Fish

Introduction and Evolution

Fish, particularly the jawless hagfishes (Class Agnatha), represent the most primitive
vertebrates and, as such, demonstrate many of the early origins of systems which are
found, often modified, in the higher vertebrates. Thus, some knowledge of fish anatomy
provides a better understanding of structures in the higher groups.

Fish are the most diverse of the vertebrate groups with over 20,000 living species known,
which can be classified into three main groups:

(a) Class Agnatha: jawless fishes.

Oldest and most primitive class of vertebrates
Lack jaws and pelvic fins
Many lack pectoral fins
Order Cyclostomata
Lampreys and hagfishes

(b) Class Chondrichthyes: cartilaginous fishes.

Skeleton entirely cartilaginous
Placoid scales
Approximately 700 species of sharks, rays & related fishes
Lack lungs and a swim bladder
Males have a clasper on the pelvic fin
Transfers sperm to female – fertilisation internal

(c) Class Osteichthyes: bony fishes.

Skeleton contains a significant amount of bone
Various types of bony scales
Most species possess lungs or a swim bladder
Most species oviparous & fertilisation is external
Terrestrial vertebrates evolved from early members of this group (Crossopterygii)
Subclass Actinopterygii: ray-finned fishes
e.g. trout
Subclass Sarcopterygii: fleshy-finned fishes
e.g. lungfishes

Fish are becoming steadily more important as domestic animals - both in terms of fish
farming for protein or sport, and as pets kept in aquaria.

A note on terminology: fish or fishes?! Generally, the plural of fish is fish if you are talking
about several individuals of the same species, and fishes if there are several individuals
from more than one species.

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General Anatomy and the Integument

Body Shape

The body of a fish is generally torpedo-shaped, tapering towards the tail. Pressure drag is
lowest when the body is long and slender, while frictional drag is lowest when the body is
short and plump so the fastest species find a compromise between the two. Projections
from the body decrease streamlining and are therefore minimal. The tail, as the main
propulsive organ, is an integral part of the body and not just an appendage – blue marlins
can attain speeds of up to 80km/hr.

Skin

The body is covered with a distinct and fairly strong skin. In many fish (not extant
agnathans), this is armoured with scales which are dermal in origin. The epidermis
consists of non-keratinised stratified squamous epithelium with scattered glands which are
mostly unicellular, but sometimes multicellular. These cells secrete the mucous that covers
most fish. This mucous layer serves several important functions: keeps the fish clean and
provides a barrier to bacterial infection, reduces drag when swimming, assists in
osmoregulation, makes the fish slippery and so difficult to grasp by predators, and may
also contain toxic or offensive chemicals which deter predators. The 50 cm hagfish Myxine
glutinosa can secrete a full bucket of mucous in response to predation.

Unicellular gland cells, known as club cells, secrete chemicals which induce fear in other
fish and so are thought to warn others of approaching danger. Poison glands, found in
many fishes, are derivatives of skin mucous glands. These glands are often associated
with fin spines which have grooves to allow the venom to be injected into the victim (e.g.
scorpion fish and stone fish).

The epidermis does not become keratinised, but remains soft and thus is easily damaged.
Remember that epidermal tissues do not contain blood vessels and so may be badly
injured without bleeding – this is significant when assessing injuries to fish skin. Since the
skin forms intimate contact with the environment, despite its mucous coat, it is still
susceptible to a variety of diseases (parasitic, bacterial, viral and fungal), particularly as
secondary infections after injury. The dermal layer is thin and contains connective tissue,
blood vessels, nerves and the scales (see Figure 2).

Many species have extensions of the skin into barbels or flaps which carry sensory organs
or are used as accessory feeding structures (e.g. catfish).

