Earth Life

Fish Scales
If you have ever been to a fish market, or caught your own fish, you will be aware that fish have scales, small almost plastic
flakes that cover their body. Most fish have scales, but not all. Scales effect the fishes life through their cost of manufacture,
restrictions they impose on body movement and through the protection the supply. Larger, heavier scales supply more
protection, but restrict movement, smaller, lighter scales offer less protection but allow for greater freedom of movement,
thus in the Common Eel, Anguilla anguilla, the scales are microscopic.
The Cyclostomes (Hagfishes and Lampreys) have no scales at all. However the fossil evidence suggests that their distant
ancestors did have some.
Scales evolved separately in the cartilaginous fish and in bony fish. The scales of sharks and rays are made of bone and
resemble teeth in that they have a soft central area called the pulp, a middle layer of dentine and a hard outer layer of
enamel. These scales are called denticles and are described as 'Placoid', they protrude through the epidermis and are not
wholly covered by it.
Such scales have a definite size and do not grow along with the animal. Instead when the animal's growth opens up a large
enough space between two or more denticles a new one grows to fill the space. The scales/denticles of cartilaginous fish
are discrete, they do not overlap as do the scales of bony fishes.
In the images below show that part of the scale which is embedded in the skin of the fish as coloured orange, (however in
reality it is not coloured like this) and the part we see as coloured grey. Apart from the placoid scales of sharks and rays,
fish scales are mostly flat and thin, they fit into flat envelopes in the skin so there is no need of a side view as is shown in
the placoid image below.

Most sharks have a complete covering of denticles arranged in a repeating

diamond sort of pattern, sharks also have a thick fibrous dermis (that part of the skin which is below the epidermis) which
supports the scales and helps protect the animal. Other cartilaginous fish that swim like sharks, such as the Guitar-fish
(Rhinobatidae) and the Saw-fish (Pristidae) also have a complete covering of denticles.
However those species that are dorso-ventrally flattened such as the rays tend to have many fewer denticles. In the Skates
(Rajidae) they are scattered in patches across the pectorals and on the head. In the Eagle Rays (Myliobatidae) they are
very few in number and in the Electric Rays (Torpedinidae) they are absent except in the modified form of the tail spines.
The Devil-fishes (Mobulidae) have none at all.
The scales of bony fishes evolved a long time ago and in their ancient form they had four layers, one of dense bone, one of
spongy bone, one of dentine and one of enamel. Such scales are called 'Cosmoid' and they only exist in the modern world
on the Ceolocanth (Latimeria chalumnae) or as fossils.

The scales of the remaining bony fishes have only two layers, a calcified one and a fibrous one. They come in two main
types 'Ganoid' and 'Elasmoid'. Ganoid scales are derived from Cosmoid scales and are the evolutionary older style and are
found on Bichirs, Gar-fish, Sturgeons and Reedfish. They are hard solid scales.
The most common form of scale is the elasmoid scale. It is the thin plate that you find on most fishes. It is often described
as coming in two forms,'Ctenoid', which have a set of fine teeth along the posterior edge, and 'Cycloid' which are simply
rounded on the outer/posterior edge. However there are intermediate forms and and these two terms are really just
adjectives that represent the extremes of a continuum.
The scales of a fish may be all of one kind, perch have ctenoid scales while herrings, minnows and trout all have cycloid
scales, or both type of scales may be found on the same fish, Sea Perches, (Epinephelus sp.), have mostly ctenoid scales
above the lateral line and cycloid below while Dab (Limanda sp.) have ctenoid scales on the upper coloured surface and
cycloid scales on the lower white surface.
Not all fish have scales, some species such the Sun-fish (Mola mola) and the Siluroidei (Naked Catfish) have none at all.
Other species, like the Common Eel appear to have no scales but they really have microscopic scales deeply embedded in
their dermis. Even in those fish that have scales they do not always cover the whole body.
The size, and distribution of scales over a fish's body often, but not always, reflect the way it lives. Thus fish that swim
quickly, or that live in fast flowing waters (Trout, Tuna etc) tend to have small scales, while fish that swim slowly in slow
moving waters tend to have larger scales, i.e. Carp.

In some S. American Catfishes the scales have become modified into bony plates to make armour, this is also the case in
Sea horses and Pipe fish. In these cases the added protection is paid for by reduced flexibility and speed. The scales of
some fish decrease in size from the head towards the tail (Carp and Herring) reflecting the need for greater flexibility
towards the tail of the fish.
While the scales of Tunny (Thunnus sp.) and Mackeral (Scomber sp.) are very small, and those of the Common (Atlantic)
Eel (Anguilla anguilla) are microscopic, other species of fish have very large scales. The Tarpon (Megalops sp.) has large
scales often reaching 5cm (2 inches) in diametre which are sometimes made into ornaments, and the scales of the Mahseer
(Borbus sp.) of India can exceed 7cm (3 inches) in diametre.

The scales of modern fish are embedded in, and grow out of, the dermis and are covered entirely by the epidermis. They
grow as the fish grow, thus, in many cases, they reflect a history of the fish's life. Experts in scale patterns can not only tell
you how old a fish is from its scales, but also how many times it has spawned and if it has been seriously ill.
Mostly a fishes scales lie shallowly in the dermis and are overlapping like the tiles on a roof, making them easily removed
(Herring Pilchard and Sardines for instance) but in other species such as Plaice they are more deeply embedded and do not
overlap.
In other fish, some, as in the Flounder (Platichthys flesus), or all, as in the Diamond Flounder (P. stellatus) are modified into
bony tubercles. Many other fish also have tubercles, such as Turbot and Black Sea Turbot (Scophthalmus spp.). In other
species of fish some of the scales have become modified into shields to protect the lateral line or into spines. In Surgeon-
fish and their allies (Teuthidae) these spines can be razor sharp. The moveable spines of Porcupine-fishes (Diodontidae)
and Puffers (Tetraodontidae) are also modified scales.
Fish Anatomy
Welcome to this brief introduction to the anatomy of fish. As with any group containing 28,000 different species there is a
great deal of diversity amongst the animals we call fish.

Accepting that there is a great deal of variety within the world of fish we may wish to ask ourselves what factors control the
shape that a fish takes. Many factors are involved but perhaps the most important is where the fish lives, i.e. in water. Water
is a very different substance than the air that we live in, it is much more supportive for one thing. This means that it is easier
for fish to fly than it is for us, and they can even float, of course we call flying in water swimming, but the fact remains that
because they live in water many fish can live their whole lives without ever touching the bottom, or resting on anything. The
other side of the coin of this supportiveness is that water is much harder to move through, we can run through air much
faster than we can run through water.
In its simplest form this means that fish don't need legs to stand on, or large wings to fly, small wings (fins) will do fine and
secondly it means that if you wish to move quickly it is best to have a streamlined shape. And this of course is what we see
in most fish, a streamlined shape with no legs and small wings (fins).

Like all creatures fish need to perceive the world they live in, and thus we find they have senses, sight, hearing and smell,
but also, because they live in water, they can sense transverse waves generated by motion and so they have evolved a
lateral line system to do this, and some have even learned to use electricity, another sense that is only really made possible
because water is such a good conductor of electricity.
Within the constraints of the basic form fish have evolved further variation to accommodate different ways of living, some
live on the bottom and have become dorsoventrally flatter, others hide in crevices and have become laterally flattened,
some have lost their streamlined shape to facilitate a life of slow movement, and so it goes on; where you live, what you eat,
who you want to hide from and how you do this all contribute to your eventual shape, both internal and external.
The many forms and shapes of the fish that fill up the waters of our world are not only beautiful but practically evolved to suit
a wide range of different life styles.
The Fish's Fin
Fins are essential to fish, the predecessors of legs they give a fish lift, steering capability, braking and momentum. In some
species they are used to hold on to the substrate while in others they allow the fish to walk, or even to fly. Without fins a fish
would just be a fancy worm.
The earliest known fishes had fins, albeit not as many as modern fish, but even the most ancient and primitive agnatha had
a caudal fin. The cephalaspids had one or two dorsal fins and an anal fin as well as a pair of primitive pectoral fins. By the
time of the placoderms pelvic had also evolved.
The shape and structure of a fish's fins reflect both its lifestyle and its evolution. The ancestors of modern fish, both bony
and cartilaginous, looked, in terms of their fin structure, much like modern dogfish and sharks. The evolution of the modern
design, with a symmetrical tail and highly manoeuvrable body fins was only possible after the evolution of the swimbladder.
The exceptions to this design are the Birchirs, Paddlefishes and Sturgeons, which are among the most ancient and primitive
of the bony fishes and still retain the larger upper lobe to the caudal fin, and some of the flying fish in which the lower lobe of
the caudal fin is larger.
The reasons for the asymmetry in the early fishes, and in modern cartilaginous fish is that in these fish the fins supply all, or
most of the lift to the fish's body. A fish is naturally heavier than water, even with lightweight bones other weight reducing
characteristics they still have a density of around 1.076, whereas fresh water has a density of 1.0 and saltwater of about
1.026. Thus a fish will naturally sink to the sea floor if nothing stops it. If the fish is happy to live on the sea floor this is not a
problem, but if it wishes to hunt nearer the surface then it has to keep making an effort to stay up.
Therefore the early fish evolved an asymmetrical tail which supplies forward momentum with the top lobe and lift with the
smaller bottom lobe, which, not being stiff, flexes in counterpoint to the upper lobe, and pectoral fins that work much the way
the wings of a bird or an aeroplane do. Thus the tail lifts the back of the fish and the pectoral fins lift the front. Of course the
fish has to keep moving, if it takes a rest it will still sink. You will probably have noticed that sharks and rays are either
moving constantly, or resting on the bottom, they do not hover in the water column.
The evolution of the swimbladder, and the neutral buoyancy it gives the fish which possess it, set fish free. They could now
devote the whole of the caudal fin to propulsion and develop the pectoral and pelvic fins to maximise their manoeuvrability,
thus modern fish can swim backwards, sideways, come to a complete halt, hover and turn on a dime. Abilities which have
greatly increased the habitats they can take advantage of and undoubtedly contributed considerably to their being the most
numerous group of vertebrates on the planet.
The fins of cartilaginous fish also differ in their basic internal anatomy from those of bony fish. The fins of all fish are
supported by finrays, however in cartilaginous fish these fin rays are inflexible, unbranched and unsegmented. Whereas in
bony fish they are branched, flexible and segmented, except in the ancient coelocanths whose finrays are unbranched.
Spines are associated with the fins of most fish. They are round in cross-section, stiff and unsegmented. They are an useful
means of defence for a fish because they effectively increase its size without a great increase in living costs. Spines, when
held stiff, make small fish much harder to swallow. In some species these spines are associated with poison glands which
increases their effectiveness.

Some Terminology.

Actinotrichia = Spines
Adipose Fin = A small fin without any strengthening rays that is only found in a few groups of fish such as the Myctophidae,
Osmeridae and Salmonidae as well as some catfish. It is located on the upper surface of the body between the dorsal fin
and the upper lobe of the caudal fin.
Anal Fin = This is a single centrally oriented fin found on the lower side of the fish behind both the pelvic fins and the anus.
Ceratotrichia = The fin rays of cartilaginous fish (inflexible, unbranched and unsegmented)
Caudal Fin = This is the tail fin, which when it is asymmetrical as in cartilaginous fish is termed HETEROCERCAL, and
when it is symmetrical as in most bony fish it is termed HOMOCERCAL
Dorsal = Referring to the upper surface of the body.
Dorsal Fin = This is one, but sometimes two, unpaired fin or fins found on the upper surface, or back of the animal. In the
bony fish they are quite manoeuvrable and can be raised, lowered or undulated. In eels and eel-like fish it may be united
with the upper lobe of the caudal fin.
Lepidotrichia = The fin rays of bony fishes (branched, flexible and segmented)
Pectoral Fins = These are paired fins that are normally located just behind the gills. They are often found lower down on the
body in evolutionarily older fish species and higher upon the body in more modern forms.
Pelvic Fins = Originally found towards the rear of the fish (abdominal placement) these are paired fins. They are absent or
reduced in eels and eel-like fish and in some bottom-dwelling species they have become modified into clasping organs.
They are the most variable, in terms of placement, of all a fishes fins. They can be found near the front of the fish, below the
pectoral fins (thoracic placement), or even, in a few cases, in front of the pectorals (jugular placement).
Ventral = Referring to the lower surface, or belly of the fish.
The fins of modern fish come in an amazing array of shapes and forms, however many of them can be related to the seven
basic patterns depicted below.
The Circulatory System in Fish

