In humans, vocal cords, also known as vocal folds or voice reeds, are folds of throat tissues that are key in creating sounds through vocalization. The size of vocal cords affects the pitch of voice. Open when breathing and vibrating for
speech
Speech is a human vocal communication using language. Each language uses Phonetics, phonetic combinations of vowel and consonant sounds that form the sound of its words (that is, all English words sound different from all French words, even if ...
or
singing
Singing is the act of creating musical sounds with the voice. A person who sings is called a singer, artist or vocalist (in jazz and/or popular music). Singers perform music (arias, recitatives, songs, etc.) that can be sung with or without ...
, the folds are controlled via the
recurrent laryngeal branch of the
vagus nerve
The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and right ...
. They are composed of twin infoldings of
mucous membrane
A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
stretched horizontally, from back to front, across the
larynx
The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
. They
vibrate
Vibration is a mechanical phenomenon whereby oscillations occur about an equilibrium point. The word comes from Latin ''vibrationem'' ("shaking, brandishing"). The oscillations may be periodic function, periodic, such as the motion of a pendulum ...
, modulating the flow of air being expelled from the lungs during
phonation
The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defini ...
.
The 'true vocal cords' are distinguished from the 'false vocal folds', known as
vestibular fold
The vestibular fold (ventricular fold, superior or false vocal cord) is one of two thick folds of mucous membrane, each enclosing a narrow band of fibrous tissue, the vestibular ligament, which is attached in front to the angle of the thyroid car ...
s or ''ventricular folds'', which sit slightly superior to the more delicate true folds. These have a minimal role in normal
phonation
The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defini ...
, but can produce deep sonorous tones, screams and growls.
The length of the vocal fold at birth is approximately six to eight millimeters and grows to its adult length of eight to sixteen millimeters by adolescence.
Testosterone
Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testes and prostate, as well as promoting secondar ...
, an
androgen
An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This inc ...
secreted by the gonads, causes irreversible changes in the cartilages and musculature of the larynx when present in high enough concentrations, such as during an adolescent boy's
puberty
Puberty is the process of physical changes through which a child's body matures into an adult body capable of sexual reproduction. It is initiated by hormonal signals from the brain to the gonads: the ovaries in a girl, the testes in a boy. ...
: The thyroid prominence appears, the vocal folds lengthen and become rounded, and the epithelium thickens with the formation of three distinct layers in the
lamina propria
The lamina propria is a thin layer of connective tissue that forms part of the moist linings known as mucous membranes or mucosae, which line various tubes in the body, such as the respiratory tract, the gastrointestinal tract, and the urogenita ...
.
Structure
Location
The vocal folds are located within the larynx at the top of the
trachea
The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
. They are attached at the back to the
arytenoid cartilage
The arytenoid cartilages () are a pair of small three-sided pyramids which form part of the larynx. They are the site of attachment of the vocal cords. Each is pyramidal or ladle-shaped and has three surfaces, a base, and an apex. The arytenoid ...
s, and at the front to the
thyroid cartilage
The thyroid cartilage is the largest of the nine cartilages that make up the ''laryngeal skeleton'', the cartilage structure in and around the trachea that contains the larynx. It does not completely encircle the larynx (only the cricoid cartilage ...
. They are part of the
glottis
The glottis is the opening between the vocal folds (the rima glottidis). The glottis is crucial in producing vowels and voiced consonants.
Etymology
From Ancient Greek ''γλωττίς'' (glōttís), derived from ''γλῶττα'' (glôtta), va ...
. Their outer edges are attached to muscle in the larynx while their inner edges form an opening called the
rima glottidis
The rima glottidis is the opening between the two true vocal cords anteriorly, and the two arytenoid cartilages posteriorly. It is part of the larynx.
Anatomy
The rima glottidis is the narrowest part of larynx. It is longer (~23 mm) in males ...
. They are constructed from
epithelium
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellul ...
, but they have a few muscle-fibres in them, namely the
vocalis muscle
The thyroarytenoid muscle is a broad, thin muscle that forms the body of the vocal fold and that supports the wall of the ventricle and its appendix. It functions to shorten the vocal folds.
Structure
It arises in front from the lower half of the ...
which tightens the front part of the
ligament
A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the:
* Peritoneal li ...
near to the thyroid cartilage. They are flat triangular bands and are pearly white in color. Above both sides of the
glottis
The glottis is the opening between the vocal folds (the rima glottidis). The glottis is crucial in producing vowels and voiced consonants.
