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BIOLOGY AND ECOLOGY
OF TOXIC PUFFERFISH
Biology and Ecology of Marine Life
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BIOLOGY AND ECOLOGY OF
MARINE LIFE BOOK SERIES
Series Author:
Ramasamy Santhanam, PhD
Former Dean, Fisheries College and Research Institute,
Tamil Nadu Veterinary and Animal Sciences University,
Thoothukkudi-628 008, India
Email: rsanthaanamin@yahoo.co.in
1. Introduction ................................................................................................ 1
2. Toxic Pufferfish: Biology and Ecology ..................................................... 9
3. Profile of Toxic Pufferfish ........................................................................ 17
4. Pufferfish Toxins ..................................................................................... 275
5. Pufferfish Poisoning ............................................................................... 285
6. Pufferfish Aquariums ............................................................................. 301
7. Diseases and Parasites of Pufferfish and Their Management ............ 319
8. Nutritional Values of Pufferfish ............................................................ 359
9. Pharmaceutical Values of Pufferfish Toxins ........................................ 375
References ................................................................................................ 383
Index ......................................................................................................... 403
LIST OF ABBREVIATIONS
INTRODUCTION
CONTENTS
warm seas including coral reefs or seagrass beds. Though most of the
marine puffers are found in coastal regions, some are oceanic (e.g.,
Lagocephalus lagocephalus) or live in the deep sea (e.g., Sphoeroides
pachygaster). Interestingly, puffers are relatively uncommon in the tem-
perate zone and completely absent from cold waters. Only 37 species are
truly freshwater pufferfish and 48 species are in brackish water (http://
animal-world.com/encyclo/fresh/Puffers/Pufferfish.php. Animal-World.
Pet and Animal Information.). The brackish water species are found
primarily in estuaries, where river mouths meet the ocean. Some will
enter periodically for feeding or breeding, and possibly stay as juveniles,
while others spend their entire lives in brackish conditions. Freshwa-
ter species occur in Amazonia, in tropical Africa, in eastern India and
Southeast Asia (Puffer Fish Facts: http://www.snorkelingpattaya.com/
blog/fish/puffer-fish.html).
Chief Taxonomical Characteristics of Pufferfish: Pufferfish range
in size from the 2.5-centimeter-long dwarf, pea or pygmy puffer to the
freshwater giant puffer, which can grow to more than 60 centimeters in
length. They normally have the appearance of a large tadpole, with bulg-
ing eyes and an elongated snout. They may be found in a variety of colors
but can sometimes be hard to identify when they are not inflated. All the
tetraodontids possess robust, oval-shaped bodies and are characterized by
a tough skin that is often covered with small spinulous scales, a beak-like
dental plate (four large teeth, fused into an upper and lower plate) divided
by a median suture, a slit-like gill opening anterior to the base of the pecto-
ral fin, no pelvic fins, no fin spines, a single usually short-based dorsal fin,
a single usually short-based anal fin, and no ribs. Pufferfish are generally
divided into two groups: the short-nosed puffers (family Tetraodontinae)
and the long-nosed puffers (family Canthigasterinae) (http://www.diveg-
allery.com/pufferfish.htm).
Locomotion: Unlike more typical fish, the body of the pufferfish is
rigid. Hence, it largely relies on its fins for motion and balance. It is
driven mainly by the pectoral fins, dorsal fin and anal fin. Caudal pedun-
cle and caudal fin serve as a rudder. Thus, the puffer is indeed quite slow,
but extremely agile. It can both forward and backward swim and climb
up and down.
Introduction 3
removal of toxic internal organs, the meat TTX is less than 2 μg.g–1 and
such species are considered fit for human consumption.
Pufferfish are prepared as meals in the name called “fugu”. This care-
fully prepared dish is named after one of the numerous species from the
Fugu genus among which is Fugu rubripes. Some restaurants in Japan
have specially licensed chefs that prepare “fugu” from very thin slices of
the flesh of certain species of pufferfish. The skin is removed and the flesh
is carefully separated from all internal organs, minimising the amount of
toxin in the portion to be consumed. Fugu is regarded as a delicacy, but
despite the care taken in its preparation, it often produces mild symptoms
due to minute traces of tetrodotoxin remaining in the flesh. In Japan seri-
ous poisoning and death from eating fugu is not uncommon.