As in other vertebrates the circulatory system of fish is comprised of both static and dynamic components. The dynamic part
is the blood with all its constituent parts that flows continuously around the fish's body. The static parts are the heart, the
veins and arteries leading to and from it and the capillaries that connect them. Fish have less blood per gram of body weight
than mammals, normally between 3% and 8% of a fish's body weight is blood, however in the hagfish and lampreys it is
greater, for them between 8% and 20% of the animal's body weight.
The blood of fish consists of plasma and the various cells that move around the body in it, erythrocytes and leukocytes.
Plasma is basically water with a variety of ions (Na +, Ca2+, K+) and small organic molecules such as urea, sugars and fatty
acids dissolved in it. In sharks and rays blood cells are created in three different organs, the spleen, the epigonal organ
which surrounds the gonads and the Leydig organ which is found in the throat near the oesophagus. However in teleost fish
blood cells are normally only produced in the spleen and the kidneys.
Blood cells, the biological boats that sail and trade in the biological sea of blood come in two sorts. Erythrocytes and
Leucocytes. Erythrocytes are by far the most common cells found in the blood plasma and their job is to move gases
around the body. they collect oxygen at the gills and carry it to all the cells of the body, and from these same cells they
collect carbon dioxide which they release into the external waters through the gills. See Gills.
The molecule that allows the erythrocytes to do their job, and which makes them red, is called haemoglobin, or hemoglobin,
and erythrocytes are full of it. The haemoglobin of hagfish and lampreys is called monomeric and consists of a single large
haem-molecule with a molecular weight of around 17,000 daltons. Most other fish have tetrameric haemoglobin which is
built up from from four different protein chains called alpha and beta chains, there are two of each in a single haemoglobin
molecule. This type of haemoglobin is very similar to ours and has a molecular weight of around 65,000 daltons.
Leucocytes are a mixed collection of cells that are united in being much less common that erythrocytes and in not being red
because they do not contain haemoglobin. The five most important types are Granulocytes, Lymphocytes, Monocytes,
Thrombocytes and Non-specific Cytotoxic Cells.
1. Lymphocytes:- are 4.5 to 12 microns in diametre and their job is in defence in detecting and marking foreign particles,
specifically they mediate antibody activity, antibodies are small molecules that find and bind to foreign materials so that
Monocytes can find them.
2. Thrombocytes:- are important in blood clotting, thus they are important in conservation of resources as they prevent
blood loss in case of injury.
3. Monocytes:- are sometimes called Macrophages, they are important in defence, their role is to eat (phagocytize)
anything they come across in the blood that might harm the fish such as bacteria, or parasite larvae.
4. Granulocytes:- are also active in defence, they specifically attack bacteria, but also seem to have a role in controlling
stress, however this is still being researched.
5. Non-specific Cytotoxic Cells:- these are another defence mechanism, this time against tumors and protozoan parasites.
There are always exceptions, this is part of the beauty of life on earth. The Icefish of the Antarctic such as
Pseudochaenichthys georgianus have no haemoglobin in their blood and rely on gases dissolved in the plasma, this helps
them become invisible, or at least harder to see. They can survive like this because 1) they live in very cold water, cold
water has a higher content of dissolved oxygen than warm water, 2) they live fairly inactive lives so their oxygen
requirements are low, their food is plentiful and easy to catch.
 At the centre of the static, comparatively, parts of the circulatory
system is the heart which is normally situated below the pharynx and immediately behind the gills. A fish's heart has four
chambers, but unlike us the chambers of their heart are not all muscular, and hey are not so built into a single organ, rather
they are located one behind the other, this is particularly evident in the sharks and rays.

Sinus venosus. The first chamber is called the sinus venosus, it is the preliminary collecting chamber. In teleosts it is filled
from two major veins called the hepatic veins and the left and right branches of the Curvierian ducts which in turn collect
blood from the paired (left and right) lateral veins the inferior jugulars, the anterior cardinals and the posterior cardinals.
However in the elasmobranchs only one hepatic vein leads into it.
Atrium. From the sinus venosus the blood flows into the atrium. the atrium is the largest of the chambers and only weakly
muscular. It pushes the blood, with weak contractions in the ventricle.
Ventricle. The ventricle is the only well muscled chamber, nearly as large as the atrium it is th work horse of the heart, its
contractions drive the blood around the body.

Bulbus arteriosus. The last chamber of the fish heart is called the bulbus arteriosus in the teleosts, but the cornus arteriosus
in the elasmobranchs. The difference between these chambers is that the cornus arteriosus of sharks and rays contains
many valves while the bulbus arteriosus of bony fish contains none. Both are alike in being primarily elastic and work to
reduce the pulsed nature of the blood leaving the ventricle giving it a more even, constant flow.
When you feel your pulse, or measure a fish's pulse there is a beat and a space, beat - space - beat - space, beat - space,
and so on. The beat is when the blood is being pumped out of the heart, which is when, in fish the ventricle, is contracting,
the space is when the ventricle relaxes and is filled with blood again. (note that a one-way valve at the exit of the ventricle
prevents the blood which was just pumped out flowing back into the ventricle). Scientists call the emptying of the heart the
systole part of the pulse and the filling of the heart the diastole part of the pulse. Systole = beat and diastole = space,
together they make one pulse.
Fish hearts have become more efficient through the ages, gram for gram the heart of a modern teleost fish is 2 to 5 times
more powerful than that of sharks and rays, and 10 to 25 times more powerful than the heart of a hagfish.
From the arteriosus the blood enters the ventral artery which takes it to the gills where it loses CO 2 and gains O2 before
heading out to the body through the dorsal artery. Various branch arteries supply the muscles and different organs of the
body through a network of ever finer capillaries. Further capillaries take the blood, now low in O2 and higher in CO2 into a
series of veins that ultimately join up with the major veins mentioned above, the hepatic veins, lateral veins, inferior jugulars,
anterior cardinals and posterior cardinals.
Not all th plasma of the blood returns through the capillaries, some of it mixes in with the fluid that surrounds the cells of
every tissue and is eventually drained away into the lymphatic vessels. This secondary circulation of lymph vessels
eventually empties into the main blood veins. However because it left the main circulatory system it is unaffected by the
beating of the heart and its flow is slow. In some fish it is helped along by small lymph hearts, muscular valved tubes which
help express the blood into the veins. Lymphatic hearts can be found in various places in those species that have them, in
the Eel (Anguila anguila) has one lymphatic heart situated in its tail.

The Brain and Nervous System of Fish

Being highly complex life forms fish need a brain and a nervous system to control their body's actions. The nervous system
of fish, much like ours, is composed of a central co-ordinating brain, a spinal cord and many, many nerves.
The Brain:- Generally speaking fish have small brains in relationship to their overall body weight. Elasmobranchs (Sharks
and Rays) in general have a slightly larger brain for the same body mass as Teleosts (Bony Fish), however there is great
variety within the teleosts scientists have learned something quite surprising about the Elephantnose Fish (Gnathonemus
petersii).

Lurking in the vegetation and then aggressively attacking anything small that comes near doesn't seem to require much
intelligence. A Pike, (Essox lucius) has a brain that amounts to only 0.077 percent of its total body mass. Which means that
when you catch a 5 kilogram Pike you have out maneuvered a fish with a brain weighing 3.85 grams or one seventh of an
ounce. In comparison a Burbot (Lota lota) is a little smarter because it has a brain that is 0.13 percent of its body weight
meaning a 5 kilo specimen would have a 6.5 gram brain, still its is a pretty dumb fish. Most fish have brains that are less
than 1 percent of their body weight, but not all.
At the other end of the continuum of fish intelligence the electric Elephantnose fish, with a brain that is 3.1 percent of its
body weight it is a lot smarter than the Pike, if it grew to weigh 5 kilos its brain would weigh 155 grams, which would make it
40 times as large as that of a Pike of similar weight. To look at it another way a human being has, on average, a brain that
weighs in at around 2.3 percent of body weight. A five kilo human would have a brain of only 115 grams, which is smaller
than that of the Elephantnose Fish.
Does this mean Elephantnose fish are smarter than people? No. Intelligence is more than just brain mass or even brain to
body ratio, while large brains do make it easier to be smart, it is the complexity of the brain and the relative proportion of
forebrain that are most important. The most complex and most forebrain dominated brains on this planet belong to dolphins
and humans. Nevertheless we can see that some fish are much smarter than others and the Elephantnose fish is known
among aquarists as the most playful and inquisitive of fish.
The brains of cyclostomes (Hagfish and Lampreys) are simple but specifically evolved to suit their lifestyles. For instance
the optic lobe is well developed in the visually oriented Lampreys but indiscernible in the blind Hagfish. In both however the
medulla is large and the cerebellum small. Together the cerebellum and the medulla make up the hind brain.
The medulla controls the operations of the inner organs such as heart rate, blood pressure, digestion and waste disposal. It
is also a relay centre for many nerves sending messages to and from the mid and or forebrain.
The cerebellum controls motor co-ordination (but it does not initiate motor activities). This means it controls the timing and
interaction of muscles once a muscular action has been initiated. The cerebellum is also important in maintaining
equilibrium.
The mid-brain of a fish consists mostly of the optic lobes, which vary greatly in size between species in accordance with
their dependance on sight, and in some species the optic lobes may be so large they completely cover the forebrain. In fish
the mid-brain is important in sorting out incoming information and it is also the main centre of learning (whereas in mammals
it is the forebrain that is the main centre of learning).
The forebrain of fish is dominated by the olfactory lobes which extend forwards and may be placed at the end of stalks.
These olfactory lobes are large in the cyclostomes and very large in the elasmobranchs reflecting the importance of smell to
these to groups of fish. The teleosts, for whom sight is often the most important sense have smaller olfactory lobes.
In many elasmobranchs and some teleosts there exists a cerebrum or pair of cerebral hemispheres These also seem to be
predominantly involved with the sense of smell (in mammals the cerebrum is much larger and involved in planning and
learning). The pituitary also arises out of the forebrain, it plays and important role in the regulation of metabolism.
A fish's brain never completely fills the cranium, the cavity in the skull where it lies protected. The remaining space is filled
up with a gelatinous material. Finally as in all vertebrates the brain, plus the gel, are surrounded by a membrane that helps
keep foreign matter and micro-organisms from contacting this most important organ.
The Spinal Cord:- The spinal cord, or nerve cord is similar in all fish. It is a thick sheath of nervous material that runs from
the base of the brain back along the fish's body through, and protected by, the neural canal of the spinal column. Normally it
extends the full length of the fish's body, but a notable exception to this is the giant Sunfish ( Mola mola) wherein the spinal
cord is actually shorter than the brain. It serves as the basis of many simple responses and as the major link to the brain for
sensory input and brain-mediated responses.
The Nerves :- Apart from the brain and the spinal cord the fish body is supplied with a vast network of nerves, the electric
wires of the body along which messages travel. Nerves are built of of numerous neurons and neurons are a one-way
system, messages either travel to or from the brain or the spinal cord along a particular neuronal path, but never both ways.
Those nerves that arise from the spinal cord are called spinal nerves and those which arise from the brain are called cranial
nerves.
Normally there is one pair of spinal nerves (left and right) for each vertebrae, thus long thin fish with many vertebrae such as
eels will have many more pairs of spinal nerves than a much shorter fish such as a gobi. In fish there are 10 pairs of cranial
nerves all with well defined roles.

Pair 1:- Sensory connecting the nasal organs to the olfactory lobes.
Pair 2 :- Sensory connecting the eyes with the optic lobes.
Pair 3 :- Connecting to muscles.
Pair 4 :- Connecting to muscles.
Pair 5 :- Mixed, part sensory part muscles.
Pair 6 :- Connecting to muscles.
Pair 7:- Mixed, part sensory part muscles.
Pair 8 :- Sensory, connects the brain with the inner ear, important for balance.
Pair 9 :- Sensory, connects the brain with the gills and the palate of the mouth.
Pair 10:- Mixed, intestines, gills, heart and lateral line.

 
The Magnetic Sense in Fish

While human science is only just discovering the basis of magnetoperception in fish, the current evidence suggests that fish
have been using it for some time. Two species which are now believed to sense the Earth's magnetic field and make use of
this information in their lives are Yellow Fin Tuna and Atlantic Salmon. Both of these species make long journeys through
the open ocean.
Yellow Fin Tuna have been scientifically observed to respond to the Earth's magnetic field and in 1984 magnetite crystals
were found in the Dermethoid Bone Sinus of this species. Then in 1990 magnetite crystals were also found in the lateral line
system of Atlantic Salmon (Salmo salar) and in the head of the Rainbow Trout (Oncorhyncus mykiss).
The ability of animals to detect the Earth's magnetic field, and to use information gained in this way is an active field of
research at the moment and undoubtedly the future will answer questions such as; Can fish remember magnetic locations
and find there way back to them? and Does a magnetic sense play role in occurrence of beaching whales, or are they
simply trying to bring to our attention the fact that, or been driven insane by the fact that, we are polluting and destroying the
worlds seas and oceans?
Fish anatomy: The Swim Bladder

Introduction
A swim bladder is a gas filled bag that sits in a fish's body cavity above its guts. The acquisition of a swim bladder, with the
neutral buoyancy it gives to it possessors, was one of the crucial steps in the evolution of modern fish, without it fish would
most surely be far less diverse, in terms of number of species, and of the habitats they make use of, than they are today.

The earliest fishes, as well as modern cyclostomes, sharks and rays all live without a swim bladder. To do this they either
have to keep moving all day every day or live on, or at least rest on the sea floor. This is because a fish is heavy than the
water it lives in, and it will sink if it stops swimming (unless it has a swim bladder). Freshwater has a density of 1.0, saltwater
has a density of 1.026 and a fish has a density of about 1.076 (this is an average figure), also bone is nearly twice as dense
as cartilage (1.1 vs 2.0) so before bony fish could become really successful nature had to solve the problem of their density,
swimming all the time is expensive in terms of energy, and resting on the bottom is not always practical, especially in
deeper waters. Furthermore, when you are using your fins to generate lift, and thus stop your self from falling to the bottom
of the ocean, you can't be using them for anything else.