Etymology
From Ancient Greek ''γλωττίς'' (glōttís), derived from ''γλῶττα'' (glôtta), va ...
are the two
vestibular fold
The vestibular fold (ventricular fold, superior or false vocal cord) is one of two thick folds of mucous membrane, each enclosing a narrow band of fibrous tissue, the vestibular ligament, which is attached in front to the angle of the thyroid car ...
s or false vocal folds which have a small sac between them.
False vocal folds
The vocal folds are sometimes called 'true vocal folds' to distinguish them from the 'false vocal folds' known as
vestibular fold
The vestibular fold (ventricular fold, superior or false vocal cord) is one of two thick folds of mucous membrane, each enclosing a narrow band of fibrous tissue, the vestibular ligament, which is attached in front to the angle of the thyroid car ...
s or ''ventricular folds''. These are a pair of thick folds of mucous membrane that protect and sit slightly higher to the more delicate true folds. They have a minimal role in normal
phonation
The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defini ...
, but are often used to produce deep sonorous tones in
Tibetan chant
The music of Tibet reflects the cultural heritage of the trans-Himalayan region centered in Tibet, but also known wherever ethnic Tibetan groups are found in Nepal, Bhutan, India and further abroad. The religious music of Tibet reflects the ...
and
Tuvan throat singing
Tuvan throat singing, the main technique of which is known as ''khoomei'' ( tyv, хөөмей, xöömej, mn, хөөмий; ᠬᠦᠭᠡᠮᠡᠢ, khöömii, russian: хоомей, Chinese: 呼麦, pinyin: ''hūmài''), includes a type of overt ...
, as well as in musical
screaming
A scream is a loud speech production, vocalization in which air is passed through the vocal cords with greater force than is used in regular or close-distance vocalisation. This can be performed by any creature possessing lungs, including human ...
and the
death growl
A death growl, or simply growl, is an extended vocal technique usually employed in extreme styles of music, particularly in death metal and other extreme subgenres of heavy metal music. Death growl vocals are sometimes criticized for their ...
vocal style.
Microanatomy
The vocal cords are composed of twin infoldings of 3 distinct tissues: an outer layer of flat cells that do not produce
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
(
squamous epithelium
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
). Below this is the superficial layer of the
lamina propria
The lamina propria is a thin layer of connective tissue that forms part of the moist linings known as mucous membranes or mucosae, which line various tubes in the body, such as the respiratory tract, the gastrointestinal tract, and the urogenita ...
, a gel-like layer, which allows the vocal fold to vibrate and produce sound. The
vocalis
The thyroarytenoid muscle is a broad, thin muscle that forms the body of the vocal fold and that supports the wall of the ventricle and its appendix. It functions to shorten the vocal folds.
Structure
It arises in front from the lower half of the ...
and
thyroarytenoid muscle
The thyroarytenoid muscle is a broad, thin muscle that forms the body of the vocal fold and that supports the wall of the ventricle and its appendix. It functions to shorten the vocal folds.
Structure
It arises in front from the lower half of the ...
s make up the deepest portion. These vocal folds are covered with a
mucous membrane
A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
and are stretched horizontally, from back to front, across the
larynx
The larynx (), commonly called the voice box, is an organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal inlet is about ...
.
Variation
Males and females have different vocal fold sizes. Adult male
voices are usually lower-pitched due to longer and thicker folds. The male's vocal folds are between 1.75 cm and 2.5 cm (approx 0.75" to 1.0") in length,
while females' vocal folds are between 1.25 cm and 1.75 cm (approx 0.5" to 0.75") in length. The vocal folds of children are much shorter than those of adult males and females. The difference in vocal fold length and thickness between males and females causes a difference in vocal pitch. Additionally, genetic factors cause variations between members of the same sex, with males' and females' voices being categorized into
voice types
A voice type is a group of voices with similar vocal ranges, capable of singing in a similar tessitura, and with similar vocal transition points ('' passaggi''). Voice classification is most strongly associated with European classical music, ...
.
Development
In newborns
Newborns have a uniform single layered lamina propria, which appears loose with no vocal ligament.
The monolayered lamina propria is composed of ground substances such as
hyaluronic acid
Hyaluronic acid (; abbreviated HA; conjugate base hyaluronate), also called hyaluronan, is an anionic, nonsulfated glycosaminoglycan distributed widely throughout connective, epithelial, and neural tissues. It is unique among glycosaminoglycans ...
and
fibronectin
Fibronectin is a high- molecular weight (~500-~600 kDa) glycoprotein of the extracellular matrix that binds to membrane-spanning receptor proteins called integrins. Fibronectin also binds to other extracellular matrix proteins such as collage ...
