-specific for the tissues of specific marine fish. Thus, the tapioca disease of salmon is caused by Henneguya salminicoL· (Kudo, 1954). Severe epidemics of wormy Pacific halibut have been reported to be caused by muscle inva­ sion of Unicapsula muscularis (Kudo, 1954). At times, 5% of the catches of barracuda contain infections of Kudoa thyrsites, which causes the muscle fibers to turn milky and soft (Kudo, 1954). Other species of Kudoa have also been found in body muscles of marine fish. Sphaeromyxa have been found in the gall bladder of Motetta and other marine fish of Europe, and Siphostoma appear in United States fish. To show the complexity of nature, it has been reported that the urinary bladder of the toadfish Opsanus tau is parasitized by the myxosporidian Sphaerosphora polymorpha. The trophozoite of the parasite is then parasitized by the microsporidian Nosema notabilis. The microsporidian parasite does not directly infect the tissues of the host fish. Lentospora cerebralis (now known as Myxosoma, according to Kudo, 1954) has been found to cause twist disease in herring reared in aquaria (Dannevig and Hansen, 1952). The gall bladders of sharks, rays, Gadidae, and Cyprinidae in the Mediterranean have been parasitized by the myxosporidians Leptotheca, Sphaerospora, Ceratomyxa, and Chloromyxum (Theodorides, 1955). Sinuolinea has been located in the urinary bladder of marine fish. Six species of the fish-inhabiting Cnidosporidia invade the cranium of several fish species, which not infrequently is found filled with a mass of soft substance constituted of spores. Such sporozoans are reported from cod, haddock, plaice, hake, etc.—for further reference see Kabata (1957). It is possible that several of these species are tissue- rather than host-specific. C.

ClLIATA

The round ciliate parasites Ichthyophthirius are a common cause of death in aquarium fishes. Marine fishes are parasitized by euryhaline species. The free-swimming ciliate enters the integument, where it forms

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a cyst, feeds on the host cells, and forms white or grey pustules. When the cyst develops into a mature trophozoite, it drops from the fish to the aquarium bottom, where it encysts and divides into young ciliates, which break through the cyst wall and may reinfect the host. The barrel-shaped ciliate Trichodina has been reported to infect puflEers of Sand Hook Bay and is assumed to be a source of infection for other fish (Nigrelli, 1940). VII. Larger Parasites

It is problematical whether extensive mortality in the natural environ­ ment occurs from the infestations of larger parasites. In crowded hatchery conditions, the infestations of the larger parasites may often cause death. Reports on the occurrence of helminth and crustacean parasites are so numerous that it will be pertinent to mention a few representative groups. In general, the larger parasites fall into three groups: flatworms (Platyhelminthes), roundworms (Nemathelminthes or Acanthocephalians), and copepods (Arthropoda). Most students of biology are familiar with theflukesand tapeworms. External-living flukes or trematodes, such as the monogenetic forms, may live in the gills, while digenetic forms live in the intestinal tract; some exceptions have been noted. Some flukes may be infective to man, for example, Heterophyes heterophyes, present in the common mullet (Belding, 1942). A. TREMATODES

The flukes are short, elongated to oval forms, flattened dorsoventrally, and are surrounded by a cuticle in place of epithelial cells (Hyman, 1951; Koratha, 1955). They are hermaphroditic, producing eggs surrounded by a capsule. Having only one host, the eggs develop di­ rectly into an adult form. Flukes move on the host by a looping manner and feed on slime epithelian cells and blood exuding from places dam­ aged by the parasites' hooks and suckers (Llewellyn, 1957). They attach to the host by means of suckers, but some species have hooks or spines. The monogenetic trematodes are primarily ectoparasitic, inhabiting the gills and occasionally the skin of fish. Benedenia melleni caused death in aquaria-held spiny-rayed fishes (Nigrelli, 1940). The digenetic flukes are generally endoparasites which live in the intestinal tract of the fish. They have a more complicated life cycle, including several metamorphic forms, and require up to four intermediate hosts. Occasionally the intermediate stages of the trematodes may be dis­ ease agents, such as the cercaria stage of the Cryptocotyle lingua (Sin­ dermann and Rosenfield, 1954). This organism invades and encysts just under the epidermis, causing the area to become black, and thus creates "pigment spot" disease. Infection occurs in inshore waters where the