Colour

The colour of fish is primarily due to dermal pigments, although the colour of underlying
tissues may also contribute. The pigment producing cells, called chromatophores, are
derived from neural crest cells and can occur anywhere in the skin but tend to be in the
dermis just below the epidermis. By altering the concentration and dispersion of pigment
granules in the chromatophores, fish can change colour and pattern. These cells are
controlled by both the endocrine and nervous systems. There are three types of

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chromatophores: melanophores (black pigment), xanthophores (yellow pigment) and
erythrophores (red pigment). The skin of fish may also contain iridophores, which consist
of crystalline guanine platelets that refract light, producing a silver or iridescent skin. A
fish‟s colour is the combination of chromatophores and the colour of the reflected light: for
example, a green parrot fish is blue light filtered through yellow xanthophores.

Scales

There are several types of scales among the fish groups and they have all arisen from a
primitive model that consisted of four layers of hard tissue: a layer of enamel that covered
an inner layer of dentine which, in turn, enclosed a layer of vascular bone; this vascular
bone was then supported by a layer of lamellar bone. Loss of one, or more, of these
layers has not only given rise to the broad range of fish scales we see in today‟s extant
species, but also to vertebrate teeth and many of the dermal (intramembranous) bones of
vertebrates!

Chondrichthyans (sharks & rays) have placoid (plate) scales that lack vascular and
lamellar bone, but retain the enamel and dentine layers around a pulp cavity, forming
projecting scales also known as surface denticles (“small teeth”) (Figure 1). The placoid
scale develops in the dermis but projects through the epidermis to reach the surface. The
teeth in sharks, as well as those of higher vertebrates, are modifications of placoid scales,
as are the spines and soft rays of fins, and the superficial bones of the skull.

In bony fishes, three types of scale are recognised but most species have teleost scales.
These scales lack enamel, dentine and vascular bone, leaving only the lamellar bone
which develops in the dermis, but does not penetrate the epidermis (Figure 2). Teleosts
scales may be classed as cycloid or ctenoid. Both are composed of concentric rings
(circuli), but ctenoid scales also have a series of small projections on the posterior margin.
As a fish grows, new circuli are added to the scales reflecting annual or reproductive
cycles (much like the growth rings in a tree).

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Figure 1. Shark Skin. a) Surface view showing regular arrangement of projecting placoid scales.
b) Section through a placoid scale of a shark. Note: Diagram on the left shows a high power
magnification of a section through the adjacent epidermis. From Kardong, p 213, 2009.

Figure 2. Skin of Bony Fishes. From Hildebrand & Goslow p 86, 2001.

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Figure 3. Teleost Scales. Note the concentric circuli and the teeth-like projections on the
posterior margin of the ctenoid scale. From Kent & Carr, p 124, 2001.

Skeleton

Skull

The skull of a typical bony fish can be divided into two parts:

(a) Neurocranium - consists of bones that surround the olfactory, optic and otic parts of the
brain, overlain by dermal bones which give form to the face.

(b) Branchiocranium - consists of bones of the jaws, hyoid (jaw support) and operculum,
and the gill arches.

Axial Skeleton

The vertebrae are divided into two regions: trunk and caudal. Each vertebra has a
biconcave centrum that develops around the embryonic notochord. Dorsal to each centrum
is a neural arch surrounding the neural canal in which the spinal cord lies. Caudal
vertebrae, found in the tail, have an additional haemal arch which extends ventrally and
protects the major blood vessels.

In many species of fishes there are both dorsal and ventral sets of ribs attached to each
vertebra. The ribs are located within the connective tissue (myosepta) that separates each
successive muscle block. The ventral ribs are serially homologous with the haemal arches
of the caudal vertebrae. Many fishes have additional small splint bones in the myosepta
around muscle blocks.

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Figure 4. Fish Vertebrae. From Kardong, p 291, 2009.

Appendicular Skeleton & Fins

The pectoral fins are supported by a pectoral girdle which is attached dorsally to the skull.
The pelvic fins are supported by the pelvic girdle which is also in articulation with the
pectoral girdle.

Figure 5. Fish Skeleton.

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Fins can be divided into two main types:

(a) Median – positioned along the median axis: dorsal, caudal & anal fins; there may also
be a fleshy adipose fin posterior to the dorsal fin.

(b) Paired - consisting of pectoral and pelvic fins.