Thus the evolution of the swim bladder set fish free, they no longer had to keep moving forwards in order to stay at the level
they wanted, and they could now use their fins to help them perform much more complicated manoeurvres, like staying still,
swimming very slowly, turning on the spot, swimming head down, or head up, or even backwards.
In the Beginning
Scientists believe that the swim bladder of modern fish evolved from a lung that early bony fish possessed. Probably these
fish lived in shallow tropical waters that had a low oxygen content, and which even have dried up in the summer, or dryer
season. The possession of a lung allowed the fish to gain essential O 2 from the air. When the environment these fish lived in
changed so that they no longer needed the lung to breath nature slowly adapted it for its new role as a buoyancy organ.
Oxygen is much more plentiful in our air than it is in water, and even though the oxygen content of the atmosphere has
varied over the millenia this was also the situation when these fish were first evolving. On average, in our modern
atmosphere, there are about 210 cubic centimetres of O 2 in one litre of air, as compared with 10 cubic centimetres in one
litre of water (keeping in mind that this is an average figure, the amount of oxygen water holds is dependant on its
temperature and its surface area to volume ratio, cold water holds more oxygen than warm water, and white water, because
of its greater surface area to volume ration holds more oxygen than blue water). The microfauna/flora living in the water also
play a role in regulating the O2 level of a body of water.
 We human beings, and some ancient lineages of
fish, such as the lungfish of South America and Africa possess a ventral lung, and scientists believe that this was the
original position of the lung. However, for swimming fish, in comparison to fish which live on the bottom of a pond, a ventral
lung is a problem in that it makes the fish top heavy and liable to flip belly up. Consider the problems you have trying to sit
on top of a beach ball in the sea rather than merely hanging beneath it. Moving the lung, or swim bladder, to the most dorsal
position possible made life much easier for fish and that is why in modern fish the guts always rest below the swim bladder
in a fish's body cavity, nevertheless they are still a little top heavy which is why they sometimes float upside down when they
are dead.

General Information
The swim bladder of modern fish is an ellipsoidal, gas-filled sac that arises as an extension of the gut. In many species of
fish the swim bladder remains connected to the gut and the fish can use this connection to control the amount of gas in the
swim bladder, this is called the 'Physostomous' condition. These sorts fish mostly live in shallow waters and swallow air at
the surface of the water, this air is then passed into the guts and then forced into the swim bladder. In some Physostomous
fish the swim bladder is constricted near the middle to make it look like two sacs, however gas freely passes from one
section to the other. In other species the connection with the gut is closed and no gas can be moved from the gut to the
swim bladder, this is called the 'Physoclistous' condition. These fish are able to control the amount gas in their swim
bladders by means of one or more areas where the membrane is very thin and richly supplied with capillaries, gas
exchange is through the capillaries and the membrane. The capillaries are both arterial and venous and they are arranged
in a counter-current system to facilitate the creation of high gas pressures within the swim bladder. These areas are known
as 'rete mirabile', some physostomous species also possess one or more rete mirabile.
As sea water is more dense than freshwater, see above, the swim bladders of marine fish are smaller than those of fresh
waters. On average the swim bladder of a marine fish comprises 5% of its volume, whereas the swim bladder of a
freshwater species will make up about 7% of its volume. The gases fish use to inflate their swim bladders are those of the
air around us N2, O2 and CO2, however the proportions are not the same as in the atmosphere and vary between different
species of fish. Thus some deep-water marine fish species have up to 84% O 2 in their swim bladder while some deep-water
freshwater species can have up to 94% N 2 in theirs.

The ability to control the pressure within the swim bladder is essential to fish, the buoyancy their swim bladder gives them
allows them to remain at a certain depth in the water without effort. However because any given pressure of gas in their
swim bladder is only neutrally buoyant for one specific depth if they change their depth they have to change the pressure in
their swim bladder. The pressure exerted by the water on a fish increases by one atmosphere (14.7 psi) for every ten
metres of depth, this means that while the pressure a fish experiences is 1atmosphere at the surface, it is doubled to 2
atmospheres just ten metres down, and doubled again to 4 atmospheres at 30 metres down. This increase in pressure
compresses the gas in the fish's swim bladder so that at 10 metres it is only half as big as it was at the surface, and at 30
metres it is only one quarter as big. Therefore to maintain neutral buoyancy a fish must have twice as much gas in its swim
bladder at ten metres depth as it has at the surface, and four times as much at 30 metres and so on.
This is why physostomous fish give of bubbles as they come to the surface and take a gulp of air before they dive down
again. Obviously a fish rising from a depth of 30 metres needs to loose gas pressure quickly if it desires to maintain neutral
buoyancy, if it does not it risks having its swim bladder greatly enlarged as the pressure is reduced. This is why some fish,
caught at a depth and brought quickly to the surface have their swim bladder expanded to fill their throat and sometimes
even extending beyond the mouth.

The Final Word

Nature is always creating diversity, which means changing things and with 28,000 species to play with there is plenty of
diversity to be found amongst the fish. Although it was the evolution of the swim bladder that allowed modern bony fish to
conquer the lakes, seas and oceans of this world many modern fish have given up their swim bladders. Foremost among
these are many species which live their whole lives on the sea or ocean floor, particularly those species which have partially
sedentary lives such as Blennies, Weavers and Flatheads. Another type of fish that now has no swim bladder are the highly
energetic fish such as Tunny, they have oils in their body that help with buoyancy and they spend their whole lives actively
swimming. For a fish that is always on the move and often changing depth quickly a swim bladder would be more of a
hindrance than a blessing so they have simply learned to do without.

The Fish's Skeleton

Because the term fish includes such a diverse array of animals it is difficult to talk in a general way about an average fish.
Most fish are active swimmers and the shape of their skeleton reflects the shape of their body, which, in most cases is
designed to allow them to move easily through the water they live in. Thus the skeleton of an average bony fish looks like
an arrow where the skull represents the arrows head, the backbone or spine the arrows shaft and the tail represents the
feathers. If we add a few barbs to our arrow to represent the spines that run along the spine we have a fairly good idea of
the basic fish skeleton.

The skeleton of a modern bony fish as shown above, and even the arrow, represent the end point of a long period of
evolution. The skeleton of the first fish was probably very like the skeleton of a modern Hagfish which is little more than an
amalgamation of pieces of cartilage. The skull of a Lamprey is a single cartilaginous trough with a few lobes and spines
while the spine is a simple sheath of cartilage surrounding the notochord. There is also a simple cage of cartilage to support
the front-most parts of the viscera. The skeleton of a Hagfish is even simpler. Sharks and Rays have a more complex
skeletal system, which, for the most part it is still only cartilage and not bone. The skull however is more complex than that
of the Lamprey and is called a chondocranium which surrounds the brain and supports the sense organs. Attached to the
skull are the jaw cartilages, called the palatoquadrate cartilage (upper) and Meckel's cartilage (lower). There are also
branchial cartilages supporting the gills. As fishes evolved the number of bones involved in the head-capsule increased and
their arrangement became more and more complex as Nature experimented with different solutions to the problems of life in
the sea.
 As can be seen in the image to the right, in the primitive bony
fishes the individual bones are larger and the amount of open space is restricted. Also, many of the bones are in different
places, or have different shapes. Look particularly at the Maxilla and the Premaxilla in this picture and in the one below. In
the primitive ray-finned fish the premaxillar is small, not moveable and carries only a few front teeth, most of the teeth are on
maxilla itself. In comparison in the more highly evolved (sometimes called 'derived') rayfinned fish the premaxilla has taken
over job of being the upper jaw bone and it is now larger, much more moveable and carries most, if not all, the teeth. The
maxilla has taken on a supportive role. You should note that all these structural changes accompany, or allow a functional
change in the way the jaw works.
The jaws of the primitive fish work simply, much the way lizard jaws do, whereas the derived fish's jaws are a much more
complex system of levers allowing the mouth to move backwards and forwards as it opens and closes. However within the
28,000 species of fish there is room for a great deal of variety. For instance Eels of the family Anguilidae ( Anguilla rostrata,
Anguilla anguilla) have no maxilla or premaxilla

As can be seen from the image at the top

of the page, and more fully from the image to the right, the skull of a bony fish is a puzzle of extreme complexity with many
moving parts. In this collection of small interlocked bones the fishes strike an ecological balance between strength and
lightness. The light-weight requirement is real because bone, being denser than water requires muscle to move, the heavier
a fish's skeleton is the more slowly it moves, both when escaping a larger enemy, and when catching smaller prey.
The earliest fishes went in for heavy armour, which made them slow moving and restricted them to living only on the bottom
of the sea. The picture below shows an artist's impression of what a cambrian fish of the genus Hemicyclaspis may have
looked like. Whatever the colours of the fish were in real-life isn't really important, what we can easily see here, and what is
known from the fossil record is that Hemicyclaspis had its entire head protected by a heavy, bony shield. Evolution has
created the modern fish, which in most instances, has a very low ratio of bone to muscle, it is fast and highly maneuverable
and it claims the whole of the sea as its domain.

The vertebral column, or spine, of a fish is the main supporting structure for the muscles that the fish uses to swim. In its
evolutionary journey it has gone from being the simple cartilaginous tube of the Hagfishes through the more complex
cartilaginous tube of other early jawless fish to partial ossification (boniness) as in the Ratfish to the fully bony tube of
modern sardines. The individual bones of the spine meet at their round centres, called 'centra' and there is usually one
vertebrae per body segment. Two flattened rods of bone arise from the upper side of the centrum, they are separated where
they arise, but meet a little way above the centrum, the space thus formed is called the 'Neural Arch'. collectively the neural
arches of all the vertebrae form a tube that encloses and protects the spinal cord. In most species of fish the combined bony
rods rise up as a Neural Spine above the Neural Arch. In some species of fish such as the Salmon a second pair of rods
extend downwards creating a 'Hemal Arch' that serves to protect various blood vessels.
Unlike those of mammals and reptiles the vertebrae of fish are not linked together, they are simply held in place by a series
of tendons. In fish both sides of the centrum are concave, the space between is filled with a ball of cartilaginous substance
that holds them a little apart allowing them to flex a bit. (There is an exception to this rule however, Garfish (Lepisostidae)
have interlocking vertebrae much like those of reptiles. In other words the centra of the vertebrae are convex on the anterior
or front face and concave on the posterior or behind face allowing them to fit into each other). The vertebrae that connect
the skull to the spine are called the Atlas and the Axis, as in all vertebrates.

In the picture above we can see representative vertebrae from three fish and two sharks, a Sturgeon, a Cod and a Salmon
a White Shark and an Angel Shark. The Sturgeon, the Salmon and the sharks are seen front on (transverse view) and the
Cod is seen side on (lateral view) to show that the processes, and hence the spines they make up, are not necessarily
vertical.

The sturgeon is an ancient fish and represents the ancestral state, in fact the vertebrae of Sturgeon are not ossified (made
into bone) and all the parts you see are in fact cartilage. The arrow represents the direction of evolution. The notochord is
broken into sections, some of which become the bones of the centrum and the rest becomes the cartilaginous balls that
keep the bones from damaging each other. The dorsal and ventral cartilages become ossified and are then called the dorsal
and ventral processes. In most fish the ventral processes of the caudal vertebrae (tail bones) move closer together to form a
hemal arch, and in some species this hemal arch forms below the other vertebrae as well, as in the salmon. In sharks and
rays you can see transitional states whereby only part of each vertebrae is calcified, these hardened, or bony, parts, which
are normally in the form of rings or struts, are embedded in a matrix of ground material that is still cartilage.

The vertebrae of the trunk (the main part of the body) and the skull support a number of additional sets of bones, all of
which may be present in varying degrees in different fish species. Extending sideways are the ribs which protect the visceral
cavity (the space where the guts are). Reaching up are a series of dorsal spines which maybe, but usually are not, in actual
contact with the vertebrae. The pelvic and pectoral fins are supported by simple pelvic and pectoral girdles which are
attached to the skull. The dorsal fin or fins and the anal fin are supported by spines that may, or may not be connected to
the vertebrae. The tail is supported by the caudal vertebrae (the Hypurals, Epurals and the Urostyle). The Urostyle is the
calcified unsegmented final portion of the old notochord. In those sharks which have highly asymmetric caudal fins
cartilaginous end of the vertebral column often extends into, and supports the larger upper lobe.
  The bones mentioned above extend only a small way, or in the case of most dorsal and anal fins not at all into the fins
themselves are. Instead spines or bristles of toughened cartilaginous material called finrays reach out into the fins from the
bones that remain encased by the flesh of the body. Movement of both the dorsal and anal fins, either sideways, or merely
to raise and lower the fin is made possible by simple hinges at the point of attachment of the finrays to the bones that
support them. For more information on the function of fins see the page on the external anatomy of a fish.
 