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snail Littorina Utorea is found. The hosts, during the life cycle of the trematode, are the snail, the gull or tern, and the herring. Massive infec­ tions may cause death of the herring. There are numerous descriptions and records of the trematode para­ sites of marine fish from various parts of the world. The majority of these have been conveniently summarized and reviewed by Dawes (1946, 1947), Yamaguti (1953), Skrjabin (1947-1955), and Manter (1955). Phil­ ippine food fishes were investigated recently with regard to the trematode family Bucephalidae Poche 1907 (Velasquez, 1959). Nagaty (1957) found several trematodes on fish in the Red Sea. A comprehensive study of diseases and parasites of marine animals off Florida has resulted in the reporting of a great number of trematodes from marine fish (Sogandares-Bernal and Hutton, 1959). Schistosoma cercariae may be so com­ mon as to cause skin disturbances in man (Hutton, 1952). A number of digenetic trematodes with high degree of host-specificity have in latter years been described from the West African coast (Thomas, 1959) from an area very poorly investigated earlier. In general, this sup­ ports the contention of Manter (1955) that host-specificity is of a high order among digenetic trematodes of marine fishes, particularly among those living in warm waters. The widespread geographical distribution of Lasiotocus longicaecum and the remarkably close resemblance of the forms on both sides of the Atlantic with those in the Pacific indicate that this species is monotypic. This coincides well with the low degree of host-specificity and with the free-ranging habits of the host fishes (Thomas, 1959). An investigation of monogenetic trematodes in the Gulf of Mexico is underway. Several papers have been published—for further references see Hargis (1955). The monogenetic Kuhnia scombii is a common para­ site of the gills of the Atlantic mackerel. A recent study was devoted to its attachment mechanism (Llewellyn, 1957). B.

CESTODES

The tapeworms are endoparasites of many fish. They have no epider­ mis, mouth, or digestive tract and attach to the host by means of suckers and hooks in the anterior end. The body is divided into segments or proglottids which are hermaphroditic, producing encapsulated eggs. Some species cast off the gravid segments. They require intermediate hosts to complete their life cycle (Hyman, 1951). Food is obtained by direct absorption through the surfaces of the segments. The head, or scolex, may be able to absorb food through the sucking disc. The tapeworms, or cestodes, generally inhabit the intestinal tract of the fish. Thus, the scolex of several cestodes has been reported from

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CARL H. OPPENHEIMER

the intestine in some fifty different species covering a wide area along the coasts of Europe, the Atlantic coast of North America, and the Arctic Ocean (Rees, 1958). Bothriocephalus scorpii has been found in the At­ lantic herring (Dollfus, 1956) and halibut (Rees, 1958). Cestodes are quite common in marine fish of India (Kambata and Bal, 1952, 1954; Rao, 1954; Ganapati and Rao, 1955). Some forms are found in the muscle such as Poecilancistrium rohustum, the spaghetti worm, which reaches lengths of 17 cm., embedded in the muscle of many species of fish in the Gulf of Mexico (W. N. McFarland, personal communication). The presence of the worm is annoying to anglers, although not harmful to man. At times, 44% of a catch may%be infected by as many as 5 worms per fish, while occasional large specimens of the croakers, Cynoscion nebulosus and Pogonius chromis, have upwards of 100 worms. The tape­ worm Diplogonoporus grandis lives as an intermediate host in marine fish and has infected humans in Japan (Belding, 1942). The plerocercoid larvae, wormlike in appearance, are found in the flesh of many species of fish. An important focus for tapeworm infections in man is the semisaline Baltic, particularly the Finnish Bay, according to Soviet investiga­ tions. Diphyllobotrium is the dominating species. Pike, pope, and turbot are the most affected species (Engelbrecht, 1958). See further Chap­ ter 16, of this volume. Elasmobranchs appear to be more heavily infested than teleosts (Robinson, 1959). Some important recent studies from various geographi­ cal areas are the following: Canada, Wardle (1932); New Zealand, Robinson (1959); India, Rao (1957) and Subhapradha (1955). C.