The fins are supported by fin rays, which may be either:

(a) Spines - normally hard, stout rays formed from a single median element and generally
present at the cranial end of the dorsal fin (see „a‟ in Figure 5); or

(b) Soft rays - which have a bilateral structure and generally show some segmentation
and branching (see „b‟ in Figure 5).

The internal skeletal support of the dorsal and anal fins is a series of bones that make up
the pterygiophores (Figure 6). Pterygiophores consist of three basal bones and several
radial bones, which lie in the median septum between adjacent vertebral spines and
articulate with the fin rays.

The caudal fin is supported by a terminal urostyle, which is formed by the fusion of several
caudal vertebrae.

Spines of fin ray

Dorsal fins
Adipose fin
Pterygiophore Caudal fin

Urostyle

Anal fin
Pectoral fin
Pelvic fin

Figure 6. Fish Fins.

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Musculature

Fish, like other vertebrates, have three types of muscles: smooth, cardiac and skeletal.
The musculature of fishes is far less complex than that of higher vertebrates.
Behind the head, the skeletal musculature is uniformly segmented with connective tissue
myosepta dividing the muscle blocks into myomeres. The myomeres of bony fish are
typically 'W'-shaped and form a series of open cones with the apices directed cranially,
overlapping considerably (Figure 7). There is also a horizontal septum that divides the
musculature into epaxial and hypaxial muscle masses (Figure 7).

Derivatives of the segmental muscles supply the paired fins and muscles of the head.
The major muscles of the head are those associated with closing the jaws and moving
the operculum that covers the gills. Muscles of the median fins (dorsal and anal) serve to
erect the fin (protractor muscles) or to depress it (retractor muscles). Similarly, the caudal
fin has dorsal and ventral flexors to raise or lower it, respectively. The paired pectoral
and pelvic fins have abductor muscles to spread or elevate them and adductor muscles
to do just that: adduct them (bring them closer to the body).

Myomeres

Figure 7. Musculature of Fish

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Locomotion

The majority of the propulsive forces for swimming are generated by the axial
musculature. Swimming is achieved by alternate contractions of the myomeres, first on
one side of the body and then the other, starting at the head. Since the vertebral column
prevents any contraction of the body, it is thrown into a series of lateral undulations, that
generate lateral thrusts against the water and so propel the fish forwards. The head acts
as a fulcrum for the tail, with the greatest propulsive forces being generated by the tail.

Figure 8. Locomotion in the Fish.

The caudal fin is clearly most important for locomotion but most of the other fins are also
able to be used for propulsion. However, their major role is in providing stability and
manoeuvrability. The dorsal and anal fins act as keels to prevent rolling and so are
situated caudal to the centre of gravity.

The pelvic and pectoral fins also act as keels to prevent rolling and pitching, but their
principal function seems to be for activities such as ascending, diving, turning and
stopping.

Digestive System

The basic pattern of the vertebrate digestive tract was established among the fishes.
Fishes show a wide variety of adaptations to different diets: cyclostomes (lampreys and
hagfishes) feed on small particulate matter including blood, detritus and particles of
tissue rasped from their prey; accordingly, they have a very simple digestive tract
consisting of a ciliated oesophagus that expands into a straight intestine, without the
presence of a stomach. Their mouth is situated within a buccal funnel which forms the
suction cup-like mechanism that allows them to „vacuum‟ for food or, as in the case of
lampreys, to attach to other fishes. The tongue of lampreys is specialised with abrasive
„teeth‟ for rasping at the skin or flesh of prey to obtain blood and tissue debris.

In the gnathostomes (jawed fish), there is generally an oesophagus, stomach and

intestine with modifications that reflect differing diets. For example, predatory fish have
well-developed teeth, a large stomach that secretes digestive acids and a relatively short
intestine (protein is easily digested and absorbed relative to plant material).

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Oropharangeal Cavity

Teeth are thought to have evolved from the bony armour of primitive fishes. In sharks,
teeth have clearly arisen from placoid scales.

In bony fish, teeth may be found on;

(a) the jaw margins.