Sight in Fish

While water is a better medium for the transmission of sound than air it is a much worse medium for the transmission of
light. Basically water absorbs light. This means that the further a ray of light travels through water the weaker, or more
attenuated it becomes.
While we who live in the air can often see for several kilometres, sometimes even more than 20 km, a fish is lucky if it can
see for 50 metres and generally it can see for far less than this. In muddy, algified or turbulent water visibility can often be
measured in mere centimetres. Furthermore, because water absorbs light, the amount of light available for fish vision
steadily decreases the deeper you go. At depths of between 150 and 750 metres even the clearest water becomes a
twilight zone, and below 1,000 metres you are in the realm of constant night. Nevertheless many fish some, or all of, their
lives at much greater depths than this.
As you know visible light is composed of a range of wavelengths, with violet being the shortest and red being the longest.
Water absorbs the red end of the spectrum more easily than the blue end, at a depth of only 1 metre 25% of the red light
entering the water has already been absorbed. What this means in practice is that the deeper you go the less color you can
see, and below 100 metres there is no real color vision at all.
For sharks and Rays this is not a problem as they do not have color vision anyway. The eyes of shallow water species are
adapted to have a maximum sensitivity to light of around 500 nanometres, and the eyes of those species that live in deeper
waters are adapted to 475-480 nanometres. Most Teleost fish however have color vision, and the eyes of the Trout
(Onkorhynchus mykiss) have three sensitivity peaks at, 455, 530 and 625 nanometres.
It should be no surprise then that sight is often far less important to many fish, as a means of perceiving the world around
them, than sound, touch, taste (chemistry) and the lateral line system.
Despite all this most fish have good eyes, the exception to this rule being the Hagfish in whom the eyes are vestigial. The
eyes of fish are in fact very similar to our own and those of other vertebrates. The main differences being: 1) They have no
lachrymal glands (tear ducts), living in water which is constantly washing their eyes they have no need of them; 2) They
have no eyelids, although some species do have extensions of the skin that cover part of the eye, and some sharks have a
nictitating membrane which can be pulled down over the eye. Scientists believe that this is mostly to protect the eye during
feeding.
The basic teleost eye, see diagram, like all vertebrate eyes, consists of an outer case called the sclera, which is only
transparent in front of the lens (when it is called the 'cornea'). The cornea has a constant thickness, unlike ours which is
faintly lens-shaped, hence it does not refract (change the path of) light rays passing through it.
On the inside the 'sclera' lies another layer of tissue called the 'coroid' layer, and inside this there is the 3rd layer, this is the
light sensitive part o the eye, called the retina. The internal part of the eye is divided into two sections by the lens and the
iris. The small part of the eye in front of the lens is filled with a fluid called the 'Aqueous Humour' and the space behind the
lens, which makes up most of the body of the eye is filled with a fluid called the 'Vitreous Humour'.
The iris of the fish's eye is normally immovable in Teleosts, and very slowly moveable in Elasmobranchs, (sharks and rays),
it takes about one hour to change from its most open to its most closed. In comparison the iris of mammals can open or
close in a matter of minutes.
The lens of a fish's eye is purely spherical, unlike ours, and it has a refractive index (light bending ability) of 1.65; this is
higher than that of any other group of vertebrates. Furthermore the lens is fixed in its shape, meaning its shape cannot be
adjusted to facilitate focusing on nearer or more distant objects.
When a mammal wants to change the focus of its eye from a nearby object to a distant one it uses special muscles to
change the shape of the lens, thus changing the part of the light entering the eye which will be land perfectly on the retina.
Because the lens of a fish's eye is fixed in shape it cannot do this, so instead it moves the lens backwards or forwards.
In teleosts the lens is pulled back towards the retina by muscles called retractors, whereas in sharks and rays it is pulled
away from the retina by muscles called protractors. Naturally, when these muscles are relaxed the lens returns to its normal
position.
An interesting aspect of the design of the fish's eye, and its placement on the side of the fish's head is that it is bifocal. This
is because, while the fish's lens is spherical the eye itself is elliptical. This means that light entering from some directions
has a short journey to the retina than light from other directions. What this means to the fish is that its eyes are close
focused to light from in front, but distanced focused to light from the side, or from behind.
Also, because the lens is spherical it tends to protrude through the iris, and this combined with the whole eye tending to
protrude from the fish's head, and the normal sinuous nature of a fish's movement means that fish often have pretty good all
round vision.
Whilst most fish have their eyes placed on the upper side of their heads, some interesting variations have evolved. A deep
sea fish called Opisthoproctus soleatus has telescopic eyes that are placed on its head so that they look straight up towards
the sky. The Mud Skippers, Periopthalmus sp., have their eyes placed on prominent turrets allowing good all round vision.
When they are out on the mud they can also withdraw their eyes into the turret to clean and lubricate them.
The Four-eyed Fish (Anableps tetrophthalmus) has eyes that rise above its body, each eye is divided into two parts with the
upper part being adapted for seeing in the air and the lower part being adapted for seeing under water (like whirly-gig
beetles of the genus Gyrinus). In the young of Idiacanthus fasciola the eyes are found on the end of stalks projecting from
the upper sides of the head. Finally many of the deepest living fish, and those that live deep in caves have very small and
often non-functional eyes.
The retina of the fish's eye is made up of 'rods' and 'cones', the rods detect only the presence or absence of light and the
cones detect colour. As I said above, most fish, except for sharks and rays can detect colour. Rods are much more common
than cones, and usually there are 4 or 5 rods for every nerve cell, whereas cones normally have only one per nerve. The
ratio of rods to cones is very variable between species, but as a general rule, the deeper a fish lives the less cones it has.
Within the cones the light sensitive chemicals, the ones that actually catch the light and convert it into an electrical impulse,
Rhodopsins in fresh water fish and Porphyropsins in marine fish, however fish that live below 500 metres often Chrysopsins
instead as these chemicals are particularly sensitive to the blue end of the light spectrum.
In most Teleosts a posterior part of the choroid layer is enlarged and filled with blood vessels, this is called the 'Falciform
Process'. The nerve that carries the electrical impulse from the retina to the brain is called the 'optic nerve', and the part of
the brain that receives this data is called the 'optic lobe'.
Many Teleosts and most Elasmobranchs possess a 'Tapetum lucidum', a layer of reflective material at the back of the eye
that sends light back passed the rods and cones. In Elasmobranchs the tapetum lucidum is found in the choroid layer of the
eye and is composed of guanine crystals. In Teleosts the tapetum lucidum is located in the retina (behind the rods and
cones) and is either composed of a) guanine crystals as in Bream ( Abramis brama) and Anchovy (Anchoa mitchilli), or b) a
melanoid lipid as in Garfish (Lepisos teidae) and Catfish (Siluridae).
In some fish other parts of the body may be sensitive to light. In Scyliorhinus species, some other juvenile fish, and in some
sharks, the 'Pineal Gland' (in the centre of the forehead) is light sensitive. Larval Sea Lampreys have light sensitive cells in
their skin, even in their tail.
Some fish, such as Rainbow Trout and Goldfish, and possibly many others, can detect UV light.
Many fish, even when unmoving have a greater monocular field of vision than man, but much less binocular vision.
Visual Range in Men and Fish
Entity Horizontal Vertical Binocular
Man 154° 150° 25°
Fish 165° 134° 12°

In some deep water fish the eyes are very large, with large lenses focusing on small retinas. This greatly increases the eyes
sensitivity to light, in Myctophium rissoi the diametre of the the anterior part of the eye is equal to 50% the length of the head
of the fish.

Hearing and Sound Production in Fish

Sound travels further and faster in water than it does in the air, about 4.4 times as fast. Because of this, and because
visibility is greatly reduced in water, it is not surprising that sound is an important part of how fish communicate with each
other and with the world around them. The seas, lakes and rivers of the world are often alive with a riot of noise, but not all
of this is made by fish. Besides other organisms there is also the noise that water makes as it moves against solid objects,
or is moved by the wind and in our modern world the noise made by mankind and his machinery.

Many things can effect the speed of sound, including not only the nature of the medium, (gas, liquid or solid) but also its
temperature and any other additive substances, such as salt in water. Basically sound travels faster through denser and
hotter materials. See table below.
Medium Temperature Speed in M/s
Air 0 331.4
Air 20 343.6
Air 30 348.7
Fresh Water Unknown 1,493
 Sea Water Unknown 1,533
Diamond Unknown 12,000
Like us fish produce sound in two main ways, intentionally and unintentionally. Unintentional sounds are produced by fish all
the time, mostly by swimming and feeding. However they make a far greater variety of sound intentionally in their efforts to
communicate with the other creatures living in their world.
The sounds that fish make are usually simpler than the complex songs of birds, or the calls of mammals. Scientists usually
describe them as grunts, scrapes, knocks, clicks, squeaks, groans, booms, thumps, rumbles and drumming. Nearly all the
species studied so far produce their sounds using either their teeth, their swimbladder or a combination of both. Those
species that produce sounds using their swimbladder usually have special muscles attached to it for exactly this purpose.
Fish create sounds for several different reasons, to stay in touch with the shoal, to warn shoal-mates of danger, to attract,
communicate with and stimulate mates, to scare intruders away from eggs and young and possibly even to echolocate in
some deep-sea species. (The Ecology of Fishes, G. V. Nikolsky, 1963. I would like to find more information on this last
point.)
Some fish are capable of making very loud sounds. One of the noisiest fish in the oceans is the Oyster Toadfish, Opsanus
tau. Because of their noisiness Oyster Toadfishes were studied by the US Navy, they kept hearing them on their sonar, and
it has been claimed that measured from a distance 60 cm the volume of sounds produced by the Oyster Toadfish can reach
100 decibels, which is equivalent to a piece of heavy machinery.
Of course many fish try to take advantage of the sounds other species make. Thus some sharks will use sound to help them
locate their prey while some smaller fish can detect the sounds larger predators make in their hunting. Recent research has
shown that some Clupeid fish, (Herrings and Shads) can detect the ultrasonic echolocation sound produced by hunting
dolphins from a distance of up to 187 metres.

Hearing

Fish hear with their ears, which, although similar to

ours in their basic functionality are missing both the pinna (earlobe), and the middle ear with its eustachian tube. The inner
ear, which is responsible for both hearing and balance is located behind the eye. The modern fish's ear consists of 3
chambers called the 'utriculus', the 'sacculus', and the 'lagena'.
The utriculus, and the 3 semi-circular canals associated with it, are entirely concerned with balance. The three semi-circular
canals are arranged in three different planes of orientation, just as in the mammalian ear, one horizontal and two vertical,
but at right-angles to each other. While this is true for the Osteichthyes and the Chondrichthyes the more ancient Lampreys
have only two canals, the horizontal one is absent, and the hagfish have only a single canal.
These canals, which are sometimes collectively called the Labyrinth, have a bulbous area at each end called the ampulla.
These ampullae each contain the cupula. Movement of the fish's body, in any plane, causes movement of the fluid inside
the canals. The movement of the fluid causes the cupulae to move, when the cupulae move they press against sensitive
hairs, which in turn send electric impulses to the animal's brain.
The 'utriculus', the 'sacculus', and the 'lagena' each contain an 'otolith' or ear-stone. This is normally calcareous and
therefore much denser than the lymph fluid that fills the chambers or even the fish's flesh,. Movement in the lymph fluid
within the chambers, caused by sound waves, causes the otolith to impact against sets of sensitive hairs within the
chambers, which then also send messages to the fish's brain. Expressed again in slightly different way, the otoliths hang in
their chamber of fluid, passing sound waves cause the molecules of the water to oscillate, the exact movement is
dependant on amplitude and wavelength of the sound waves, this movement of the water molecules causes the fish to rock
back and forth in the water a little and because the otoliths are denser than the lymph fluid they move less and or more
slowly and thus impact on the sensitive hairs.
It is interesting to note that the shape of the otoliths is different for nearly every species of fish, and scientists can often
identify a species of fish just from the otoliths, quite why this should be so is a mystery waiting for a scientist to unravel it.
The basic fish ear is fairly limited in its range of use. However many teleost fish have their inner ear connected to their swim
bladder, either directly, as in fish like herrings, or indirectly via a chain of small bones called Webarian Ossicles. This allows
them to use their swim baldder as a sort of drum to detect a greater range of sounds.
In the cartilaginous fish, which have no swim bladder, this is obviously not possible. Instead they have duct, filled with
endolymph, that connects the sacculus to the external environment. This duct passes through an area called the parietal
fossa, this parietal fossa is covered by a layer of skin which may act as a sort of ear drum. Nevertheless Sharks and Rays
have a relatively low hearing range, even though they have been shown to directionally locate sources of low frequency
sound pulses from up to 100 ms. The shark's ear also contains an organ called the Macula Neglecta whose exact function
is not known, however it is suspected that it may well respond to movements in lymph fluid caused by sound waves
contacting the skin that covers the parietal fossae.

