In the auditory system, the columella contributes to hearing in amphibians, reptiles and

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...

s. The columella form thin, bony structures in the interior of the

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...

and serve the purpose of transmitting sounds from the eardrum. It is an evolutionary

homolog In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of prima ...

of the

stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...

, one of the auditory ossicles in mammals. In many species, the extracolumella is a cartilaginous structure that grows in association with the columella. During development, the columella is derived from the dorsal end of the hyoid arch.

Evolution

The evolution of the columella is closely related to the evolution of the jaw joint. It is an ancestral homolog of the

, and is derived from the hyomandibular bone of fishes. As the columella is derived from the hyomandibula, many of its functional relationships remain the same. The columella resides in the air-filled tympanic cavity of the middle ear. The footplate, or proximal end of the columella, rests in the oval window. Sound is conducted through the oval window to the interior of the otic capsule. This motion ultimately stimulates sensory cells in the inner ear. EB1911 Reptiles - Evolution of the Ossicular Chain of the Ear

EB1911 Reptiles - Evolution of the Ossicular Chain of the Ear

In the transition of

tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...

from sea to land, the earliest appearance of functional columella appeared in

temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carbo ...

Extracolumella

Crocodilians Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...

evolved to lift the head and body off the ground, isolating the head from ground vibrations. Under selective pressure to detect airborne sound vibrations, the columella in crocodilians have become more slender and reduce their mass. The extracolumella, a cartilaginous outgrowth on the distal end of the columella, couples the columella to the tympanum to conduct sound from the exterior air. Birds and modern crocodilians have evolved a trifurcated columella, which forms a Y-shaped support structure on the surface of the

tympanic membrane In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear from the middle ear. Its function is to transmit sound from the a ...

. In birds, this is thought to increase the surface area of the columellar footplate, thus lowering the threshold of hearing and improving the detection of airborne sound waves.

Anatomy in amphibians

Frogs

frogs A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" '' Triadobatrachus'' is ...

, the extracolumella is simple and club-shaped.

Anatomy in reptiles

In reptiles, the columella function to transduce sound through the

middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles, which transfer the vibrations of the eardrum into waves in the ...

as part of the auditory pathway. The columella is relatively straight and moves in a piston-like motion in response to vibration. Due to the rigid bony structure, the columella primarily responds to low-frequency vibrations transmitted through the ground.

Crocodilians

In crocodilians, the columella arises from a proximal and a distal component which develop into the columella and extracolumella, respectively. It is typically trifurcated, with three finger-like projections supporting it against the tympanic membrane. The extracolumella remains cartilaginous while the columella ossifies during development. The connection between the columella and extracolumella remains flexible over the animal's lifetime.

Snakes

Snakes have lost a tympanic membrane, and hence a distal attachment for the columella. The columella is instead connected to the

quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms upper pa ...

of the

jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...

. Thus, snakes are able to detect and localize ground vibrations through the lower jaw, rather than the sides of the head.

Worm lizards

In Amphisbaenia, the extracolumella is particularly lengthened and firmly connects with a layer of skin over dentary bone of the lower jaw. This connection appears to facilitate detection of airborne vibrations in the facial area. The embedding in the skin often occurs at a specially enlarged labial scale. As a result, the amphisbaenian is able to detect substrate vibrations as it burrows through the ground while protecting the internal ear from damage. Amphisbaenians otherwise lack an external ear structure, likely due to selective pressure to protect the middle and inner ears from damage as the animal burrows.

Birds

In birds, the columella is anchored to the conical tympanic membrane at an acute angle, rather than a 90-degree angle relative to the plane of the tympanic membrane. This is thought to provide a lever advantage in conducting airborne sound from the distal to the proximal end of the columella.

Development in chickens

In chick embryos, the primordial columella arises from a

mesenchymal Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every ...

condensation. Chondrification of the columella occurs earlier than the extracolumella. During endochondral ossification, the columella ossifies from two origins of

periosteum The periosteum is a membrane that covers the outer surface of all bones, except at the articular surfaces (i.e. the parts within a joint space) of long bones. Endosteum lines the inner surface of the medullary cavity of all long bones. Structu ...

: the shaft and the footplate.

Homology in mammals

Within mammals and other synapsids the columella has evolved into the stapes, a homologous bone within the newly evolved inner ear. As the tympanic cavity evolved to reduce in size, the columella shortened in length. The stirrup-shaped articular processes of the columella inspired a new name for this auditory ossicle, the stapes. The auditory ossicles continue to function in conducting transmitting sound through the auditory pathway; however, they have lost their function in conducting low frequency ground vibrations. Later-arising reptiles with columella likely evolved stronger limbs and a more crawling posture, which removed the body from the ground and prevented the transmission of ground-conducted sounds. The skin over the ear evolved into the eardrum, which allowed for the detection of high-frequency airborne vibrations. In mammals, the newly specialized ossicles function to transduce and amplify these vibrations along the auditory pathway.

Artificial columella

In humans, artificially made columella may be produced as

autograft Autotransplantation is the transplantation of organs, tissues, or even particular proteins from one part of the body to another in the same person ('' auto-'' meaning "self" in Greek). The autologous tissue (also called autogenous, autogen ...

s from

cortical bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, and ...

. These prostheses are used as replacements for the stapes in ear surgery to correct for hearing problems (such as

cholesteatoma Cholesteatoma is a destructive and expanding growth consisting of keratinizing squamous epithelium in the middle ear and/or mastoid process. Cholesteatomas are not cancerous as the name may suggest, but can cause significant problems because of th ...

or re-perforation).

References

{{Tetrapod osteology, S. Reptile anatomy Bird anatomy Amphibian anatomy Auditory system Ossicles