(b) roof of the mouth.
(c) tongue.
(d) pharynx (gill arches)

Fish are primarily homodont (all teeth the same). The teeth are clearly associated with
the type of diet. Skates and rays have grinding molariform teeth in their oral and
pharyngeal cavities and feed on clams, other molluscs and crustacea. Grazers such as
parrot fish bite their food off with a “beak” resulting from the fusion of individual teeth at
the front of the jaw. This is followed by titurition by pharyngeal grinding teeth.

In sharks, teeth are continually replaced and are not set into the jaw bone.

Alimentary Canal

Fish generally have a very distensible oesophagus which allows them to swallow food in
large units. The stomach of predatory fish is elongate while that of omnivorous species is
generally sac-shaped. The stomach of the mullet is extensively muscularised to form a
powerful grinding organ (akin to the gizzard in birds); the lining is also heavily
strengthened with connective tissue. Remember that lampreys and hagfishes subsist on
a very simple diet of particulate matter and so lack a stomach altogether.

The stomachs of predatory fish contain gastric glands which secrete HCl and
pepsinogen, important in the digestion of proteins. An interesting modification of the
stomach occurs in puffer-fish and porcupine fish which enables the stomach to become
inflated with air or water – this distends the fish so that it appears almost globular in
shape. This probably has a protective function because when the fish is in this distended
shape it serves to erect spines all over the body.

The intestine also shows interspecies variation depending on food type. Typically, it is
short in carnivores and elongated, with many folds, in herbivores (plant material takes
longer to digest). The intestine of sharks contains an elaborate spiral fold known as the
spiral valve which acts to slow the passage of food, and to increase the surface area of
the intestine, all of which serve to facilitate digestion and absorption. Features with
similar functions in other species include transverse folds (rugae) and villi. Glands
throughout the intestine secrete digestive enzymes and mucous.

In most species of bony fishes, pyloric caecae are present at the junction of the stomach
and the duodenum. These can vary in number from one caecum to approximately 200
caecae in some species, and serve as additional areas for digestion and absorption.

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The liver produces bile salts which may be concentrated in a gall bladder before being
discharged into the intestine. Fish, like other vertebrates, have both an endocrine
(insulin) and exocrine pancreas (digestive enzymes). In lampreys and hagfishes, the
exocrine pancreas is dispersed across the liver and throughout the submucosal layer of
the intestine, while the endocrine pancreas forms a discrete clump of cells adjacent to
the bile duct. In sharks and some species of bony fish, the pancreas is a single structure
with clearly defined endocrine and exocrine components, while in other species of bony
fish it is a diffuse structure scattered throughout the omentum and liver, and so is termed
an hepatopancreas.

Figure 9. Fish Digestive Tract

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Figure 10. Digestive Tracts of Various Fishes. From Kardong, p 520, 2009.

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Circulatory System

Fish (with the exception of lungfish) have what is termed a single circulatory system: de-
oxygenated blood is returned from the body via veins to the heart; blood then leaves the
heart via arteries to the gills where it is oxygenated before continuing on to the rest of
body. Contrast this with the situation in higher vertebrates where oxygenated blood is
first returned from the pulmonary circulation to the heart, from where it is pumped to the
body.

The hearts of chondrichthyans and bony fish consist of four chambers: sinus venosus,
atrium, ventricle and bulbus arteriosus (conus arteriosus) (Figure 11). Blood returning to
the heart enters the thin-walled sinus venosus. This opens medially into the atrium
(auricle) which is also fairly thin-walled but has muscle fibres which can contract to pump
blood to the ventricle. The atrio-ventricular opening is surrounded by a sphincter and
pocket-shaped valves which prevent the return of blood to the atrium. The ventricle has
thick, muscular walls and pumps blood into the elastic bulbus ateriosus and the ventral
aorta. Note that while the sinus venosus, atrium and ventricle consist of cardiac muscle,
the bulbus arteriosus is composed of smooth muscle. The ventral aorta is median and
divides to form the aortic arches which send afferent branchial arteries to each gill arch
(Figure 11). From the gills, efferent branchials carry the oxygenated blood to the carotid
artery which supplies the head. Oxygenated blood is delivered to the rest of the body via
the dorsal aorta which branches segmentally.