,
fibroblast
A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s, elastic fibers, and collagenous fibers. While the fibrous components are sparse, making the lamina propria structure loose, the hyaluronic acid (HA) content is high.
HA is a bulky, negatively charged glycosaminoglycan, whose strong affinity with water procures hyaluronic acid its viscoelastic and shock absorbing properties essential to vocal biomechanics.
Viscosity and elasticity are critical to voice production. Chan, Gray and Titze, quantified the effect of hyaluronic acid on both the viscosity and the elasticity of vocal folds by comparing the properties of tissues with and without HA.
The results showed that removal of hyaluronic acid decreased the stiffness of the vocal cords by an average of 35%, but increased their dynamic viscosity by an average of 70% at frequencies higher than 1 Hz. Newborns have been shown to cry an average of 6.7 hours per day during the first 3 months, with a sustained pitch of 400–600 Hz, and a mean duration per day of 2 hours.
Similar treatment on adult vocal cords would quickly result in edema, and subsequently aphonia. Schweinfurth and al. presented the hypothesis that high
hyaluronic acid
Hyaluronic acid (; abbreviated HA; conjugate base hyaluronate), also called hyaluronan, is an anionic, nonsulfated glycosaminoglycan distributed widely throughout connective, epithelial, and neural tissues. It is unique among glycosaminoglycans ...
content and distribution in newborn vocal cords is directly associated with newborn crying endurance.
[ These differences in newborn vocal fold composition would also be responsible for newborns inability to articulate sounds, besides the fact that their lamina propria is a uniform structure with no vocal ligament. The layered structure necessary for phonation will start to develop during the infancy and until the adolescence.]
The fibroblast
A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s in the newborn Reinke's space Reinke's space is a potential space between the vocal ligament and the overlying mucosa. It is not an empty space, but contains cells, special fibers and extracellular matrix. It plays an important role in the vibration of the vocal cords. Edema of ...
are immature, showing an oval shape, and a large nucleus-cytoplasm ratio. The rough endoplasmic reticulum and Golgi apparatus, as shown by electron micrographs, are not well developed, indicating that the cells are in a resting phase. The collagenous and reticular fibers in the newborn the vocal cords are fewer than in the adult one, adding to the immaturity of the vocal fold tissue.
In the infant, many fibrous components were seen to extend from the macula flava towards the Reinke's space. Fibronectin is very abundant in the Reinke's space of newborn and infant. Fibronectin is a glycoprotein that is believed to act as a template for the oriented deposition of the collagen fibers, stabilizing the collagen fibrils. Fibronectin also acts as a skeleton for the elastic tissue formation. Reticular and collagenous fibers were seen to run along the edges of the vocal cords throughout the entire lamina propria. Fibronectin in the Reinke's space appeared to guide those fibers and orient the fibril deposition. The elastic fibers remained sparse and immature during infancy, mostly made of microfibrils. The fibroblasts in the infant Reinke's space were still sparse but spindle-shaped. Their rough endoplasmic reticulum and Golgi apparatus were still not well developed, indicating that despite the change in shape, the fibroblasts still remained mostly in a resting phase. Few newly released materials were seen adjacent to the fibroblasts. The ground substance content in the infant Reinke's space Reinke's space is a potential space between the vocal ligament and the overlying mucosa. It is not an empty space, but contains cells, special fibers and extracellular matrix. It plays an important role in the vibration of the vocal cords. Edema of ...
seemed to decrease over time, as the fibrous component content increased, thus slowly changing the vocal fold structure.
Children
The infant lamina propria is composed of only one layer, as compared to three in the adult, and there is no vocal ligament. The vocal ligament begins to be present in children at about four years of age. Two layers appear in the lamina propria between the ages of six and twelve, and the mature lamina propria, with the superficial, intermediate and deep layers, is only present by the conclusion of adolescence. As vocal fold vibration is a foundation for vocal formants, this presence or absence of tissue layers influences a difference in the number of formants between the adult and pediatric populations. In females, the voice is three tones lower than the child's and has five to twelve formants, as opposed to the pediatric voice with three to six.
