Chapter 18
Chapter 18
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o Highly-efficient respiratory system allows fishes ancient seafloors for decaying organic matter,
to extract oxygen from an environment that holds annelids, molluscs, or other bottom-dwelling
little oxygen per unit volume; efficient locomotor invertebrates.
structures allows fishes to move through a buoyant, o Most modern fishes are predators and spend
but viscous medium; highly efficient sensory much of their lives searching for food. Some feed on
systems include typical vertebrate systems, but also invertebrate animals floating or swimming in the
a lateral-line system that detects low-pressure plankton or living in or on a substrate. Many feed on
waves and electroreception; and efficient other vertebrates.
reproductive mechanisms have the potential to o The kind of food that one fish eats at different times
produce overwhelming numbers of offspring. in its life varies. For example, as a larva, a fish may
feed on plankton; as an adult, it may switch to a
18.3 Evolutionary Pressures Locomotion larger prey, such as annelids or smaller fish.
o The streamlined shape of a fish and the mucoid o Fishes usually swallow their prey whole. Teeth
secretions that lubricate its body surface reduce capture and hold prey, and some fishes have teeth
friction between the fish and the water. that are modified for crushing the shell of molluscs or
o Water’s buoyant properties also contribute to the the exoskeleton of arthropods.
efficiency of the fish’s movement through the water, o To capture prey, fishes often use the suction
o Fishes move through the water using their fins and closing the opercula and rapidly opening the mouth
body wall to push against the incompressible creates, which develops a negative pressure that
surrounding water. sweeps water and prey inside the mouth.
o Muscle bundles of most fishes are arranged in a o Herring, paddlefishes, and whale sharks are filter
zigzag pattern. Because these muscles extend feeders.
posteriorly and anteriorly in a zigzag fashion, o long gill processes, called gill rakers, trap plankton
contractions of each muscle bundle can affect a while the fish is swimming through the water with its
relatively large portion of the body wall. mouth open.
o Very efficient, fast-swimming fishes, such as tuna o Carps feed on a variety of plants and small animals.
and mackerel, support body movements with a o Lampreys are external parasites for at least a
vertical caudal (tail) fin that is tall and forked. portion of their lives.
o The forked shape of the caudal fin reduces surface o The fish digestive tract is similar to that of other
area that could cause turbulence and interfere with vertebrates. An enlargement, called the stomach,
forward movement. stores large, often infrequent, meals. The small
intestine, however, is the primary site for enzyme
Nutrition and the Digestive System secretion and food digestion.
o The earliest fishes were probably filter feeders o Sharks and other elasmobranchs have a spiral
and scavengers that sifted through the mud of valve in their intestine, and bony fishes possess
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outpockets of the intestine, called the pyloric ceca, adequate levels of oxygen in their bloodstream, they
that increases absorptive and secretory surfaces. must pass large quantities of water across the gill
surfaces and extract the small amount of oxygen
Circulation and Gas Exchange present in the water.
o All vertebrates have a closed circulatory system o Most fishes have a muscular pumping mechanism.
in which the heart pumps blood, with red blood cells o Some elasmobranchs and open-ocean bony fishes,
containing the hemoglobin, through a series of such as the tuna, maintain water flow by holding
arteries, capillaries, and veins. their mouths open while swimming. This method is
o In fishes, blood flows from the venous system called ram ventilation.
through the thin-walled, muscular atrium. From the o Spiracles are modified pharyngeal slits that open
atrium, blood flows into a larger, more muscular just behind the eyes of elasmobranchs and are used
ventricle. as an alternate route for water entering the pharynx.
o The ventricle is the primary pumping structure. o Gas exchange across gill surfaces is efficient. Gill
o Anterior to the ventricle is the conus arteriosus, (visceral) arches support gills. Gill filaments
extend from each gill arch and include vascular folds
which connects the ventral aorta.
of epithelium, called pharyngeal lamellae.
o In teleosts, the conus arteriosus is replaced by an
o Branchial arteries carry blood to the gills and into
expansion of the ventral aorta called the bulbus
gill filaments. They break into capillary bed in
arteriosus.
pharyngeal lamellae.
o The pulmonary artery is a vessel to the lungs that o Gas exchange occurs as blood and water move
has developed as a branch off aortic arch VI in opposite directions of either side of the lamellar
lungfishes. epithelium. This countercurrent exchange
o The atrium and ventricle of the lungfish heart are mechanism provides very efficient gas exchange by
partially divided. These partial divisions help keep maintaining a concentration gradient between the
less oxygenated blood from the body separate from blood and the water over the entire length of the
the oxygenated blood from the lungs. capillary bed.