In the tail, the dorsal aorta is termed the caudal aorta and passes through the haemal
arches of the vertebrae. Blood from the tail passes into a caudal vein, lying just below the
dorsal aorta, and also into the renal portal system. From the kidney, blood passes into a
posterior cardinal vein which joins the anterior cardinal vein above the heart.

The heart of the lungfish differs to that of other fish in that the atrium is partially divided
by an interatrial septum to form left and right atrial chambers (Figure 11, Figure 12).
Pulmonary veins deliver oxygenated blood from the lungs to either the sinus venosus or
the left atrial chamber (depending on species). Systemic deoxygenated blood is
delivered to the right atrial chamber. The ventricle is also partially divided by an
interventricular septum, and the bulbus arteriosus contains a spiral valve, both of which
also help to keep oxygenated and deoxygenated blood separate. As the oxygenated and
deoxygenated blood streams exit the bulbus arteriosus, they pass to different aortic
arches: some associated with gills, and others lacking gills. The oxygenated blood flows
to the aortic arches lacking gills and continues directly to the dorsal aorta, thus entering
the general circulation. Deoxygenated blood is passed through those aortic arches that
are associated with gills and so the blood becomes oxygenated before entering the
dorsal aorta. (If you are interested in this extraordinary adaptation in the lungfish, much
more detail can be found in Kardong, p 471-472).

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 Heart

Figure 11. Circulation in the Fish

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Figure 12. A Comparison of the Circulatory System in Lungfishes vs Other Fishes

Gills and Respiration

The gills are the chief organs of respiration, although a few groups of fish have
developed specialised modifications for air breathing (see below). The gills are supported
by skeletal structures called branchial bars (branchial arches, gill bars). Attached to the
branchial bars are the gill filaments which are, in turn, composed of many lamellae
(primary and secondary), which are covered with a very thin epithelium and contain a
vast network of capillaries. Due to their delicate nature, gills are susceptible to parasitic
and bacterial attack. Along the anterior margin of the gill arch are rigid gill rakers which
strain the water before it passes over the gills, removing potentially damaging particulate
matter. In some species, gill rakers are modified to function as food harvesting
structures.

In sharks and rays, the gills are divided into individual chambers by interbranchial septa
(Figure 13). In the bony fish, these septa are reduced such that the gills all lie within a

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common chamber protected by the operculum (Figure 13). The operculum plays an
important role in ventilation in bony fish, forming part of the dual pump mechanism. The
dual pump creates alternating negative and positive pressures that first draw water into
the buccal chamber and then, secondly, force it across the gills (Figure 14). During the
suction phase, when water is drawn into the buccal chamber, the opercular valve at the
opercular margin seals the internal gill chamber. The mouth is opened and water flows
into the buccal and gill chambers, forcing the buccal chamber to expand (Figure 14). The
mouth then closes, the pharyngeal floor is raised and the buccal chamber contracts,
opening the opercular valve and forcing water out. In sharks, gill flap valves (gill septa)
replace the role of the operculum seen in bony fishes.

Sharks and other fast swimmers (e.g. tuna or mackerel) also swim with their mouths
open, allowing water to pass constantly over the gills; this process is known as ram
ventilation and serves to increase water flow across the gills, and so improve ventilation,
during periods of activity. Many sharks do not have well-developed respiratory
movements and can only take in sufficient water when swimming using ram ventilation -
they suffocate when immobilised.

The arrangement of capillaries in the gill filaments is such that a counter current is
produced with the water. Water high in oxygen is taken into the buccal chamber and
driven across the gill filaments which contain capillaries carrying deoxygenated blood in
the opposite direction to that of the entering water. Because the water and blood are
flowing in opposite directions, gas exchange is optimised with some fish able to extract
as much as 85% of the oxygen from the water (Figure 15).

Figure 13. Gill Structure. From Hildebrand & Goslow, p 221, 2001.