NEMATODES

Nematodes or roundworms are commonly found in marine fishes (Kahl, 1939; Rees, 1946; Olsen, 1952; Chalupsky, 1955). The sexes are separate and they have a simple life cycle. The worms are cylindrical, the anterior end equipped with hooks or teeth. A digestive system is present, and food is acquired from tissues and intestinal contents. Food may be absorbed through the epidermis. In most of the Gulf of St. Lawrence and in some other marine areas of eastern Canada, the infection of the flesh of cod with larval nematodes —chiefly Porrocaecum decipiens (Krabbe)—constitutes a considerable economic problem (Heller, 1949; Templeman et al, 1957; Scott and Mar­ tin, 1957, 1959). Seals are the hosts for the final adult stage, releasing eggs which reinfect young fish. The infection level is related to the preva­ lence of seals, chiefly three species. The relative importance of each seal species, as to the incidence of larval nematodes in cod from different fishing grounds off the Canadian Atlantic mainland, was studied by Scott

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and Fisher (1958). In all areas studied, there is an increased incidence of infection with age. In latter age groups, 70% of the cod were infected. No direct relationship was established between the frequency of this nematode in cod flesh and the distribution of seals, other than a drop in offshore waters as compared to inshore coastal waters (Scott and Martin, 1959). A number of other fishes were likewise infected, particularly the smelt, the American plaice, and the sea raven. No case of herring infection was found. Two other nematodes were frequently encountered (Templeman et al, 1957). Porrocaecum larvae have been reported from the European side of the Atlantic by Kahl (1939), and in both Greenland and the Barents Sea by Grainger (1959)—in all cases occurring in the flesh. Besides cod, he found larvae in the flesh of ling, redfish, angler, and wolf-fish. The occurrence of nematode worms in cod has been reviewed by Dollfus (1953). Another species, Anacanthocheilus, was in addition encountered in cod fillets from Iceland, Bear Island, and the Barents Sea. Punt (1941) studied nematode parasites in many North Sea fish. The incidence of infection and the number of nematodes per fillet usually is related directly to the size of the cod (Scott and Martin, 1957). The cod may become infected throughout its entire life, but the highest rate of infection occurs in small-sized cod. Changes in feeding habits which accompany growth in the size of cod may explain the fact that young cod frequently appear to be more exposed to infection than large specimens. Seasonal variations in the incidence of nematodes in different fishing locations are attributed to seasonal migrations of the cod. More intensive catching in certain shallow inshore waters is considered to account for a higher incidence of nematodes in this period (Scott and Martin, 1957). The nematode Contracoecum aduncum is harassing the Baltic cod (for further references, see Schulman, 1959). The larvae infest the liver and greatly reduce its economic value. Regional differences prevail, too. Thus, the degree of infestation explains why the ratio of liver weight to total live weight is 4.5 times higher in Riga Bay as compared to the waters of Liban (Schulman, 1959). D.

COPEPODS

The brachiurian Argulus (often classified as a copepod) is a common parasite on the integument of many fish and elasmobranchs. At times, the incidence of infecting brachiurians may be great enough to produce open lesions on the fish. Usually the parasite irritates the epithelium