Hearing Ranges in Fish

Fish can hear a wide range of sounds, however there is a great variety of capabilities between species. The data in the
table below has been collected from a number of papers and serves well to demonstrate great variability in hearing potential
within the world of fish.
The Hearing Ranges of Fish
Common Name Scientific Name Hearing Range in Hz
Atlantic Salmon Salmo salar, (Lin. 1758) 40 to 350
Bonito/Tuna Euthynnus affinis, (Can. 1849) 100 to 900
Red Piranha Pygocentrus natereri, (Kner. 1858) 80 to 1,500
Goldfish Carasius auratus, (Lin. 1758) 40 to 3,200
Brown Bullhead Amereius nebulosus, (Lesueur 1819) 100, to 4,000
Stone Moroko Pseudorasbora parva, (T & S. 1846) 100 to 8,000
Atlantic Cod Gadus morhua, Lin. 1758 20 to 38,000
American Shad Alosa sapidissima, (Wilson 1811) 200 to 180,000
Gulf Menhaden Brevoortia patronus, Goode 1878 200 to 180,000

Electrical Sensing in Fish

Nerve impulses are electric impulses, and water, especially if it contains dissolved minerals (called electrolytes), is a good
electrical conductor. This means that every time an animal that lives in the water uses a muscle it gives off a minute
electrical signal. It should come as not surprise to learn that many species of fish have learned to detect these signals.
With electrical sensing a fish can detect and locate prey that is hidden from sight and against the direction of the flow of the
water, perhaps buried in the mud or sand even if their prey item is not moving. Rays in particular use electrical sensing to
detect buried prey. Electrically sensitive fish can also detect the approach of both conspecifics and predators.
In most electrically sensitive species the sensors, called 'external pit organs' in teleosts and 'ampullae of lorenzini' in
elasmobranchs are located on the animals head, however they may also be scattered over the body, or along the lateral
line. The sensors consist of canals of electrically conductive gel that open to the water at certain points called pits.
It is known that fish can detect the weak electrical field generated by their own movement through the Earth's magnetic field,
an ability which would obviously be useful for migratory species. Species which can sense electrical impulses of other
animals but not generate their own special fields include Sharks, Rays, Skates, Catfish and Paddlefish.
The next step in evolution is to generate an electrical field specifically for the purpose of electrolocation. Fish that can
purposefully generate an electrical discharge in excess of the normal electrical patterns given off by all living things can
perceive, not only living organisms, but all the world around them electrically. This has happened in 2 families of Rays and
in 10 families of Teleost fish. Although the most famous electrical fish are the Electric Catfish of Africa (Gymnarchus
niloticus) and Electric Eels of South America (Electrophorus electricus) many other interesting, albeit less powerful, species
exist.
Family Common Name No. Species Region
Torpedinidae Electric Rays 14 Marine, various
Rajidae Skates 200 Marine, various
Mormyridae Elephant Fish 198 F.W., Africa
Malopteridae Electric Catfish 2 F.W., Africa
Gymnotidae Naked-backed Knifefishes 9 F.W., S. America
Apteronotidae Ghost Knifefishes 64+ F.W., S. America
Hypopomidae Electric Knifefishes 35+ F.W., S. America
Sternopygidae Glass Knifefishes 41+ F.W., S. America
Rhamphictthydae Sand Knifefishes 6 F.W., S. America
Uranoscopidae Stargazers 4 Marine,
1. The three families with + contain many undescribed species.
2. The Uranoscopidae contains about 50 species but only the four are electrically sensitive.
These fish have electricity generating cells that constantly give off an electrical pulse that in tern creates an electrical field
around them. Everything that touches this field changes the feedback image the fish receives from its field. thus the fish can
build up an image of its world that is not dependant on sight, sound, smell or touch. Such an ability is very useful if you live
in muddy water, as most electrical teleosts do. Also most electric fish are nocturnal. Furthermore these fish can control and
change their own electrical field to improve their image of the world, or to change another electrically sensitive fish's
perception of them.
Apart from generating an electrical field for the purposes of perception many electrical fish can release an electric discharge
into the water around them. By doing so they can both deter predators and stun their own prey.
As all muscular movement generates electrical impulses, movement can distort a fish's electrical image of the world. For
this reason most electric fish (fish that generate a special electric field for perceptive purposes) tend to move slowly and to
hold their body straight, using only their elongated dorsal fin to swim with.
Most fish, especially the electrically stronger species, produce their electrical field as a result of timed pulses of electrical
discharge, but a few of the weakly electrical fish use a continual wave to produce their field. Most species, even the most
powerful, normally only produce relatively weak discharges to maintain their field, saving the power bursts for special
occasions.
The cells producing the electricity are called 'electrocytes'. Each electrocyte produces only a very little electricity, to produce
a large voltage a fish must have many such cells arranged in a batteries that are able to discharge simultaneously. These
electrocytes are normally located in the head in marine species and in the tail in fresh water species.
Fish produce their electrical fields at individual frequencies. When two fish of th same species meet who are using similar
frequencies they change their frequencies to make them less similar in order to avoid having their perception jammed by the
other fish's output. In other species males and females use different frequency ranges to help them recognise potential
mates and to avoid jamming each other. In Apteronotus leptorhynchus the females use frequencies between 600 hz and
800 hz while the males use frequencies of between 800 hz and 1050 hz.
Not all fish can produce electrical discharges of equal potential difference, it is ecologically expensive to carry around so
much generating equipment. The champion is without doubt the Electric Eel (Electrophorus electricus) which can generate a
potential difference of up to 600 volts. The Electric Catfish ( Gymnarchus niloticus) can generate about 300-350 volts, a
Torpedo Ray about 220 volts and some Stargazers can produce 50 volts, however most electric fish are limited to between
8 and 40 volts.
The two largest electric fish, The Electric Eel of South America and the Electric Catfish of Africa prey specifically on the
smaller and weaker species of electric fish. They can turn off their own electrical field to help keep their presence unknown
to their prey and then when a smaller species comes close they suddenly release a large killing discharge. The smaller,
weakly electric fish have been observed trying to hide from such predators by also turning their electrical field off, or by
turning it of and on in a irregular way.
The voltage of an electrical discharge rapidly diminishes as it travels away from its source. Electrolocation is ultimately a
close range sense with a working sphere for most species of between 1 and 2 metres.
Finally, as I said earlier, anything within a fish's electrical field with a different conductivity will distort the the electrical flow
pattern set up by the fish and thus change the field potential around the fish. Detecting these continual changes allows the
fish to map the physical world around it. Of course a fish's ability to make sense of this flow of information is dependant on
its brain size an complexity.
Mormyrids are well known for their large brain to body weight ratios which are far higher than in any other fish, equal to a
human beings in fact. Does this equate with higher intelligence? The answer seems to be yes. Aquarists (people who keep
fish) have known for a long time that the Nile River basin Mormyrids, which use more complicated signals than the Amazon
River basin Gymnotids, are playful. They are well known for making toys out of objects in their aquaria, carrying them
around with their noses. Also scientist have long considered the Mormyrids to be the easiest of fish to teach tricks in the
laboratory. Both playfulness and learning are signs of intelligence.

 
Ionic Balance and Osmoregulation in Fish

Fish live in water, but so in a way do we. We carry our water around with us, but we inevitably loose some and we need to
take more in. This water we have as a part of our body is essential to us, even a 10% loss can be very dangerous for us.
Scientists tell us that 70% of our body is water. Something similar applies to fish, they too are mostly water, in their blood, in
every single cell and around the outside of every single cell there is water. Water is the cradle of life.
The water that fish live in, and even the water we drink is not pure H 2O. It is the nature of water for mineral ions (Na +, K+,
Mg2+, Cl- SO42- etc) to dissolve in it, in brief it is an excellent solvent. The ions that are dissolved in a body of water give it its
ionic balance. Of course the same applies to the water that invests the cells of our, or a fish's body. We and the fish like to
maintain the ionic concentrations, the ionic balance, of our personal waters at a level that is optimum for our biochemistry.
For most species this internal balance is not in harmony with the balance of the environment. The mechanisms that fish use
to maintain and internal ionic balance that is different to that of the water they are living in is called osmoregulation.
It is easy to understand that fresh and marine waters do not have the same ionic balance, although the balance that they do
have is often fairly stable, but in places where they meet the ionic balance is often highly variable over time and place. All
this makes problems for the fish, which over the millions of years of their evolution they have solved in a variety of ways.
The ionic balance of sea water is about 1,000 milligrams of dissolved salts per litre, and that of freshwater normally around
8 to 10 milligrams of dissolved salts per litre or mgs/l.
Cell membranes and even the skin of fish is not 100% waterproof. We know that the basic physical laws of the universe
tend to work towards creating an even homogeneous environment, they push towards a balance. Thus water naturally
diffuses from an area of low ionic content towards an area of higher ionic concentration, and ions, if possible, diffuse from a
high concentration towards a lower one.
What does this mean for a fish? It means that if the ionic content of the water it is living in is lower than the ionic content of
its internal environment (fresh water) it will be constantly gaining water, some through its skin, but most through its gills. This
gain in water will change its internal ionic balance and disrupt its metabolism. To avoid this happening it will need to be
constantly pumping water out of its system. If however the ionic content of the water it is living in is higher than the ionic
content of its internal environment (sea waters) it will be constantly losing water, and in order to stay alive it will need to
drink the water it lives in, and because this water brings a lot of salts with it it will need to find a way to get rid of those
excess salts.
You may have noticed that I said 'if' and 'if' in the previous paragraph. This is because not all fish are in one or either of
these situations. It is possible to avoid confrontation with the environmental balance simply by maintaining an internal ionic
balance that is the same as, or pretty close to that of the external environment and this is exactly what the Hagfish do. Their
internal environment has an inorganic ionic balance of around 1,150 mgs/l. They are they only vertebrate to use this
strategy, although it is common amongst invertebrates, which suggests that it is the old way of doing things, more modern
animals have found that their metabolism works better with an inorganic ionic balance of around 350 mgs/l and so they
strive to maintain this balance.
The ionic balance of a body of water is dependant on both its inorganic ions like those mentioned above, and on organic
ions. The amount of organic ions is usually relatively low, but one group of mostly marine fish, the sharks and rays have
evolved to use the organic ions that their body naturally creates to help them avoid dehydrating in the sea. The
elasmobranchs, like the teleosts like to have an internal inorganic ion content of around 350 mgs/l, so to avoid conflict with
sea they raise their overall ionic balance by maintaining a large amount of organic ions, mostly urea, but also some
trimethylamine oxide in their water. A shark has a total ionic concentration of around 1,007 mgs/l. How they avoid poisoning
themselves with the urea is a more complicated question that is beyond the scope of this introduction, but the
trimethylamine oxide is an important factor.
This simple strategy is also used by the ancient Coelcanth (Latimera chalumnae) Interestingly the Bull Shark or Cub Shark
(Carcharhinus leucas), a species that commonly frequents fresh waters as well as marine environments is able to adapt the
amount of organic salts in its internal environment, a Bull Shark swimming 1,000 kilometres up a major river has a urea and
TMAO balance of only one third of what it had when it was in the sea a month or two before.
The marine teleosts however have not gone along this path, they evolved another way of dealing with the imbalance. Their
preferred internal ionic balance is about 350 mgs/l or one third of that of the sea. Therefore they are always losing water,
they compensate for this by drinking water, but because the water is salty they now have too high a concentration of salts in
their internal environment. They solve this problem by actively excreting salts in concentrated form back into the sea. This is
not easy, it is like pushing pebbles up a hill, as soon as you stop pushing they all fall back down the slope again. To achieve
their goal fish have special cells in their gill filaments and the skin of their opercular that concentrate salt and then excrete it.
Because they are pushing against the gradient, this process uses up energy and a percentage of a fish's daily intake of
food, and thus its energy, is spent on the constant battle to keep the salt out.
Freshwater teleosts obviously have a different problem, they are constantly absorbing water involuntarily and have to work
to get rid of it again. They do this by producing copious quantities of dilute urine. A freshwater fish may produce the
equivalent of 30% of its total body weight in urine every day. For example a 1 kg freshwater Pristis microdon, or Largetooth
Sawfish produces about 250 millilitres of urine a day, in comparison a 1 kg marine Squalus acanthias or Piked Dogfish
produces about 8 ml of urine a day and Scyliorhinus canicula or Small-spotted Catshark produces only 3 ml of urine a day.
Most of the later vertebrates like to maintain an internal ionic balance less than that of the teleost fishes, reptiles,
amphibians, birds and mammals all have internal ionic concentration that are normally less than 300 mgs/l. Because the
balance of life is so delicate, and because ionic interactions are so essential to life, so intricate a part of our essential
biochemistry getting the best ionic environment is very important. It seems that the most complex life forms on this planet
have found that ionic concentrations lower than that of sea water, but greater than that of fresh water are the most efficient
to work with. In the fish we can see the direction of change from the earliest habit of simply putting up with the dictates of
the external environment that the first fish inherited from their invertebrate ancestors towards the complex maintenance of
an independent optimum internal ionic environment that is the legacy and blessing of our modern biochemistry.
Fish anatomy 1: The Digestive Tract

As with all animals digestion in fish involves the breakdown of eaten food into its smaller component parts, amino acids,
vitamins, fatty acids etc. which can then be used to build up new fish body. The breaking apart or breaking down of the
eaten material is called anabolism, the building up of new material is called catabolism and these two together make up the
whole of metabolism. Grammatically it follows from this that the respective adjectives are anabolic, catabolic and metabolic.
As anybody who has watched a gold fish knows quite well fish eat and defecate. Like all animals the fish's body is basically
a long tube that is twisted up on itself a bit in the middle and has a layer of muscles and ancillary organs around it. This tube
has the mouth at one end and the anus or cloaca at the other. Mostly we consider the mouth to be the entrance to the tube
and the anus to be the exit, food items come in and faeces go out. Different things happen in different parts of the tube and
for the sake of study and understanding we give the various parts names.
Mouth - Pharynx - Oesophagus - Gizzard Stomach - Intestines - Rectum.

However not all fish have all these parts, some, like many of the Cyprinids and Cyprinidonts, lack a stomach, while a gizzard
is only found in a relatively few species.