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http://www.arlis.org/docs/vol1/52386062/v04_id131_con_gillfun.jpg

Figure 14. Fish Ventilation

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Figure 15. Countercurrent Exchange

Various fish species have developed adaptations for air breathing. An example is
mudskippers, which leave the water for brief periods – to scavenge for crustaceans or to
evade predators – and have a highly folded and well vascularised inner wall of the
operculum and gill chamber which enables them to extract oxygen from the air. The
lungs of lungfish (Order Dipnoi) appear to be derived from the gas bladder (see below).
However, it is not certain whether these „lungs‟ are homologous with the lungs of higher
vertebrates. In the Australian lungfish (Neoceratodus forteri), there is a single lung (the
other species have paired lungs) which is connected to the oesophagus via a trachea.
Thus, swallowed air passes from the oesophagus to the lung which is divided into many
small compartments (faveoli) that are highly vascularised for gas exchange.

Gas Bladder

The gas bladder is a characteristic of bony fishes (note that it is absent in agnathans and
elasmobranchs) and has various functions across the fish species:

(a) Respiration: In fishes where the gas bladder is joined to the alimentary canal
(physostomous gas bladder), it is used as a supplementary organ of respiration. Air is
swallowed and passed to the gas bladder, via the pneumatic duct, where oxygen is
absorbed into the blood – this is a similar situation to that seen in the lungfishes except
that these air-breathing fishes utilise a gas bladder for gas exchange rather than lungs.

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 (b) Sound Conduction: The gas bladder may also act as a sound conductor, or
resonator. In some fishes, extensions of the gas bladder make direct contact with the
inner ear, while in others a series of small bones (Weberian ossicles) connect the gas
bladder to the inner ear. Because the gas bladder is filled with gas (!) it can also act as a
resonator to enhance sound detection.

(c) Sound Production: The gas bladder may be used to generate sound by abruptly
releasing air from the bladder (akin to burping!), grinding teeth, or strumming the bladder
via specialised muscles which originate from the dorsal body wall and insert onto the
bladder making its walls vibrate. Sounds play an important role in breeding behaviour,
the defence of territories, and as warning sounds.

(d) Hydrostatic Control: One of the most universally important functions of the gas
bladder amongst the bony fishes is as a hydrostatic organ, regulating buoyancy. In fish in
which the gas bladder serves primarily as a hydrostatic organ, rather than a respiratory
organ, it is known as a swim bladder. Fishes which have a physostomous gas bladder
(see above) regulate air volume in the bladder by gulping or releasing air via the
pneumatic duct (Figure 16).

Fishes in which the pneumatic duct has been lost (physoclistous swim bladder) adjust
gas volume in the bladder entirely by exchanging gases with the blood. The transfer of
gas into the bladder occurs at a highly vascularised portion of the bladder wall called the
gas gland with its associated rete mirabile; gas is removed from the bladder at the oval, a
pocket at one end of the swim bladder in which gas diffuses from the bladder into the
surrounding capillaries (Figure 16).

Rete
Mirabile

Figure 16. Swim Bladders. From Kardong, p 423, 2009.

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Excretion

The key role of the kidneys in fish is in osmoregulation. While they also excrete some
nitrogenous waste, the gills are the main site for the removal of nitrogenous wastes from
the body in the form of ammonia.

In freshwater fish, body fluids are hyperosmotic compared to the surrounding water and
so there is an influx of water through the gills, oral membranes and intestine, and a loss
of salts to the environment. This is countered by the kidneys which produce a large
volume of very dilute urine to remove excess water from the body – freshwater fish
excrete approximately 10x the volume of urine produced by marine fish. The kidneys
have large, well-developed glomeruli to facilitate the production of dilute urine. The gills
and oral surfaces also provide a site for the active absorption of salts.