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CARL H. OPPENHEIMER

layers of the fish, and may cause the fish to swim in an erratic fashion attempting to brush the parasite off. Copepods are common inhabitants of the gills and surfaces of marine fishes. Walford (1958) gives the history of Lernaeocera branchialis, which begins its life as a free-swimming form and later assumes the role of a parasite in the gill covers of flounders. Leaving the flounder, as a free-swimming form again, it next takes a cod as its host. In the cod gills the organism loses its copepod appearance and develops into a mass of tissue which eats its way into the host tissues. In Cuxhaven, up to 20% of the codfish examined were infected and below normal in weight. The parasitic copepod Lepeophtheirus salmonis is widely distributed as a sea louse on salmonids along the North Atlantic coasts, of both Europe and America, as well as in the North Pacific. Very heavy infes­ tations have been discovered on the southeast coast of Nova Scotia (White, 1942), even to the extent of killing the fish (White, 1940). The skin in the occipital region was removed through these parasitic attacks, thus exposing the underlying red flesh. This species is marine and suc­ cumbs gradually when the fish enters fresh water for spawning. Species of the parasitic genus FenneUa parasitize widely different hosts such as cetacea, flying fish, and swordfish (Gnanamuthu, 1951). Several major fishing areas have been scanned with regard to the incidence of parasitic copepods: off Angola, by Nunes-Ruivo (1956); off Madras in India, by Gnanamuthu (1951), Subhapradha (1951). VIII. Atypical Cell Growth

Tumors and atypical cell growths frequently occur in marine fish (Nigrelli, 1953). Often aquarium fish will be found with tumors (fibromas) or growths on the head which are presumed due to traumatic injury from the aquarium walls. Certain cell hypertrophy may be caused by fungi, viruses such as lymphocystis, and microsporidians such as the encystment of Glugea hertwigi in smelt. Thyroid tumors have been re­ ported for marine fishes in aquaria and in the natural habitat (Nigrelli, 1952). The thyroid tumors were malignant, infiltrating and destroying gill tissue, cartilage, and bone of the hyoid-mandibular-branchial com­ plex. Epidermal fin tumors have been reported in the gobiid fishes (Tavolga, 1951). In 1945-1947, an extensive epidemic of tumor disease, frequently named the "cauliflower" disease, in the silver and golden eel was reported in the area between the West Baltic Sea and the Kattegat along the German, Polish, and Scandinavian coasts (Christiansen and Jensen, 1947). New epidemics were reported in 1956 (Engelbrecht, 1958). The growth, consisting of epithelial tumors of a papillomatous structure, were

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primarily found on the head. Occasionally they appear on the fins or other parts of the body. Attempts to transmit the disease to healthy fish failed. This disease is also reported in cod (Schäperclaus, 1953). Whether it is due to a virus or initiated by a chemical traumatic effect is still an open question. The disposal of large quantities of mustard gases has taken place in these areas. IX. Therapy of Marine Fish Diseases

The question of therapy of marine fish diseases raises special problems. It is quite impossible to control a naturally occurring disease due to environmental conditions. Thus, therapy must be confined to the aquarium or fish hatchery. Unfortunately, most of the reports for the control of fish diseases are confined to fresh-water fishes, and it is difficult to extrapolate such treatment to the marine environment. Materials injected for internal therapy will not be affected by sea water. However, the pH and presence of salts may affect water therapy. For example, aureomycin loses its antibiotic effect in sea water of pH 8.2. Therefore, the extrapolation of fresh-water disease therapy to the marine environment should be made with caution. Various types of bactericides, sulfonamides, antibiotics, dyes, and copper sulfate have been employed for the control of disease in freshand sea-water aquaria. In general, the bacterial diseases are best controlled with antibiotics or sulfonamides. The microorganisms causing tail rot and P. ichthyodermis can usually be controlled with a mixture of penicillin and streptomycin injected into the fish at the concentration recommended per unit weight. External lesions may be controlled with direct application of merthiolate or other bactericides. Furunculosis may be controlled with aureomycin or terramycin. Vibrio infections respond to sulfonamide therapy. Davis (1956), in his book "Culture and Diseases of Game Fishes," adequately covers the therapy of most fresh-water fish diseases. It is imperative to note that the first steps in the therapy of aquarium diseases is to properly prohibit the spread of the disease by controlling the water which may pass through diseased areas, to sterilize periodically the aquaria walls with chlorine solutions, and to keep residue foods from decaying in the tanks. Experience has also shown that fish lose their resistance to disease when subjected to abnormal temperatures, pH or adverse water conditions. Many fish diseases are more prevalent during the warm summer months, and precautions should be taken to maintain a lower temperature in the aquarium during the summer. An important factor in the cultivation of marine fish is to make certain that food is not contaminated with disease organisms. Often freshly

