VERTEBRATE EAR

(Dr. Girish Chandra)

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.
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

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 eustachean 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.