Marine fish, however, live in an environment in which they are hypo-osmotic and so they
tend to lose water and gain salts, putting them at risk of dehydration. In this case the
kidneys produce a very concentrated urine so as to maximise the excretion of excess
salts and to minimise the loss of water. In contrast to the kidney of freshwater fishes, that
of marine fishes lacks glomeruli and so produces very little fluid that would need to be
resorbed via the renal tubules. The kidneys of marine fishes also lack a distal tubule –
normally the distal tubule resorbs salt but also allows for water to be excreted, and so by
losing the distal tubule the potential for water loss is eliminated. Thus, the urine of marine
fishes is produced by the selective secretion of solutes into the renal tubules to produce
a very concentrated urine.

Sharks solve their osmotic problems rather differently by retaining high levels of urea in
the blood plasma (ureotelism) so that blood osmolarity approximates that of sea water;
hence, large losses of water to the environment do not occur. Excess salts are eliminated
via a special gland called the rectal gland which passes salts into the faeces.

Reproduction

Testes are internal, usually paired (except in cyclostomes) and elongate, and are
suspended by mesorchia below the swim bladder. Sperm leaving the testis may move
down the meonephric (archinephric) ducts of the kidney or, in the higher bony fish, a true
sperm duct may develop.

The ovaries are also internal, longitudinal and suspended from the dorsal body wall in a
similar manner to the testes. Most bony fishes have paired ovaries, while cyclostomes
have only one ovary. In many species of sharks, two ovaries initially develop but only one
(usually the left) persists and is functional. The size of the ovaries varies considerably
with the stage of sexual maturity. In most bony fishes, the eggs pass from the ovary
directly into an oviduct. However, in trout, salmon and spotted barramundi, the eggs
rupture directly into the body cavity and pass to the exterior through pores in the body
wall near the anus.

Fertilisation is usually external, in which case there are behavioural mechanisms to bring
the two sexes into close proximity. However, in some species, fertilisation is internal and
is associated with the development of special intromittant organs. In male sharks, the

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pelvic fins are specialised to form claspers which are inserted into the female‟s cloaca.
During copulation, sperm exits the male cloaca and enters a groove in the clasper where
it is then flushed into the female cloaca via water squirted from siphon sacs in the male‟s
body wall. Male top-minnows have elongated rays in their anal fin which forms a
gonopodium. During mating the gonopodium transfers a package of sperm
(spermatophore) to the cloaca of the female to facilitate internal fertilisation.

Once the eggs are laid, there may be some incubation behaviour by the adults or the
eggs may be left attached to vegetation or floating in the water. The spotted barramundi
incubates the eggs in its buccal cavity until they hatch. The male kurtus, a north
Australian fish, broods the eggs on his forehead until hatching. Fish which bear live
young may mature the eggs in the oviduct (e.g. sharks) or while still in the ovary (species
of bony fish).

Figure 17. Reproductive Strategies.

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The Nervous System

The fish brain is an enlargement of the anterior end of the spinal cord. The olfactory
(cerebral) lobes, optic lobes and cerebellum are particularly prominent. The sense of
smell is quite important to fish. Scents are detected through the olfactory epithelium of
the nasal sac, below the external nares, which has nerves running directly to the
olfactory bulbs of the brain.

Figure 18. Evolutionary Development of the Brain. From Kent & Carr, p 400, 2001.

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The lateral line system is present in many fish species. It consists of long grooves, called
lateral line canals, in the skin particularly in the region of the head and extending along
the sides of the body and tail. Within the lateral line canals are sensory organs called
neuromasts, which are associated with afferent nerve pathways. Neuromasts consist of
an area of sensory tissue made up of receptor cells (hair cells) which send hair-like
extensions into a gelatinous cupula. Neuromasts communicate with the environment
through pores in the epidermis. The main function of the lateral line system is to provide
information about the direction of travel and of localised disturbances in the water caused
by small currents or vibrations. Cave-dwelling or deep sea fishes that lack vision are able
to accurately navigate around obstacles via information relayed through the lateral line
system. In surface-feeding fish the lateral line system detects the oscillations of insects
on the water‟s surface. There is also some evidence to suggest that the lateral line
system can detect low frequency sounds; however, this is still controversial.

Figure 19. Lateral Line and Neuromast. From Hildebrand & Goslow, p351, 2001.

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Figure 20. Fish Brain and Neuromast.

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