Cultured Pufferfish: Three common edible species of pufferfish,
Takifugu obscurus, T. flavidus and T. rubripes are cultured in various loca-
tions in the Yangtze River, the East Sea, the Yellow Sea, the South Sea and
the Bohai Sea of China. These cultured pufferfish, which are very popular
among consumers, not only have a high growth rate but yield non-toxic
meat, which still retains a delicious taste. They are regarded as a high qual-
ity fish because of their high protein content and special flavor.
Aquarium Use: Some pufferfish species can be kept as aquarium fish
in freshwater, brackish and saltwater aquariums. In freshwater aquariums,
they can be used for the biological control of snails.
Toxicity: The toxin of pufferfish is variously used as a defensive bio-
toxin to ward off predation. Predators that do not head the danger signals,
eat puffers anyway and may die from choking, or from toxic poisoning.
Majority of the pufferfish species are toxic with a potent neurotoxin viz
tetrodotoxin (TTX), a potent neurotoxin and the paralytic saxitoxin (STX)
is seen in certain species as the secondary toxin. Depending on the nature
of actions of TTX, Nader et al. (2012) reported three toxic levels as detailed
below:
Weakly toxic (10–100 MU/g tissue) (2–20 µg/g tissue)
Moderately toxic (100–1000 MU/g tissue (20–200 µg/g tissue)
Strongly toxic (>1000 MU/g tissue) (> 200 µg/g tissue)
(1 MU = 0.2 μg of TTX/g tissue)
Introduction 5
Sabrah et al. (2006) on the other hand classified the toxic levels of TTX
as follows:
Non-toxic < 10 MU/g (< 2 µg/g tissue)
Weakly toxic ≥ 10–99 MU/g (≥ 2 – 198 µg/g tissue)
Moderately toxic ≥ 100–999 MU/g (≥200–1998/g tissue)
Strongly toxic ≥ 1000 MU/g (≥ 2000/g tissue)
The above classifications could not be applied presently to the vari-
ous species of pufferfish as the toxic level of different species varies
with their areas of catch or size. Further, detailed research/information is
needed on this aspect. The toxic (poisonous to eat) and less toxic (harm-
less) species among the recorded pufferfish species of the world are
given in the following table.
Among the pufferfish species, only one species viz. Takifugu chinen
sis is under the “Critically Endangered” category of the International
Union for Conservation of Nature (IUCN). The global population of
T. chinensis has been estimated to have declined by over 99.99% over
the last 40 years. This estimate is based on landing statistics from the
East China Sea and Yellow Sea provided by the Sekai National Fisheries
Institute in Nagasaki and the National Fisheries University in Shimono-
seki City. The population decline of this species is largely due to its large
scale consumption and over exploitation. On the other hand, Takifugu
plagiocellatus) is under the “Endangered” category and it is due to its
coral habitat degradation. “Vulnerable” category of 5 species is result-
ing from habitat modifications caused by deforestation and conversion
of lands in agricultural areas, increasing urbanization, and overharvest-
Introduction 7
ing for the aquarium trade. The long felt gap on the conservation status
of this globally important group of fishes calls for further, immediate
research on this aspect.
KEYWORDS
• edrophonium
• golden poison frog
• paralytic shellfish toxin (PST)
• pufferfish
• pufferfish poisoning
• saxitoxin (STX)
• Tetrodotoxin (TTX)
CHAPTER 2
TOXIC PUFFERFISH:
BIOLOGY AND ECOLOGY
CONTENTS
Fins: Fins of the pufferfish are unique and they are used in a different
manner. Puffers use mainly their pectoral fins along with dorsal and
anal fins for moving. The tail fin is used for the purpose of changing
direction only. Such fins allow easy movement even backwards.
Body: The bodies of pufferfish look mostly like a ball, however, they are
perfectly adapted to their environment and living habits. The head is large and
they have big eyes which are wide apart. Such eyes allow puffers to hunt effec-
tively. Also, most of these fish have toxic skin which avoids natural predators.
Teeth: Tetraodontids lack true teeth, the jawbone itself being modified into
four fused beak -like structures. There are two teeth in the upper mandible and
two in the lower mandible. These teeth grow continuously at a faster rate, so
such a diet is essential in order to maintain them at a reasonable length. It is
also crucial to feed them food which is hard enough to naturally grind their
teeth. Such food includes normally aquatic snails, cockles and crustaceans
with shells.
Toxic Pufferfish: Biology and Ecology 11
tion, due to a lack of oxygen getting into vital body organs (Fagundes, http://
www.bv.fapesp.br/en/bolsas/138343/morphological-description-of-diges-
tive-tube-of-sphoeroides-testudineus-linneaus-1758-tetraodonti/; Zhao et al.,
2010).