The Mouth
Food is brought into the body via the mouth, and the jaws of modern teleost fish are a mechanical wonder, and the way the
many bones work together is quite inspiring. However there is, as always a large variety in fish as a whole and the mouths
of a Basking Shark, a Yellowfin Tuna and a Seahorse are quite different in both form and function. Lips are rare in fish, most
species have a hard edge to their mouth. Some suction feeders that take in small prey items have small protractible lips that
help give the mouth the form of a tube with a circular opening.
The tongue of fish is generally very simple, being a thick, horny and immovably pad in the lower jaw which may often be
decorated with small teeth. In fish the tongue is not necessary for the manipulation of food as it is in terrestrial animals
because the food items remain buoyed up by the water and can me moved threw the mouth adequately by control of the
water flow and the placement of the teeth. The tongues of the Sharks and Rays (Elasmobranchs)are a little more movable.
However the tongues of Hagfish and Lampreys are armed with teeth and highly movable. However the musculature behind
this movability is quite different in the two groups.
The teeth of most fish are the fore-runners of vertebrate teeth with an outer layer of enamel and an inner core of dentine. A
fish may have teeth at the front of its mouth and along the jaws and in the pharynx as well as on its tongue.
The teeth of Elasmobranchs are simply embedded in the gum, and not attached to the cartilage that supports the jaw. In
fish there is a continuum stretching from the Paddlefish Polyodon spathula where the teeth are embedded in the gums and
not connected to the jaw bones at all through a few species like the Pike ( Esox lucius) wherein the teeth are loosely
attached to the jaws by means of fibrous ligaments to the majority of fish which have the teeth ankylosed, or tightly and
immovably bound by fibrous tissue, to the bones of the jaws etc. In a few species of the Characidae the teeth are implanted
in special sockets of the jaw bones.
Most species of fish, as well as sharks and rays, have polyphyodont teeth, meaning the teeth are continually replaced as
they wear out or are lost. In the Elasmobranchs the teeth are arranged in parallel rows situated behind the functional set.
Those teeth waiting to replace lost or damaged teeth lie flat against the gum and point back into the mouth where they may
assist in preventing food from escaping but take no part in biting. When a tooth, or some teeth, need to be replaced the gum
moves forward pulling the new teeth both forward and erect. In true fish the new teeth grow either at the base of the old
teeth, or in between the old teeth when these teeth are not too closely packed. The Holocephali (Chimaeras) and the
Dipnoi/Dipnomorpha (Lungfish) do not replace their teeth but instead have teeth that are continually growing from the base.
The buccal cavity (the empty space in an empty mouth) secretes mucus to aid in the swallowing of food, but there are no
special organs involved, no salivary glands, and this mucus is a lubricant only, it contains no digestive enzymes such as
mammalian saliva does and is lined with squamous epithelium.

The Pharynx
Immediately behind the mouth is the pharynx which is the continuation of the tube started at the mouth and in which the are
found the gill clefts, through which water flows out of the alimentary canal and into the gills. It is short which leads to the
oesophagus. It is lined with squamous epithelium. As I mentioned above the pharynx may possess teeth, both upper and
lower and as many as 4 rows of them. These pharyngeal teeth may be specialised for grinding like molars, comb-like for
breaking up fine materials of sharp and pointed for piercing prey, in some species they are even hinged so that they fold up
to allow food to pass and hang down again afterwards to prevent its escape. For the most part however pharyngeal teeth
seem to have evolved in order to assist in the act of swallowing food.

The Oesophagus
After the pharynx comes the oesophagus, a muscular tube that leads to the stomach. It is constructed of two layers of non-
striated muscle, one of which is longitudinal and the other circular, strangely in some species of fish the longitudinal muscle
is the inner layer while in others the circular muscle is the inner layer. With so many species generalisations only apply to
the majority, there are always exceptions. The Tench (Tinca tinca) for instance is unusual in having striated muscle all
through the oesophagus and stomach ant into the intestines. The walls of the anterior portion of the Oesophagus are lined
squamous epithelium while those of the posterior section are lined with columnar epithelium, the whole contains many
mucus cells, the mucus keeps the food lubricated and helps it to move along the tube.

The Gizzard
The gizzard is really a highly muscular modification of the first part of the stomach. Its main purpose is to grind up coarse
food items into smaller pieces thus facilitating their later digestion. In those fish which have a gizzard, such as Shad, it is the
place where digestion begins because as well as its muscular activity the gizzard also secretes digestive enzymes into the
food.

The Stomach
The stomach of fish is less well delineated than it is in the higher vertebrates, and in some cases it is considered to be
absent. Where a true stomach is found to exist it is a muscular bag, or tube with a highly acidic internal environment. Unlike
the Oesophagus, and the Intestines mentioned below the stomach is surrounded by a triple layer of non-striated muscle.
The outer layer being longitudinal and the inner layer being circular with the middle layer running at an oblique angle to the
two of these. Inside these muscular layer is a layer of columnar epithelium. The acidity of the stomach changes depending
on whether it is full or not. Secretion of Hydrochloric acid is stimulated by the stretching or expansion of the stomach walls
caused by the presence of food, so the the stomach is more acid when it is full that when it is empty. In most fish the pH of
the stomach varies between 2 and 4. The main enzymes active in the stomach are Pepsins. The stomach may have the
form of a swollen tube, it may be U-shaped with the open part of the U facing the mouth or it may be a blind sac with the
entrance and exit valves quite adjacent to each other. The valve between the stomach and the intestines is called the
pyloric valve.

Pyloric Caeca
At th hind end of the stomach, before, just at the beginning of the intestines many fish have some thin blind tubes called
Pyloric Caeca. Not all fish have them, Wrasses, Pipefish and many Catfish do not have any. In those species that do have
the number is variable, and may even differ between individuals of the same same species, the Sand Eels or sand Lances
(Ammodytes sp.) possess only one, Turbot (Psetta maxima) has two, Perch (Perca fluviatilis) has 3 in comparison Whiting
(Micromesistius australis) has around 100 and Mackeral (Scomber ) may have 200. Most sharks and rays do not have
pyloric caeca, the exceptions being the Greenland Shark (Somniosus microcephalus)and some skates. The function of
these Pyloric Caeca is poorly understood, but they may secrete Trypsin and enzymes active in the intestines, it is also
considered likely that they are important in neutralizing the acidity of the chyme (the partially digested food that leaves the
stomach) before it reaches the intestines, where the environment is alkaline in contrast to the stomachs acidity. It is possible
that the pyloric caeca play a fuller or more complex role in the digestive cycle in some groups of fish than they do in others.

Intestines
The intestine is a long thin tube with a thin double layer musculature, the outer layer being longitudinal and the inner layer
being circular. It is the sight of the final digestion and absorption of the food a fish eats. In the sharks, rays and many
piscivorous bony fish the intestine is little longer than the distance to the anus, but it may be longer, and as a general rule
the intestines become longer as the diet moves through being omnivorous to detritivorous to herbivorous. When the
intestines are considerably longer than the body length they are coiled up, or even wound around other organs such as the
swim bladder i.e. Stone Rollers ( Campostoma sp.) Because the absorption of nutrients takes place across its walls it
important that it have a large surface area. In the sharks and rays , and in a few other ancient fish such as Lampreys,
Lungfish, Paddlefish and Sturgeons, the internal surface area of the intestines is greatly increased by a structure called the
Spiral Valve. In effect this is a corkscrew like structure that runs down the centre of part of the intestine, being twisted helps
it to pack more surface area into a given length. The spiral valve is simple in the ancient fish, but often highly evolved in the
sharks and rays.

The Rectum
The rectum is the end of the intestines and through it faeces pass out of the fish's body and into the surrounding water. In
the lungfish, sharks and rays the rectum opens into the cloaca which also receives wastes (urine) from the kidneys and
material from the reproductive organs. In bony fish the rectum reaches the outside environment through the anus, which is
normally situated just in front the urinary and reproductive openings. However in some fish the digestive tract may be curled
back on itself, and in the Electric Eel (Electrophorus electricus) the anus is situated in the fish's throat. Most of what is
excreted by fish is undigested material and dead bacteria. Fish usually convert nitrogenous wastes into ammonia which is
secreted into the water through the gills, 80% to 90% of a fish's nitrogenous waste is dealt with in this way, the rest will be
formed into urea and pass out through the rectum. In sharks and rays all the nitrogenous wastes are converted into urea.
While most, or all of the digestion that occurs within the fish's digestive tract is the result of activity by enzymes produced by
the fish itself it should be noted that many herbivorous and omnivorous fish derive nutrients from the activity of gut
microbes, single celled archaea, bacteria and fungi that feed on cellulose and other plant products that the fish finds difficult
to digest itself and in the process give out excess byproducts such as fatty acids which are useful to the fish.

The Pancreas
The Pancreas is well developed in the lungfish, sharks and rays and most juvenile fish, however in many teleosts it
becomes quite reduced and diffuse in the adults. In sharks and rays it is quite distinct from the liver, but in those teleosts
wherein it is found it is often partially embedded in the liver. The pancreas secretes enzymes such as trypsin (attacks
proteins), amylases (attack carbohydrates) and lipases (attack fats) into the intestines either through sharing one of the
hepatic ducts (those belonging to the liver), or through its own pancreatic duct.

The Liver
Is a large organ that play various roles in the fishes body, it is the site of glycogen storage, it produces a variety of
substances, including enzymes that help with the digestion and it is a major chemical factory producing various hormones
as well as numerous other important molecules. The liver has no specific shape in fish and generally molds itself into the
space around the stomach and the heart however it has a tendancy to reflect the fish's body shape, being long and tin in
eels and wide in rays and skates. The liver is often very large in some sharks and may extend along the body cavity to the
cloaca. The liver usually has two separate lobes, but it may have only one (some members of the Salmonidae) or even
three as in the Mackeral (Scomber scomber). The gall bladder is usually found somewhere within the liver, it secretes
substances that attack fats and help them to be broken down. The liver always has at least one, and sometimes as many as
eight ducts leading into the first part of the intestines. In many cases the pancreas will share one of these ducts.

Gills: Gaseous Respiration in Fish

Just like you and me fish need a constant supply oxygen in the form of O 2 in order to run their metabolism. Without oxygen
they can't turn their food into energy or make any new fish body. All the free oxygen on this planet on this planet, was, or is
being, released into the air by plants, the atmosphere at the moment is about 21% O 2. However oxygen will dissolve in
water, in a similar sort of way that the bubbles in your Coca Cola are dissolved into the liquid, which is mostly water, that
makes up the drink.
Fish could of course breathe air like Seals and Whales, and some do, but if they wish to stay safe under the water for longer
periods of time it would be much easier if they could get the oxygen they need from the water, and this is exactly what they
do. In fact they were doing this long before any vertebrate animals learned how to breath the air.
So fish live in the water and they breathe the water, to do this they have special organs called gills. Gills are wonderfully well
designed, and they have to be because although the water does hold some oxygen it never holds any where near as much
as the air, and to make things more difficult the amount of oxygen a body of water can hold decreases the warmer it
becomes, and also, salt water holds less oxygen than fresh water. If the air above a body of water is 21% O 2, this means
that 210 parts per thousand are O2, but if we do the math for one of the figures below, such as cold salt water we see that it
contains only 7.58 parts per thousand O 2. What this means to the fish is that their gills need to be a lot more efficient at
extracting O2 from the water than our lungs need to be at extracting it from the air.
Comparison of the O2 available in water
with the air above it.

Type of Water Temperature Ratio

 = 4.35% of
Fresh Water 15ºC
the air
= 2.33% of
Fresh Water 35ºC
the air
= 3.61% of
Salt Water 15ºC
the air
= 1.92% of
Salt Water 35ºC
the air

Fish solve the problems of extracting the O 2 they

need from the water they live in in a variety of ways. Firstly they have different life styles, obviously a fish that spends most
of its life resting on the bottom of the ocean waiting for its dinner to swim by needs less O 2 than a fish which actively chases
smaller fish for its dinner. However most of the problem is solved in the design of the gills.
A fish's gills are situated one set on either side of the body and near the back of the head They are open to the gullet at the
front, and open to the external environment behind. They are designed so that water can flow continually passed them,
coming in through the mouth, and/or the spiracle in sharks and their allies, and passing out through the single external gill
opening in fish or through one of the 5 to 7 gill clefts in sharks and rays. In fish there is a bony plate protecting the gills, this
is called the operculum, and it is hinged and has muscles attached to it so it can be regularly opened and closed.
This ability to have water continually passing over the gills is one of the major factors making gills more efficient than lungs.
With lungs the air comes in, fills the space, and then has to be expelled before any more O 2 rich air can be brought in. With
gills there is no time wasted getting rid of the old air/water and no energy wasted reversing the direction of the flow..
 In sharks and rays the number of gills is usually 5,
but there are some species with 6 or 7 sets, in fish the number of gills is 4 on either side of the body. Each gill is supported
by a gill arch and protected by gill rakers. Each gill arch supports one set of paired gill filaments. The gill rakers help make
sure that no extraneous material gets into the gill filaments to clog them up. Each paired gill filament in turn supports
numerous lamellae (sing. lamella), extending out from both sides of the filament body. It's here in the lamellae that the
uptake of O2 actually occurs.
The lamellae are very fine structures, however there exact dimensions depend on the normal activity levels of the fish in
question. The more active the fish the thinner they are and the less distance there is between them. Also the absolute
thickness of the individual lamellae walls varies, this is important in considering to facility with which O 2can diffuse from the
water to the fish's blood, the thinner the membrane the more quickly. and easily the O 2 can pass across it.
Thus in sluggish fish like the American Brown Bullhead (Amerius nebulosus) the lamellae are 25 µ thick, 45 µ apart and the
lamellae walls are 10 µ thick giving you 14 lamellae per mm, whereas in a highly active fish such as the Atlantic Herring
(Clupea harengus) the lamellae are 7µ thick, 20 µ apart and the lamellae walls are >1µ in width, giving you 32 lamellae per
mm.

