ORGANS OF HEARING AND TACTILE RESPONSES

VERTEBRATE EAR

The ears are the organs of hearing and balance. They have three parts, namely external,
middle and inner ears.

External ear - The fleshy part outside the head is called the pinna. It is made up of elastic
cartilage and skin. It is followed by the external auditory meatus. This passage is lined with hairs
and ceruminous glands. These glands produce cerumen or earwax. The hair and wax prevent
foreign objects from reaching the ear drum. The ear drum or tympanic membrane is an oval,
three layered structure. It separates outer and middle ears.

Middle ear - It is an air filled cavity. It contains three auditory ossicles called the malleus, incus
and stapes. The handle of malleus is in contact with the inner surface of the ear drum. The head
of the malleus is attached to the incus. While the stapes on one side is attached to the incus, its
other side fits into the oval window. The oval window leads to the inner ear.

Inner ear - This region has tunnels and chambers inside the temporal bone called the bony
labyrinth. The bony labyrinth
contains three regions called the
cochlea, vestibule and
semicircular canals. The oval
window found in between the
middle and inner ears
communicates with the vestibule
of the inner ear. The organs of
the inner ear perceive the sound.

Actual ear is a delicate

organ located in the temporal region of the skull. It is called membranous labyrinth, which
evolved in primitive vertebrates for maintenance of balance and posture. Hearing function of this
organ evolved for the first time in anurans when they started living in terrestrial environment,
where sound waves travel faster and to longer distances. A typical membranous labyrinth
consists of three canals and three sacs. The three canals are attached at right angle to each other
and to the sacs in anterior, posterior and horizontal positions. The three sacs which are also
collectively called vestibule are: Utriculus, sacculus and lagena, each one of which possesses a
depression inside with a sensory organ called macula that detects the position of the body in
relation to gravity, or static sense. Each canal ends in an ampulla that also carries a sense organ
called crista, which is responsible for detection of the movement of head in any direction, or
kinetic sense. Both these senses together maintain the balance and posture of the animal during
its activity. Endolymph fills the membranous labyrinth and is secreted and drained in meninges.
Perlilymph surrounds the membranous labyrinth for protection.

Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

 The structure of crista is like any neuromast organ of fishes. It consists of sensory cells
with hairs on the top and nerve fibres attached below and many supporting cells that nourish and
protect the sensory cells. Hairs of sensory cells are embedded in a gelatinous mass called cupula
terminalis in which some crystals are embedded just above the hair tips. As the animal moves
these crystals also move and touch the hairs of sensory cells, which produce nerve impulses that
take the kinetic sense to the brain. This is called dynamic equilibrium maintained by the
semicircular canals.
Macula is the sensory organ of sacs. Its sensory cells also have hairs on the top and nerve
at the base but the cells are of two types, namely, cylindrical cells and flask shaped cells which
bear stereocilia and kinocilia embedded in a gelatinous mass. A paper thin otolith made of
calcium carbonate and proteins floats over the cilia and its position can be disturbed by the
animal bending or tilting in any direction in relation to gravity, disturbing the cilia in the process
and generating a nerve impulse that informs the brain of static sense.

CYCLOSTOMES
Cyclostomes have degenerated membranous labyrinth due to their parasitic mode of life.
Petromyzon has only two semicircular canals, namely the anterior and posterior and only two
sacs, utriculus and sacculus, lagena being absent. As they have no paired fins, lampreys and
hagfishes do not seem to have three dimensional sense of things. In hagfishes, the anterior and
posterior canals also fuse together to make a ring like structure. Sacculus and utriculus also fuse
together to form a single sac.