o A spiral valve in the conus arteriosus helps direct
blood from the right side of the heart to the Swim Bladders and Lungs
pulmonary artery and blood from the left side of the o The Indian climbing perch spend its life almost
heart to the remaining aortic arches. Thus, the entirely on land. These fishes, like most bony fishes,
lungfishes show a distinction between a pulmonary have gas chambers called pneumatic sacs.
circuit and a systematic circuit. o In nonteleosts fishes and some teleosts, a
pneumatic duct connects the pneumatic sacs to
Gas Exchange the esophagus or another part of the digestive tract.
o Fishes live in an environment that contains less
than 25% of the oxygen present in air. To maintain
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o Most zoologists believe that lungs are more primitive o Openings, called external nares, in the snouts of
than swim bladders. fishes lead to olfactory receptors.
o The eyes of fishes are similar in most aspects of
Buoyancy regulation structure to those in other vertebrates. They are
o Bone has a specific gravity twice that of water. lidless, however, and the lenses are round.
o You can float on water because of two large, air-filled o Receptors for equilibrium, balance, and hearing
organs called lungs. are in the inner ears of fishes, and their function is
o Fishes maintain their vertical position in a column of similar to other vertebrates.
water in one or more of four ways. o Semicircular canals detect rotational movement,
One way is to incorporate low-density and other sensory patches help with equilibrium and
compounds into their tissues. balance by detecting the direction of the
gravitational pull.
A second way is to use fins to provide lift. o Fishes lack the outer and/or middle ear, which
A third adaptation is the reduction of heavy conducts sound waves to the inner ear in other
tissues in fishes. vertebrates.
The fourth adaptation is the swim bladder. o Vibrations strike the fish, are amplified by the swim
o The pneumatic duct connects the swim bladders of bladder, and are sent through the ossicles
garpikes, sturgeons, and other primitive bony fishes (modifications of vertebrae) to the skull.
to the esophagus or another part of the digestive o Running along each side and branching over the
tract. head of most fishes is a lateral-line system. The
o Most teleosts have swim bladders that have a lost lateral-line system consists of sensory pits in the
functional connection to the digestive tract. The epidermis of the skin that connect to canals that run
blood secretes gases into the swim bladder using a just below the epidermis. In these pits are receptors
countercurrent exchange mechanism in a vascular that are stimulated by water moving against them.
network called the rete mirabile (“miraculous Lateral lines are used to detect either water
net”) currents, or a predator or a prey that may be causing
o Gases may be reabsorbed into the blood at the water movements, in the vicinity of the fish. Fishes
posterior end of the bladder, the ovale. may also detect low-frequency sounds with these
receptors.
Nervous and Sensory Functions
o The central nervous system of fishes, as in other ELECTRORECEPTION AND ELECTRIC FISHES
vertebrates, consists of a brain and a spinal cord. o All organisms produce weak electrical fields from the
Sensory receptors are widely distributed over the activities of nerves and muscles. Electroreception
body. (electrogeneration) is the detection of electrical
fields that the fish or another organism in the
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environment generates. It has been demonstrated in o As with all vertebrates, the excretory structures in
over 500 species of fishes in seven families of the kidneys are called nephrons. These filter
Chondrichthyes and Osteichthyes. bloodborne nitrogenous wastes, ions, water, and
o Spiny dogfish sharks, the common laboratory small organic compounds across a network of
specimen, locate prey by electroreception. capillaries called a glomerulus.
o Electroreceptors are located on the heads of sharks o Freshwater fishes live in an environment
and are called ampullary organs. containing few dissolved substances. To control
o An electric fish (Gymnarchus niloticus) lives in excess water build-up and ion loss, they never drink
freshwater systems of Africa. Their electrical sense is and only take in water when feeding. They also
an adaptation to living in murky freshwater habitats produce large quantities of very dilute urine. The
where eyes are of limited value. numerous nephrons of these fishes often have large
o The fishes best known for producing strong electrical glomeruli and relatively short tubule systems.
currents are the electric eel (a bony fish) and the o Marine fishes face the opposite problems. Their
electric ray (an elasmobranch). environment contains 3.5% ions, and their tissues
o The electric eel (Electrophorus) occurs in rivers of contain approximately 0.65% ions. They combat
the Amazon basin in South America. The organs for water loss and accumulation of excess ions by
producing electric currents are in the trunk of the drinking water and eliminating excess ions by
electric eel and can deliver shocks in excess of 500 excretion, defecation, and active transport across gill
volts. surfaces. Their nephrons often possess a small
o The electric ray (Narcine) has electric organs in its glomeruli and long tubule systems.