The length of the vocal fold at birth is approximately six to eight millimeters and grows to its adult length of eight to sixteen millimeters by adolescence. The infant vocal fold is half membranous or anterior glottis, and half cartilaginous or posterior glottis. The adult fold is approximately three-fifths membranous and two-fifths cartilaginous.
Puberty
Puberty
Puberty is the process of physical changes through which a child's body matures into an adult body capable of sexual reproduction. It is initiated by hormonal signals from the brain to the gonads: the ovaries in a girl, the testes in a boy. ...
usually lasts from 2–5 years, and typically occurs between the ages of 12 to 17. During puberty, voice change is controlled by sex hormones
Sex hormones, also known as sex steroids, gonadocorticoids and gonadal steroids, are steroid hormones that interact with vertebrate steroid hormone receptors. The sex hormones include the androgens, estrogens, and progestogens. Their effects are ...
. In females during puberty, the vocal muscle thickens slightly, but remains very supple and narrow. The squamous mucosa also differentiates into three distinct layers (the lamina propria) on the free edge of the vocal folds. The sub- and supraglottic glandular mucosa becomes hormone-dependent to estrogens and progesterone. For women, the actions of estrogen
Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
s and progesterone
Progesterone (P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the m ...
produce changes in the extravascular spaces by increasing capillary permeability which allows the passage of intracapillary fluids to the interstitial space as well as modification of glandular secretions. Estrogens have a hypertrophic and proliferative effect on mucosa by reducing the desquamating effect on the superficial layers. The thyroid hormone
File:Thyroid_system.svg, upright=1.5, The thyroid system of the thyroid hormones T3 and T4
rect 376 268 820 433 Thyroid-stimulating hormone
rect 411 200 849 266 Thyrotropin-releasing hormone
rect 297 168 502 200 Hypothalamus
rect 66 216 386 25 ...
s also affect dynamic function of the vocal folds; (Hashimoto's thyroiditis
Hashimoto's thyroiditis, also known as chronic lymphocytic thyroiditis and Hashimoto's disease, is an autoimmune disease in which the thyroid gland is gradually destroyed. Early on, symptoms may not be noticed. Over time, the thyroid may enlarg ...
affects the fluid balance in the vocal folds). Progesterone has an anti-proliferative effect on mucosa and accelerates desquamation. It causes a menstrual-like cycle in the vocal fold epithelium and a drying out of the mucosa with a reduction in secretions of the glandular epithelium. Progesterone has a diuretic effect and decreases capillary permeability, thus trapping the extracellular fluid
In cell biology, extracellular fluid (ECF) denotes all body fluid outside the cells of any multicellular organism. Total body water in healthy adults is about 60% (range 45 to 75%) of total body weight; women and the obese typically have a lower ...
out of the capillaries and causing tissue congestion.
Testosterone
Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testes and prostate, as well as promoting secondar ...
, an androgen secreted by the testes, will cause changes in the cartilages and musculature of the larynx for males during puberty. In women, androgens are secreted principally by the adrenal cortex
The adrenal cortex is the outer region and also the largest part of an adrenal gland. It is divided into three separate zones: zona glomerulosa, zona fasciculata and zona reticularis. Each zone is responsible for producing specific hormones. It is ...
and the ovaries and can have irreversible masculinizing effects if present in high enough concentration. In men, they are essential to male sexuality
Human male sexuality encompasses a wide variety of feelings and behaviors. Men's feelings of attraction may be caused by various physical and social traits of their potential partner. Men's sexual behavior can be affected by many factors, incl ...
. In muscles, they cause a hypertrophy of striated muscles with a reduction in the fat cells in skeletal muscle
Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
s, and a reduction in the whole body fatty mass. Androgens are the most important hormones responsible for the passage of the boy-child voice to man voice, and the change is irreversible. The thyroid prominence appears, the vocal folds lengthen and become rounded, and the epithelium thickens with the formation of three distinct layers in the lamina propria.
Adulthood
Human vocal cords are paired structures located in the larynx, just above the trachea, which vibrate and are brought in contact during phonation. The human vocal cords are roughly 12 – 24 mm in length, and 3–5 mm thick. Histologically, the human vocal cords are a laminated structure composed of five different layers. The vocalis muscle, main body of the vocal cords, is covered by the mucosa, which consists of the epithelium and the lamina propria.[Hirano, M., S. Kurita, and T. Nakashima. Vocal fold physiology : contemporary research and clinical issues. in Vocal Fold Physiology, Conference. 1981. San Diego, Calif.: College-Hill Press.] The latter is a pliable layer of connective tissue subdivided into three layers: the superficial layer (SL), the intermediate layer (IL), and the deep layer (DL). Layer distinction is either made looking at differential in cell content or extracellular matrix
In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide stru ...