FISHES
Fishes use membranous labyrinth for balance and posture only, since the middle and
external ears are absent but some fishes can perceive water borne sounds to some extent. For
example, in fishes of the family Ostariophysi four pieces of Weberian ossicles (scaphium,
claustrum, intercalarium and tripus) connect the swim bladder with membranous labyrinth
and transport sound vibrations for hearing.
Fish membranous labyrinth is well developed with three canals and three sacs, and crista
and macula well formed. In elasmobranchs a recessus utriculus is attached on the lower side of
utriculus. Macula of utriculus is sometime called pars neglecta in fishes.

AMPHIBIA
Urodela and apoda do not have external and middle ear and consequently inner ear is
meant for maintenance of balance and posture only. Hearing power evolved in anurans by the
modification of lagena which produces an organ of reception of sound waves, called basilar
papilla. On the surface, on either side of head, a tympanic membrane is stretched over a ring of
cartilage called annulus tympanicus, which is a modification of pterygoquadrate cartilage of the
first visceral arch. Sound vibrations received by this membrane are transmitted to basilar papilla
through a columella bone, which is a modification of hyomandibular cartilage of the hyoid
arch. Thus, in frog a mechanism has evolved to receive sound vibrations from outside and
transmit them to the sense organ of lagena, so that the nerve impulse can convey information
about the sound to brain.

REPTILES & BIRDS

Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

 In reptiles and birds lagena elongates and basilar papilla elaborated to form the organ of
corti, which is much more sensitive to perceive sound vibrations. Columella bone is called
stapes, which is further elongated by the addition of an extrastapaedial cartilage. For better
protection, tympanic membrane sinks into the temporal bone, producing an auditory canal,
through which sound vibrations travel from outside to the tympanic membrane. In crocodiles
lagena is coiled like cochlea of mammals. In reptiles both Eustachian tubes join together to
open into pharynx by a single opening.
In snakes there are no external and middle ear cavities and hence sound vibrations travel
through ground to the lower jaw and from there to quadrate bone to columella, which carries
them to lagena of inner ear for perception of hearing. Therefore, snakes cannot hear sounds
travelling through air.
Birds have essentially similar mechanism of hearing as do the reptiles, except that ear is
much more sensitive in birds due to elongation of lagena.

MAMMALS
Mammals possess the best developed hearing power among all vertebrates. Lagena
elongates to form a spirally coiled cochlea. In middle ear cavity, instead of one bone, mammals
have three ear ossicles, namely, incus ( a derivative of pterygoquadrate), malleus (a derivative of
articular or meckel’s cartilage) and stapes (derived from hyomandibular), which transmit sound
vibrations from tympanic membrane to the fenestra ovalis that leads to scala vestibuli of
cochlea. Mammals also have various sizes of ear pinna to collect
sound waves and direct them to the auditory canal.
Cochlea is a specialized and highly sensitive sense organ of
hearing. Its cross section reveals three long chambers, namely,
scala vestibuli, scala media and scala tympani, the middle
chamber is filled with endolymph while the other two are filled
with perilymph. The organ of corti is attached to the basilar
membrane and carries sensory hair cells and supporting cells and
the cochlear nerve at the base. A tectorial membrane floats in the
middle of scala media and touches the hairs of sensory cells when it
vibrates by the sound vibrations, resulting in the generation of a
nerve impulse that travels via the cochlear nerve to the brain. Base of cochlea is more sensitive
to high frequency vibrations while the apical portion can detect very faint vibrations. Sound
waves from stapes enter scala vestibuli through fenestra ovalis or oval window, then travel to
the entire length of cochlea and return back via scala tympani and escape to the middle ear cavity
through fenestra rotunda or round window.
Human ear is capable of hearing sound waves ranging from 20 to 20,000 hertz but
different mammals possess capacity to hear sounds of different frequencies. For instance,
whales, dolphins and porpoises can hear ultrasonic sounds of up to 150,000 hertz and elephants
can also hear infrasonic sounds of 10-14 hertz.

Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016

Dr.C.V.Narasimha murthy. Associate professor (Contract), VSUPGCenter,Kavali. M.Sc.Zoology, notes-2016