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CARL H. OPPENHEIMER

caught or frozen marine fishes are used as food for aquaria fishes. Inspec­ tions of the food fishes should be undertaken to ascertain whether in­ fections are present. The larger external parasites and some external sporozoans such as Eimeria may be partially controlled by placing the fish in fresh water. Solutions of copper sulfate from 0.1 to 0.5 p.p.m., have been used to control Oodinium. External lesions may respond to treatment with mercurochrome or other germicides applied directly. Acriflavine was found nontoxic to killifish and was effective against protozoan infections at 10-hr. exposures of 10 p.p.m. of the dye (Pickford, 1952). X. General Relationships A.

DISEASE VERSUS CATASTROPHES

It is rather difficult to discuss fish diseases without a definition of the term. In man, the term "disease" encompasses a multitude of complex reactions of the human body to environment from malnutrition to virus infections. Thus, one must define what fish disease is and how it affects a living population of fish. Mass mortality of fish in the marine environment is commonplace. If the mass mortality is caused by disease, it is called an epidemic, whereas other mortalities may be considered as catastrophes. The toxicity of Gymnodinium and Gonyaulax red tides to fish has long been known (Galtsoff, 1954). Although fish are sensitive to adverse salinities and temperatures and can often avoid such conditions, often mass catastro­ phes caused by abnormal environmental conditions are found along the coasts of the continents. Such mortalities are of interest to the paleon­ tologist, who finds the remains of large masses of fish fossils in the geo­ logical record. Catastrophes may be caused by volcanism, abrupt salinity change, freezing, sudden change of temperature, toxins from micro­ organisms, lack of oxygen, or production of noxious gases such as H2S, and stranding on beaches due either to severe storms or to pursuit by predators (Brongersma-Sanders, 1957). Even the effects of currents which carry fish eggs and larvae into adverse conditions may, in broad terms, be considered as a mass mortality. B.

EFFECT OF DISEASE ON FISH ABUNDANCE

One can only guess at the gross effect fish diseases have on the abun­ dance of fish in the sea. Within the writer's experience, pandemics or large-scale epidemics of fish have never been reported. Several reports have been made describing localized epidemics. It is unlikely that disease is a primary factor in maintaining an under-

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populated sea. The total population of the ocean may depend on the limiting concentration of some essential mineral such as phosphate, nitrate or vitamin B12 necessary to some part of the food chain. Thus, the death of a fish may not affect the long-range total population, because bacterial activities may release bound-up nutrients such as P 0 4 , N0 3 , and other component minerals, or produce vitamin B12 during the decomposition activities. The by-products from the fish are then used by algae for the primary production of organic matter in the sequence of organic carbon cycles. C.

MAN AS HOST TO FISH DISEASES

It is unique that most of the microorganisms pathogenic to fish are not pathogenic to man. Fish have been shown to harbor human pathogens, such as Clostridium botulinum type E., Clostridium tetanus, and Erysipelothrix, but the organisms are apparently not pathogenic to the host fish. Aronson (1926) reported the presence of tubercles in croakers, sea bass, and a sergeant major, and described the causative organism as Mycobacterium marinum, which was not pathogenic to rabbits or guinea pigs. Other organisms in sea water near sewage sources which might be pathogenic to man are Escherichia, which can be found in estuaries (Rittenberg et at, 1958), paracolon bacteria, Shigella, and Streptococcus. Normally, human pathogens do not survive more than 2 weeks in sea water. Lethal cases of C. botulinum type E toxin poisoning have reportedly been caused from the organisms in home-pickled herring and homecanned salmon (see Chapter 3, Volume II), Section IV on Botulinus poisonings for further reference). The origin of the microorganisms could not be established. Clostridium botulinum type E has been found in the intestinal contents of fish from the Sea of Azov. Type A strains of C. botulinum have been found in sea water. Therefore, it is possible that the lethal cases of botulinum may have originated when the fish were in the natural environment or soon after catching. Erysipelothrix will grow quite readily in sea-water nutrient medium. Although the microorganism does no apparent damage to fish, fish handlers commonly contract the disease through fish-spine punctures or through cuts in the skin (Sheard and Dicks, 1949; Svenkerud et ah, 1951). D.