(extracellular matrix) content. The most common way being to look at the extracellular matrix content. The SLP has fewer elastic and collagenous fibers than the two other layers, and thus is looser and more pliable. The ILP is mostly composed of elastic fibers, while the DLP has fewer elastic fibers, and more collagenous fibers. In those two layers, which form what is known as the vocalis ligament, the elastic and collagenous fibers are densely packed as bundles that run almost parallel to the edge of the vocal fold.
There is a steady increase in the elastin content of the lamina propria as humans age (elastin is a yellow scleroprotein, the essential constituent of the elastic connective tissue
Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
) resulting in a decrease in the ability of the lamina propria to expand caused by cross-branching of the elastin fibers. Among other things, this leads to the mature voice being better suited to the rigors of opera.
The extracellular matrix of the vocal cord LP is composed of fibrous proteins such as collagen and elastin, and interstitial molecules such as HA, a non-sulfated glycosaminoglycan
Glycosaminoglycans (GAGs) or mucopolysaccharides are long, linear polysaccharides consisting of repeating disaccharide units (i.e. two-sugar units). The repeating two-sugar unit consists of a uronic sugar and an amino sugar, except in the case ...
.[ While the SLP is rather poor in elastic and collagenous fibers, the ILP and DLP are mostly composed of it, with the concentration of elastic fibers decreasing and the concentration of collagenous fibers increasing as the vocalis muscle is approached.] Fibrous proteins and interstitial molecules play different roles within the extracellular matrix. While collagen (mostly type I) provides strength and structural support to the tissue, which are useful to withstanding stress and resisting deformation when subjected to a force, elastin fibers bring elasticity to the tissue, allowing it to return to its original shape after deformation.[ Interstitial proteins, such as HA, plays important biological and mechanical roles in the vocal cord tissue.][ In the vocal cord tissue, hyaluronic acid plays a role of shear-thinner, affecting the tissue viscosity, space-filler, shock absorber, as well as wound healing and cell migration promoter. The distribution of those proteins and interstitial molecules has been proven to be affected by both age and gender, and is maintained by the ]fibroblasts
A fibroblast is a type of biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework ( stroma) for animal tissues, and plays a critical role in wound healing. Fibroblasts are the most common cells o ...
.[
]
Maturation
Vocal fold structure in adults is quite different from that in newborns. Exactly how the vocal cord mature from an immature monolayer in newborns to a mature three layer tissue in adults is still unknown, however a few studies have investigated the subjects and brought some answers.
Hirano et al. previously found that the newborns did not have a true lamina propria, but instead had cellular regions called maculae flavae, located at the anterior and posterior ends of the loose vocal fold tissue. Boseley and Hartnick examined at the development and maturation of pediatric human vocal fold lamina propria. Hartnick was the first one to define each layer by a change in their cellular concentration. He also found that the lamina propria monolayer at birth and shortly thereafter was hypercellular, thus confirming Hirano's observations. By 2 months of age, the vocal fold started differentiating into a bilaminar structure of distinct cellular concentration, with the superficial layer being less densely populated than the deeper layer. By 11 months, a three-layered structure starts to be noted in some specimens, again with different cellular population densities. The superficial layer is still hypocellular, followed by an intermediate more hypercellular layer, and a deeper hypercellular layer, just above the vocalis muscle. Even though the vocal cords seem to start organizing, this is not representative of the trilaminar structure seen in adult tissues, where the layer are defined by their differential elastin and collagen fiber compositions. By 7 years of age, all specimens show a three-layered vocal fold structure, based on cellular population densities. At this point, the superficial layer was still hypocellular, the middle layer was the hypercellular one, with also a greater content of elastin and collagen fibers, and the deeper layer was less cellularly populated. Again, the distinction seen between the layers at this stage is not comparable to that seen in the adult tissue. The maturation of the vocal cords did not appear before 13 years of age, where the layers could be defined by their differential fiber composition rather than by their differential cellular population. The pattern now show a hypocellular superficial layer, followed by a middle layer composed predominantly of elastin fiber, and a deeper layer composed predominantly of collagen fibers. This pattern can be seen in older specimens up to 17 years of age, and above. While this study offers a nice way to see the evolution from immature to mature vocal cords, it still does not explain what is the mechanism behind it.