fins that are capable of producing pulses of 50 o Elasmobranchs have a unique osmoregulatory
amperes at about 50 volts. Shocks that these fishes mechanism. They convert some of their nitrogenous
produce are sufficiently strong to stun or kill prey, wastes into urea in the liver. This adaptation required
discourage large predators, and teach unwary the development of tolerance to high levels of urea,
humans a lesson that will never need to be repeated. because urea disrupts important enzyme systems in
the most tissues of most other animals. In spite of
Excretion and Osmoregulation this unique adaptation, they must still regulate the
o Fishes, like all animals, must maintain a proper ion concentrations in their tissues. They possess a
balance of electrolytes (ions) and water in their rectal gland that removes excess sodium chloride
tissues. This osmoregulation is a major function of from the blood and excretes it into the cloaca.
the kidney and gills of fishes. o A cloaca is a common opening for excretory,
o Kidneys are located near the midline of the body, digestive, and reproductive products.
just dorsal to the peritoneal membrane that lines o Diadramous fishes migrate between freshwater
the body cavity. and marine environments. Salmon (e.g,
Onchorhynchus) and marine lampreys
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(Petromyzon) migrate from the sea to freshwater to known brooders include the seahorses
spawn, and the freshwater eel (Anguilla) migrates (Hippocampus) and pipefishes (e.g., Sygnathus).
from freshwater to marine environments to spawn. o The male Brazilian catfish (Loricaria typhys)
o Diadramous fishes require gills capable of coping broods embryos in an enlarged lower lip.
with both uptake and secretion of ions. o Sunfishes and sticklebacks provide short-term
o Up to 90% of nitrogenous wastes are eliminated as care of post-hatching young. The Cichlidae engage
ammonia by diffusion across gill surfaces. The in long-term care.
remaining 10% of nitrogenous wastes is excreted as 18.3 Further Phylogenetic Considerations
urea, creatine, or creatinine. These wastes are o Two important series of evolutionary events occurred
produced in the liver and are excreted via the during the evolution of the bony fishes.
kidneys.
One of these was an evolutionary
explosion that began about 150 mya and
Reproduction and Development resulted in the vast diversity of teleosts living
o Mating may occur in large schools, and one today.
individual releasing eggs or sperm often releases The second series of events involves the
spawning pheromones that induce many other adults evolution of terrestrialism.
to spawn. o Most cladistics and anatomical evidence indicated
o The vast majority of fishes are oviparous meaning that the lungfish lineage gave rise to no other
that eggs develop outside the female from stored vertebrate taxa.
yolk. Some elasmobranchs are ovoviviparous, and o The Tetrapodomorpha is a group that includes the
their embryos develop in a modified oviduct of the osteolepiform sarcopterygians. Osteolepiforms
female. Other elasmobranchs, including the gray reef possessed several unique characteristics in common
sharks and hammerheads, are viviparous. with early amphibians. These included structures of
o In guppies (Lebistes), eggs are retained in the the jaw, teeth, vertebrae, and limbs, among others.
ovary, and fertilization and early development occur These sarcopterygians probably represent early
there. stages in the transition between fish and tetrapods.
o Some fishes have specialized structures that aid in o Another group of Tetrapodomorpha may still be
sperm transfer. Male elasmobranchs have closer to the tetrapod ancestor. In 2004, a new 375-
modified pelvic fins called claspers. During million-year-old fossil, Tiktaalik, was discovered
copulation, the male inserts a clasper into the in the Canadian arctic. This fish had fins, gills, and
cloaca of a female. scales, but it also possessed many more tetrapod
o Clusters of embryos may be brooded in special characters than other sarcopterygian fossils,
pouches attached to some part of the body, or they including a dorsoventrally compressed and widened
may be brooded in the mouth. Some of the best- skull and striking tetrapod forelimb skeletal
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homologies. It lacked the opercular supports and
dorsal and anal fins present in other sarcopterygians.
o Tiktaalik is the first sarcopterygian fossil that shows
evidence of a pectoral girdle and a freely movable
neck.
o The pectoral girdle of tetrapods attached the
forelegs to the vertebral column, and other fishes
lack a neck.
o Tiktaalik may not be the direct ancestors of
amphibians, but this “fishapod” certainly
demonstrates the evolutionary pre-adaptations that
allowed vertebrates to become terrestrial.
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