DISEASE IN THE NATURAL ENVIRONMENT

Usually, the microbiologist or physiologist is introduced to marine fish diseases when a specific incidence of an infected fish is brought to the laboratory, from reports of an extensive fish kill nearby, or from a fish which has died in an aquarium. Very rarely are references made to

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fish diseases discovered during the routine examination of fish on board a fishing vessel. The aqualung free-diving apparatus has provided some opportunity to observe fish in their natural habitat (Oppenheimer, 1953). More prevalent cases of infection have been noted on occasions. For instance, during several summers (1951-1953) fish with skin and eye lesions were common in the aquarium at the Scripps Institute of Ocea­ nography. At the same time, Conrad Limbaugh during free diving oper­ ations reported the presence of similar types of infected fish in the waters outside the surf zone near Scripps and off the Coronado Islands, July, 1952 (personal communication). Limbaugh reported that from 5 to 10% of the fishes showed evidence of infection which appeared as white blotches on the sides or fins of the tail surfaces. Some of the fish were blinded in either one or both eyes by a coating of white material. Pike-perch, kelp bass, sheepshead, and blacksmith were identified. In September, 1955, Limbaugh further noted a mass mortality of the queenfish Seriphus politus under the pier of the Scripps Institute of Oceanography. From 20 to 40% of the fishes were infected with open lesions. The following day hundreds of dead queenfish were noted under the pier. Many of the fish with lesions were inspected for the presence of patho­ genic bacteria. Several pure cultures of marine bacteria, including P. ichthyodermis, were isolated which produced the characteristic lesions in inoculated fish. Soviet investigations maintain that the composition of the parasitic fauna frequently distinguishes one race of a fish species from another. This is said to apply to local strains in the White Sea and to cod, herring, and "navaga" (Poljanskij and Byochovskij, 1959). On the other hand, studies of the Baltic show that in the Finnish Bay and other less saline eastern parts are encountered a number of fresh­ water species of coccidia, nematodes, trematodes and copepodes, while in the western and southern more saline sections of the Baltic several typical Atlantic species of these same groups are found (Shulman, 1959). Engelbrecht (1958) investigated fish parasites in two areas ofi the Ger­ man Baltic coast and concluded that fish in such brackish water are less subject to attack than either fresh- or salt-water fish. Two parasites are specific to brackish water and both the other two groups have difficulties in adapting to a brackish environment. E.

DETECTION OF DISEASED FISH

Often the lesions of fish are not noticed because of the physical prop­ erties of the refractive index of the water. In the water the lesions ap­ pear as cottony growths with perhaps an opaque membrane over the