The presence of all these lamellae greatly increases the surface area of the gills, meaning that a large amount of water is
available for gaseous exchange at any particular moment of time. In active fish, such as the Atlantic Mackerel (Scomber
scombrus), which has nearly the same gill dimensions as the Atlantic Herring, there may be as much as 1,000 square mm
of lamellae surface for every gram of body weight. Such a fish weighing 1 kg will have approximately 1 square metre of gill
surface area. Having such a large amount of gill area obviously helps the fish in its battle to extract enough O 2 from the
water it lives in.
 The Lamellae are basically
collections of blood vessels sandwiched between two sets of membranes. The membranes keep the blood and the water
separate whilst allowing the dissolved gases, O 2 and CO2, to diffuse naturally from one liquid to the other, depending on
which has the greater concentration of either gas. The laws of physics compel gases to seek equilibrium, if you have
different concentrations of a particular gas on either side of a permeable membrane (i.e. a membrane that very small gas
molecules can pass through, but not larger organic molecules) the the gas will move from the side with the greater
concentration to the other without you having to do anything.
To ensure a continuous gradient of gaseous difference between its blood and the water flowing past its gills fish use a
counter-current system. This means that the blood flows along the vessels in the lamellae in the opposite direction to which
the water is passing on the outside of the lamellae.
While it may not seem entirely intuitive that a counter-current system is much more effective than a system in which the both
fluids flow side by side in the same direction, scientists have tested the system by artificially controlling the water flow, and
they have found that it is in fact five times more efficient. The system works because the blood enters the gills as the water
is leaving them, at this point both of them have their lowest O 2 and highest CO2 concentrations, conversely, the blood leaves
the gills as the water is entering them, at which point both have their highest O 2 and lowest CO2 concentrations. Thus the
difference remains, and O2 moves out of the water and into the blood during the whole operation.
Fish can breathe the water by swimming forwards and letting some of the passing water flow in through the mouth, across
the gills, and then out. They can control the amount of blood flowing through their gills, increasing the amount of oxygen
they take from the water if they being more active. When they are moving slowly, or resting in the water fish can breathe the
water by synchronously expanding and contracting the buccal cavity (the mouth and throat), and the opercular cavity, which
is the space behind the operculum where the gills are. In doing this they can pump a continuous stream of water across
their gills.
Sharks do not have an operculum, or an opercular cavity, but have instead a branchial cavity and a series of branchial flaps,
one for each set of gill but which work in a similar way. Sharks also have a pair of spiracles in front of their gills (one on
each side), they can pump water in through these spiracles and over their gills. In those species of Sharks and Rays which
often, or even habitually rest on the sea floor these spiracles are on the top of the animal's head, allowing it to breathe
without sucking in sand which would damage the delicate gills.
Most fish can also absorb some O2 through their skin, and while they are in their larval form this is often the main source of
respiration. The smaller a fish is the higher is its ratio of skin to body mass, however as it grows it increases its mass much
faster than its surface area and soon the skin is not able to take in all the O 2 the body needs and so other forms of
respiration/gaseous exchange take precedence. But in some fish scientists have learned that the skin continues to take up
O2 throughout the animal's adult life, although the O 2 taken up is only used by the skin, and not passed on to the muscles,
two examples of this are the Crucian Carp (Carassius carassius) and the Yellow Perch (Perca fluviatilis). In adult fish, only
the Ameirus melas (check genus spelling) the Scaleless Black Bullhead uses its skin to take up O 2 for the body, in this
species about 5% of the total gaseous exchange occurs through the skin.
  

Sense of Smell in Fish 

Scents disperse more slowly in water than they do in air, nevertheless smell is an important source of information for fish.
So much so, that many fish have four nostrils, two in and two out. In this way they are constantly able to access new scent
information. Also, it is important to remember that no fish can breath threw its nostrils, they are only for detecting smells.
Fish (Osteichthyes) have their nostrils on the upper side of their head, while Sharks and Rays (Chondrichthyes or
elasmobranchs) have theirs on the lower side of their head.
Not all fish have an olfactory pit (= nostrils, olfactory rosette [O.R.] and connective channels), Puffer fish, and their relatives
in the Tetraodontidae have lost theirs completely and so have no sense of smell. Hagfish and Lampreys have only a single
central nostril and O.R.; in Hagfish water in through the nostril, over the O.R. and then continues on into the gut, but in
Lampreys water is pumped in and out of a muscular pocket called the 'hypophysical pouch'.
Sharks and Rays, and some fish, such as Wrasses and Cichlids have only two nostrils, in the elasmobranchs, and in some
of the fish, each nostril is divided into two halves by a flap of skin (an inlet and an outlet half) and water is pumped through,
and over the O.R. by beating cilia.
The 'olfactory rosette' is a specialised area of epithelium within the 'olfactory pit'. It is where the scents are actually detected
and its surface is greatly folded, thus allowing a large number of sensory cells to be packed into a small area. These
sensory cells are basically the same as those used for taste, however nerves from the O.R. lead to the olfactory centre of
the brain, a distinct and separate part of the brain from that used to analyze tastes.

Fish use smell in a number of distinctly different ways.

1. Fish use smell to recognize places in their environment, even over considerable amounts of time, for instance Salmon
remember the smell of the river they were spawned in.
2. Some fish, such as Tetalurus natalis recognise each other as individuals by scent, or smell.
3. Fish use smell to communicate danger, Minnows release and alarm pheromone into the water if their skin is damaged.
4. Fish use smell in reproduction, pheromones released by females can trigger courting behavior in males (Bathygobius
soporator), or appease male aggression to invasive conspecifics (a conspecific is another member of the same
species).
5. Fish use smell to find food. A shark zeroing in on the source of an attractive scent such as blood uses the same sort of
zigzag approach that a male moth uses when homing in on a receptive female. By testing continually across the flow of
the scent, and assessing its changing strength an animal can quickly focus on the source. 
Fish Muscles
Gram for gram fish have more muscle than any other vertebrate, a male salmon or tuna can be nearly 70% muscle, which is
one reason why fish are so good to eat. The muscles of fish are layered rather than bundled as in the other vertebrates.
Each segment, or sheet, of muscles is called a myomere or myotome and is separated from its neighbor by a sheet of
connective tissue. Other sheets of connective tissue, called septa (pl =septum) occur along the vertical midline of the body
separating the muscles of the left and right sides of the body, and horizontally separating the muscles of the upper and
lower halves of the body. The muscles of the upper half of the body are called 'epaxial' and those of the lower half are called
'hypaxial'. Salmon are a particularly good fish for studying the basic shape of fish muscle because the coloration of their
muscles allows you to see the edges of each myotome, as in the picture above and the photos below.

The myomeres are not flat sheets of muscle, but are folded
into a 3-d shape. In their placement along the body they are angled against the line of the body, with the innermost edge
nearer the front of the body and the outermost edge nearer the tail. This means that if you cut a fish in half across the long
axis of the body you will cut through a number of myomeres. This angling, in combination with the complex folded structure
of the myotome means that the muscles fit into each other along the fish's body with the outer edge looking a little like a W
on its side and the extended edges of the fold forming cone like projections. By looking at the two photographs of sections
of a salmon's muscles seen below (longitudinal section on the left (this is rotated 90 degrees to the right so that the epaxial
muscles are to the right and the hypaxial to the left) and transverse on the right) while keeping in mind the diagram of a
single myotome to the left it is possible to get a good idea of basic structure of a fish's musculature. Notice how the muscles
appear to be zigzag in the LS photo and a series of inset ellipses in the TS. However cooking a small salmon, or wild caught
sea-trout, and then carefully teasing out the individual myotomes is a very informative exercise.
Fish muscles come in three different types, red, pink and white. Most fish have a mixture of two, or all three, types of
muscle, but keep the types in discrete groupings, however in the salmonid fishes the red and white muscle types are mixed
to form a mosaic type of muscle. The colours these muscles show is related to the amount of haemoglobin present in the
muscles, with red muscle having plenty of haemoglobin present and white very little, if any. However when looking at fish
muscles it is well to be aware that some fish, that feed on crustaceans, particularly salmonids, develop a pink colour to their
muscles as a result of a carotenoid pigment they acquire from their food, in the same way that flamingoes get their pink
colour from the crustaceans they eat.
              

Red muscle, also known as slow muscle, is red because it has a high number of capillaries present in it and thus has a high
haemoglobin content. Being well supplied with oxygen red muscle is used for steady, constant-effort swimming and is found
in active fish, particularly those that live in the open waters of seas and oceans, nevertheless red muscle seldomly makes
up even as much as 20% of a fish's total muscle mass. White muscle, or fast muscle, has thicker fibres than red muscle and
possesses many less capillaries, and so it has a much reduced blood flow, and therefore, a reduced oxygen availability.
Most white muscle activity is anaerobic (glycogen is converted to lactate). White muscle fibres can produce tensions that
are up to 2.7 times greater than those of red muscle, but they are more energetically wasteful and therefore the cost to the
animal is higher. Finally white muscle can only work for short periods of time, a couple of minutes maximum is not unusual,
before they exhaust their supply of glycogen and need to rest. All this means that white muscles are convenient for short
quick bursts of movement, in which capacity they out compete red muscle easily, but that they are no good for prolonged
swimming. Pink muscle is intermediate between the two, and is good for continued swimming efforts lasting a few tens of
minutes at a relatively high speed. Of course, like all animals, fish use all their muscles in concert as they go about their
daily lives.

The Lateral Line Sense in Fish

As your science teacher taught, a mechanical wave is a rhythmical movement of energy that passes through matter, as the
wave passes it causes the molecules of the matter it moves through to oscillate (either backwards and forwards, or up and
down). The waters of this planet are full of waves, not just on the surface where we see them, but everywhere, these waves
are caused by anything that moves in the water and by the movement of the water itself, and it would be very useful, if you
lived in the water, to be able to detect these waves. The lateral line system is the fish's way of doing this.
By being able to detect these waves a fish can learn a lot about the world it is living in. For instance when a fish swims it
creates a bow wave (amongst others) by pushing some water in front of it. When this wave impacts an obstacle it reflects a
distorted form of its self back towards the fish. By being able to sense this reflected wave a fish can avoid bumping into
things, such as the glass of a fish tank, even when all its other senses are inoperable.
The 'lateral line' system is a way of detecting wave energy as it passes over or bounces off the fishes body. It works using
mechanoreceptors similar to those responsible for the senses of hearing and balance. The basic sense receptor, called a
'neuromast' consists of a cup like base which rests in the epidermis and a cylindrical gelatinous cupola which extends
beyond the epidermis into either, a supporting mass of lymph fluid or, in a few deep sea fish, directly into the water. Any
mechanical wave that passes through the water sets up reciprocal movements in the lymph fluid that surrounds the cupola.
Within the supporting cup are sense cells which possess hairs that reach up into the cupola. When the cupola is moved by
a passing pressure wave the hairs are bent and this triggers a different electrical impulse to be sent to the brain. I said a
different electrical impulse because the neuromasts are constantly sending off a regular series of impulses to the brain. This
baseline signal is changed when the hairs are bent, bending to one side results in an increase in the impulse's frequency
and bending to the other side results in a decrease in the impulse's frequency.
Fish have these neuromasts located all over their body, some are individually housed in shallow pits, but many others are
located in special epidermal (skin) canals that open regularly to the surface through pores. These canals and canal pores
make up the lateral line that we see running along the fish's flanks. However most fish have an additional series of canals
that run over their head. One canal passes above the eye and on to the end of the fish's snout, two more pass below the
eye and then on to the upper and lower jaws respectively. These lines may give rise to a few, or even many, branch lines.
The lateral line system is an essential component of a fish's sensory arsenal and is undoubtedly one of the reasons fish
have been so successful as a group. It is also a wonderful example of Nature taking an already existing adaptation and
retooling it to work in another capacity. 

Are Fish Cold Blooded?: Thermoregulation in Fish

Generally speaking the answer is yes, but sometimes it is no. Most fish ectothermes meaning their core body temperature is
normally close to that of their environment. However this does not mean that they are completely complacent about body
temperature. By choosing a particular environment a fish can gain some control over it body temperature. Furthermore
some sharks, and some fish such as Tuna possess biological mechanisms that enable them to maintain parts of their body
at a temperature greater than that of the environment.