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surface. When the fish is removed from the water, the lesion loses its characteristic appearance, and unless it is an open bleeding sore, the lesion may not be noticed. In the natural habitat it is difficult to get near enough to the fish to see that they are diseased. Also diseased fish may be ready prey for other fish and thus be noticeable only when large-scale infections are present. The mortality of adult fishes and the incidence of pathogenic microorganisms has been reported in adult fish, yet we have to rely entirely on laboratory and hatchery rearing experiments and operations to obtain information on the mortality of eggs and larvae. The eggs and larvae of fish collected from the seas have been studied by many ichthyologists. Unfortunately, the small organisms must be collected by nets for examination and it is quite impossible to determine the difference between those which were dead in the water and those killed during collection. An experienced investigator can recognize larvae which are in an advanced state of decomposition, but possible microorganisms specifically responsible for death have usually been masked by the adventitious population of bacteria and other microorganisms which immediately attach to the dead eggs and larvae or are present in the plankton haul. A large bacterial population may not be indicative of an abnormal incidence of diseases in the natural habitat. The shallow bays of the Texas coast and other similar bays may contain up to 100,000,000 bacteria per milliliter of sea water and yet harbor no increased incidence of disease. One would expect that the eggs and larvae would be most susceptible to disease and that the survival of fish would depend on the relative survival of the young forms. Although aquaria research has demonstrated the apparent fragility and critical stages in the early development of fish, very little is known of their diseases. In large-scale hatching of cod, herring, and plaice in Norway, it has been noticed that the gas content and abrupt temperature change will adversely affect the survival of eggs and larvae (Dannevig and Dannevig, 1950). Bacteria and other unicellular organisms will also affect the survival of cod eggs. At the State Fish Hatchery in Arendal, Norway 2- to 3-month-old plaice and herring larvae were presumed killed by Vibrio anguillarum and the myxosporidia Lentospora cerebralis (Dannevig and Hansen, 1952). It is within reason to assume that if the larger fish are attacked by pathogenic or parasitic microorganisms, the eggs and larvae will be similarly affected. It is possible that the vulnerable young forms are more susceptible to microbial attack than the adults. Thus the fluctuations in year classes may be, in part, due to the mortality of the young forms from microbial activities.

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F.

OPPENHEIMER

FlSH SLIME

Very little is known about the relation of slime or mucus excreted by fish to disease. Fish slime is a glycoprotein which is secreted by integument cells (van Oosten, 1957). It is assumed that different types of mucus are excreted by different fish according to the type of mucusexcreting cells. Most investigators have found the slime of fishes to be a good source of food for bacteria and to provide a habitat for numerous species of bacteria which may vary seasonally (Gibbons, 1934). A primary protective mechanism of mucus has been described where the mucus is continually washed away during the movement of the fishes through water which in turn removes the bacteria before they can attack the skin. Large populations of bacteria cause the rapid decomposition of the fishes if they are not properly handled after being caught. The mucus of some eels has a form of protective action by its ability to coagulate mud and particles in the water (van Oosten, 1957). It is possible that the fish can precipitate or coagulate bacteria and thus clear the water. G.

CONTROL

In order to reduce the incidence of nematodes, spread by feces from seals to a number of important food fishes, it has been suggested that the seal be eradicated from vital fishing areas (Scott and Martin, 1957). Alternative measures are rated as ineffective. Candling is employed to detect and remove parasites from cod fillets. In recent years this has become inevitable. Through this method only three-fourths of the infections are spotted. All minor attacks pass unobserved (Power, 1958). ACKNOWLEDGMENT The author is indebted to Drs. Ross Nigrelli and William McFarland for their critical review of the manuscript, to Sandy Wilson for her editorial assistance, and to Professor George Borgstrom who has added considerable material from his per­ sonal experiences and files to make the chapter more complete. REFERENCES

Anonymous. (1960). Bacterial disease attacks bluebacks. Food Field Reporter 10 pp. (August 14, 1960). Arai, Y., and Matsumoto, K. (1953). On a new sporozoa, from the muscle of yellowfin tuna. Bull Japan. Soc. Sei. Fisheries 1 8 ( 7 ) , 293-298. Aronson, J. D. (1926). Spontaneous tuberculosis in salt water fish. /. Infectious Diseases 39, 315-320. Aronson, J. D. (1938). Tuberculosis of cold-blooded animals. Am. Assoc. Advance. Sei. Symposium Ser. 1, 80-86. Baker, J. A., and Hagen, W. A. (1942). Tuberculosis of the Mexican platyfish (Pfotypoecilus maculatus). J. Infectious Diseases 70, 248-252.

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