Is body temperature important? The answer is often yes. Everything that happens in the body of a fish or a person is the
result of chemical reactions, and within certain limits chemical reactions work faster the more energy they have and warmth
is energy. So a warm body means that your metabolism works more quickly, food is digested faster, growth is quicker,
muscles respond more quickly and with greater strength, brains think faster etc. However there is also a cost, warmth is
energy, it has to come from somewhere, and that somewhere is our food. To maintain a body at a temperature above that of
the ambient, above that of the surrounding environment we burn food, and the greater the difference between the ambient
temperature and our body temperature the more food we have to burn. So to have a more effectively functioning brain and
muscles we need to keep them warm, but to do this we need to eat more food, and preferably to eat high energy food, and
obviously before we can eat the food it has to be found and captured. So it is a trade off, birds and mammals go for
maintaining a higher body temperature and paying a higher cost, reptiles and amphibians choose ectothermy as do most,
but not all fish.
Behavioral thermoregulation occurs when a fish actively seeks out areas of water with higher or lower temperature. For
instance juveniles of the Bear Lake Sculpin (Cottus extensus) live at the bottom of lakes feeding on the lake floor during the
day. Here the temperature is normally around 5°C, however after dark they stop feeding and rise to he surface where they
hang around digesting their food. The temperature near the surface of the lake is about 14°C. By doing this they are able to
digest much more food during the night than they could have done by staying on the lake bottom and so they can grow
more quickly. The difference is considerable, at 5°C it would take them 33 hours, or a day and a half to digest the food they
can eat in one day, but at 14°C it takes them only 4.5 hours to digest the same amount of food, so by coming to the surface
at night they are making much more efficient use of their time. And then by returning to the colder water as soon as its food
is digested it slows down its metabolism again and thus conserves energy.
Sometimes the strategies that evolve in simple living creatures are truly amazing and almost seem to reflect intelligence.
Another example of behavioural thermoregulation is the Sockeye Salmon (Oncorhynchus nerka). Sockeye Salmon are
crepuscular feeders, meaning they hunt for food, and therefore feed only at dawn and dusk. During the full day and the full
night they rest. Scientists have learned that in Canada, in Summer, when the days are long and the nights are short that the
salmon adjust their behaviour to maximize their efficiency. After feeding at dusk the salmon rest throughout the night at
depths of around 11 metres where the temperature is about 15°C, however after their dawn feeding they retreat to depths of
about 37 metres where the temperature is only about 5°C. Why? Because the night is short they need to be warm to digest
all they have eaten before the next feeding session, but because the days are long they have more time and so can digest
their food fully at a lower temperature, and in doing so they reduce their metabolic costs.
Behavioural thermoregulation is not uncommon in fish, or in reptiles, amphibians and insects. It is easy to implement and
has the advantage of not being permanent, meaning the animal can, in the best environmental circumstances choose to be
warm or cool as it needs. However it has the problem that the environment is not always at its best, and even when it is it
takes time to warm up enough to allow quick muscular action. Animals that practice physiological thermoregulation have the
edge when the ambient temperature is not optimum, which even in the sea is quite often.
Physiological thermoregulation is where the fish controls its core body temperature by means of internal physiological and
metabolic activities. This is also how we maintain our body temperatures, but while it is universal among mammals it is rare
in fish. It occurs in only a few species, all of which are marine and swim constantly. These include various Tuna, some
Mackerels, the Mackerel Sharks (Lamna nasus and L. ditropis) and the Great White Shark (Carcharodon carcharias).
Both sharks and bony fish that maintain an increased body temperature do so by means of a counter-current exchange
system whereby blood vessels carrying blood that is hot as a result of muscular activity pass along side, and give up some
of their heat to, blood that is going to parts of the body the animal wishes to keep warm. The organ where the heat
exchange takes place is called retia mirabilia. A fish may have more than one retia mirabilia, Mackeral Sharks for instance
have three, one in the swimming muscles, one in the body cavity near the guts and one around the brain.
As with any other successful adaptation there are many other minor changes that work along side the major one in
harmonizing the animals metabolism. For example Albicore Tuna have evolved a unique form of haemoglobin that
increases its affinity for O2 as it gets warmer while its CO2 affinity increases as it gets cooler. This is the reverse of the
normal situation, but it does prevent O2 from being lost from the warming arterial to the cooling renal blood in the retia
mirabilia.
The next step up from recycling the heat from muscular effort to heat parts of the body, which obviously only works while the
animal is using it muscles (which is why all the species that do so are constant swimmers), is to use purely metabolic
processes to generate heat. The only example I know of in the fish is the Swordfish (Xiphia gladius). Swordfish have
specifically modified eye muscles with an exceptionally high mitochondrial volume. Mitochondria are the organelles wherein
food is turned into energy, they are found in every cell of your body. The swordfish does not use these special eye muscles
for anything except normal eye movements (which does not generate much heat) and to heat the blood going to its brain. In
other words the mitochondria work purely to make heat for the purpose of keeping the precious brain warm, it is not merely
recycling heat that was a byproduct of normal muscular activity as are the sharks and the tuna. This is much closer to the
mechanisms used by birds and mammals to maintain their body temperatures. Evolution takes a long time in comparison
with a short life spans, but who can say what fish will be swimming in the sea and oceans in another 50 million years.
Remember, as I said above chemical reactions occur more quickly at warmer temperatures, warmer guts digest food more
quickly, a warmer heart beats more strongly, warmer muscles contract more strongly and respond more quickly and warmer
brains work better. The contraction power of the muscles of Bonito ( Sarda chiliensis lineolata) doubles with a 10°C increase
in temperature. All this can give a predator and advantage that makes the difference between life and death. All the fish and
sharks that maintain a raised body temperature are active constant hunters. The tuna and mackerels hunt smaller fish and
the sharks hunt the tuna and mackerel amongst other things. Endothermic animals need to eat a lot more than ectothermic
ones, sometimes as much as 3 or 4 times as much.
 

Bibliography
 
The Fish Anatomy Menu
Anatomy Fins Blood Nerves Magnetism Swim-bladder
Skeleton Sight Scales Hearing Electricity Osmoregulation
Digestion Gills Smell Muscles Lateral Line Thermoregulation
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CLASSES AND ORDERS OF FISHES
CLASS MYXINI (Hagfishes )
 Order Myxiniformes

CLASS CEPHALASPIDOMORPHI (Lampreys )

 Order Petromyzontiformes

CLASS ELASMOBRANCHII (Sharks , Rays , Guitarfish , Sawfish )

 Order Hexanchiformes (Cow and frill sharks )
 Order Heterodontiformes (Bullhead and Horn Sharks )
 Order Orectolobiformes (Whale , Nurse , carpet , goblin sharks , and relatives)
 Order Lamniformes (Tiger, Mackerel , Basking , and Thresher Sharks )
 Order Carchariniformes (Houndsharks , Cat, Weasel, Requiem , Hammerhead Sharks , and relatives)
 Order Squaliformes (Dogfish , Bramble Sharks )
 Order Pristiophoriformes (Saw Sharks )
 Order Squatiniformes (Angel Sharks )
 Order Pristiformes (Sawfish )
 Order Torpediniformes (Electric Rays )
 Order Rajiformes (Skates and rays )
 Order Myliobatiformes (Stingrays )
CLASS HOLOCEPHALI (Chimaeras )
 Order Chimaeriformes
CLASS SARCOPTERYGII (Lungfishes , Coelacanths )
 Order Coelacanthiformes (Coelacanths )
 Order Ceratodontiformes (Australian Lungfishes )
 Order Lepidosireniformes (South American and African Lungfishes )
CLASS ACTINOPTERYGII ( = Osteichthyes , or Bony Fishes)
 Order Acipenseriformes (Sturgeons and Paddlefishes )
 Order Polypteriformes (Bichirs )
 Order Lepisosteiformes (Gars )
 Order Amiifores (Bowfins )
 Order Osteoglossiformes (Bonytongues , Arapaimas , Mooneyes , Elephantfishes , and knifefishes )
 Order Elopifores (Tarpons Tenpounders )
 Order Albuliformes (Bonefishes )
 Order Notocanthiformes (Halosaurs , Spiny Eels )
 Order Anguilliformes (eels )
 Order Saccopharyngiformes (Bobtail Eels, Swallowers, Gulpers)
 Order Clupeiformes (Herring , Shad , Sardines , Anchovies , Milkfishes )
 Order Cypriniformes (Minnows , Carps , Loaches , Suckers)
 Order Characiformes (Tetras , Hatchetfishes , Piranhas , and others)
 Order Siluriformes (Catfishes )
 Order Gymnotiformes (Knifefishes )
 Order Salmoniformes (Pikes , mudminnows , Smelts , Salmon , trout , and others)
 Order Stomiiformes (Bristlemouths , lightfishes , Viperfishes , Dragonfishes , Snaggletooths , Loosejaws , and
others)
 Order Aulopiformes (Greeneyes , pearleyes , Barracudinas , Daggertooths , Sabertooths , Lancetfishes , and
others)
 Order Myctophiformes (Lanternfishes , Brotulas , and others)
 Order Batrachoidiformes (Toadfishes )
 Order Lophiiformes (Anglerfishes )
 Order Gobiesociformes (Clingfishes )
 Order Atheriniformes (Silversides , Rainbowfishes )
 Order Cyprinodontiformes (Rivulines , Killifishes , Goodeids , Livebearers , four-eyed fish)
 Order Beloniformes (Medakas , Sauries , needlefishes , halfbeaks )
 Order Lampriformes (Opahs , Velifers , Crestfishes , Tapertails , Ribbonfishes , Oarfishes , Threadtails , and
others)
 Order Beryciformes (Pinecone fishes, Lanterneye fishes, Squirrelfishes , Soldierfishes , Pricklefishes ,
Bearfishes , and others)
 Order Cetomimiformes (Whalefishes )
 Order Zeiformes (Dories , Oreos , Boarfishes )
 Order Gasterosteiformes (Sand eels , Sticklebacks , Tubesnouts , Seamoths )
  Order Syngnathiformes (Trumpetfishes , Cornetfishes , Snipefishes , Pipefishes , and Seahorses )
 Order Synbranchiformes (Swamp-eels , Spiny-eels )
 Order Scorpaeniformes (Scorpionfishes , Stonefishes , Thornyheads , waspfishes , Velvetfishes , Searobins ,
Gurnards , Greenlings , Cottids , sablefishes , Sculpins , Flatheads, Poachers , Snailfishes , and others)
 Order Perciformes (Perches , Snooks , Basses , Groupers , Soapfishes , Basslets , Flagtails , Bigeyes ,
Cardinalfishes , Tilefishes , Silagos , Remoras , Jacks , Pompanos , Roosterfishes , Dolphinfishes , Snappers ,
Grunts , Breams , Croakers , Drums, Gatfishes , Spadefishes , batfishes , Scats , Butterflyfihes , Angelfishes ,
Cichlids , Damselfishes , Anemonefishes , Hawkfishes , Kelpfishes , Jawfishes , Mullets , Wrasses , Eelpouts ,
Gunnels , Swallowers , Gapers , Sand Lances , Stargazers , Blennies , Kelpfishes , Gobies , rabbitfishes ,
Surgeonfishes ,Barracudas , Mackerels , Swordfish , Marlin , Tunas , and many others)
 Order Pleuronectiformes (Flounders , Halibut , Tonguefishes )
 Order Tetraodontiformes (Triggerfishes , Filefishes , Puffers , Boxfishes , Porcupinefishes , Ocean Sunfishes )
 Order Gadiformes (Cods , Haddocks , Hakes , Burbots , Grenadiers )
 Order Percopsiformes (pirate and trout perches , Cavefishes )
 Order Ophidiiformes (Cuskeels , Pearlfishes )

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MEER Marine biology Table of Inde CD- Link Support
References Books
home home Contents x ROMs s MEER
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 Phylum Chordata
The Phylum Chordata includes some of the most familiar of all animals, - including mammals , birds , reptiles ,
amphibians , and fishes. These groups are all members of the subphylum vertebrata. Also included in the phylum
chordata are two invertebrate phyla, the Urochordata and Cephalochordata , which together with the vertebrates
comprise approximately 45,000 living species of Chordates .

The characteristic features of the phylum Chordata are:

 A dorsal, hollow nerve chord
 A notochord at some time during development
 Pharyngeal gill slits at some time during development
 A muscular post-anal tail at some during development

Although the evolutionary history of the chordates is controversial, they appear to have a common ancestor with most
other deuterostomes , such as the echinoderms , lophophorates , and hemichordates . The chordates are sometimes
placed in the superphylum Deuterostomia , which also includes the phyla Chaetognatha , Hemichordata , and
Echinodermata . Pharyngeal gill slits may have originated in some early deuterostomes as a means of pumping water
through the mouth and out the gill slits in order to increase respiratory gas exchange. Primitive chordates may have then
modified these gill slits to function as filter feeding devices. Along with other basic features of the phylum, early
chordates were thought to have a tadpole-like larval form, similar to that possessed by many modern day tunicate
larvae. The tunicates appear to have been early evolutionary offshoots of the chordate line that lead to the other
vertebrate groups. There are other theories of chordate origins, including close relationships with other groups such as
arthropods and mollusks , however, the echinoderm - chordate theory is presently the most accepted view.

At least some forms of chordates can be found throughout all depths and types of marine habitats, (as well as most
terrestrial and freshwater habitats) that can support life. Fishes, mammals , and birds can be found worldwide in all seas
and oceans. Some forms, such as the reptiles , are somewhat more limited in their distribution, and there are no truly
marine amphibians .

The Phylum Chordata includes:

Subphylum Cephalochordata (about 20 species)

Subphylum Urochordata (=tunicata), about 3,000 species
Appendicularia
Ascidiacea
Thaliacea
Subphylum Vertebrata (about 42,000 species)
Amphibia
Aves
Chondrichthyes
Mammalia
Osteichthyes
Reptilia
Superclass Agnatha
Cephalaspidomorphi
Myxini
Pteraspidomorphi

Fishes
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There are about 25,000 described species of fishes living today, more than the number of species in all other w search_sm

vertebrate groups combined. Ichthyologists (biologists who study fish)have various definitions for what a fish is, 22218516
but the most basic would be as "aquatic vertebrates that have gills throughout life and limbs, if any, in the
befree
shape of fine" (Nelson, 1994). Some ichthyologists exclude the agnatha (jawless vertebrates such as lampreys
and hagfish ) from the fishes, but we include them here as one of five classes of extant (living) fishes. q
 Epinephalus tauvina, class Osteichthyes

Carcharodon carcharias, great white shark, class Chondrichthyes

Fishes as a group are extremely diverse in form, physiology, behaviour , trophic position, habitats, and general life
history. Fishes of the Osteichthyes (bony fishes) include an enormous variety of shapes and sizes, however, a
generalized form that comes to mind for most people might be one similar to Epinephalus tauvina (a member of the sea
bass family, the Serranidae ). The class Osteichthyes contains most of the species of extant (living) fishes.

The sharks , rays , chimaeras , and their relatives are members of the class Chondrichthyes , the cartilaginous fishes,
and consists of about 800 species worldwide.

There are several other groups of living fishes, including the sarcopterygii (lobe finned fishes, such as lungfish and
Coelacanths ), the myxini (hagfish ), and Cephalaspidomorphi (lampreys ).