Macula flavae
Maculae flavae are located at the anterior and posterior ends of the membranous parts of the vocal cords. The histological structure of the macula flava is unique, and Sato and Hirano speculated that it could play an important role in growth, development and aging of the vocal cords. The macula flava is composed of fibroblasts
A fibroblast is a type of biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework ( stroma) for animal tissues, and plays a critical role in wound healing. Fibroblasts are the most common cells o ...
, ground substances, elastic and collagenous fibers. Fibroblasts were numerous and spindle or stellate-shaped. The fibroblasts have been observed to be in active phase, with some newly released amorphous materials present at their surface. From a biomechanical point of view, the role of the macula flava is very important. Hirano and Sato studies suggested that the macula flava is responsible for the synthesis of the fibrous components of the vocal cords. Fibroblasts have been found mostly aligned in the direction of the vocal ligament, along bundles of fibers. It then was suggested that the mechanical stresses during phonation were stimulating the fibroblasts to synthesize those fibers.
Impact of phonation
The viscoelastic
In materials science and continuum mechanics, viscoelasticity is the property of materials that exhibit both viscous and elastic characteristics when undergoing deformation. Viscous materials, like water, resist shear flow and strain linearly wi ...
properties of human vocal fold lamina propria are essential for their vibration, and depend on the composition and structure of their extracellular matrix
In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide stru ...
. Adult vocal cords have a layered structure which is based on the layers differential in extracellular matrix distribution. Newborns on the other hand, do not have this layered structure. Their vocal cords are uniform, and immature, making their viscoelastic properties most likely unsuitable for phonation. Hyaluronic acid plays a very important role in the vocal fold biomechanics. In fact, hyaluronic acid has been described as the extracellular matrix molecule that not only contributes to the maintenance of an optimal tissue viscosity that allows phonation, but also of an optimal tissue stiffness that allows frequency control.[ ]CD44
The CD44 antigen is a cell-surface glycoprotein involved in cell–cell interactions, cell adhesion and migration. In humans, the CD44 antigen is encoded by the ''CD44'' gene on chromosome 11. CD44 has been referred to as HCAM (homing cell adhes ...
is a cell surface receptor for HA. Cells such as fibroblast
A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s are responsible for synthesizing extracellular matrix molecules. Cell surface matrix receptors in return, feed back to the cells through cell-matrix interaction, allowing the cell to regulate its metabolism.
Sato et al. carried out a histopathologic investigation of unphonated human vocal cords. Vocal fold mucosae, which were unphonated since birth, of three young adults (17, 24, and 28 years old) were looked at using light and electron microscopy. The results show that the vocal fold mucosae were hypoplastic, and rudimentary, and like newborns, did not have any vocal ligament, Reinke's space, or layered structure. Like newborns, the lamina propria appeared as a uniform structure. Some stellate cell
Stellate cells are neurons in the central nervous system, named for their star-like shape formed by dendritic processes radiating from the cell body. Many stellate cells are GABAergic and are located in the molecular layer of the cerebellum. Ste ...
s were present in the macula flava, but started to show some signs of degeneration. The stellate cells synthesized fewer extracellular matrix molecules, and the cytoplasmic processes were shown to be short and shrinking, suggesting a decreased activity. Those results confirm the hypothesis that phonation stimulates stellate cells into producing more extracellular matrix.
Furthermore, using a specially designed bioreactor, Titze et al. showed that fibroblasts exposed to mechanical stimulation have differing levels of extracellular matrix production from fibroblasts that are not exposed to mechanical stimulation. The gene expression levels of extracellular matrix constituents such as fibronectin, MMP1, decorin, fibromodulin, hyaluronic acid synthase 2, and CD44
The CD44 antigen is a cell-surface glycoprotein involved in cell–cell interactions, cell adhesion and migration. In humans, the CD44 antigen is encoded by the ''CD44'' gene on chromosome 11. CD44 has been referred to as HCAM (homing cell adhes ...
were altered. All those genes are involved in extracellular matrix remodeling, thus suggesting that mechanical forces applied to the tissue, alter the expression levels of extracellular matrix related genes, which in turn allow the cells present in the tissue to regulate the extracellular matrix constituent synthesis, thus affecting the tissue's composition, structure, and biomechanical properties. In the end, cell-surface receptors close the loop by giving feedback on the surrounding extracellular matrix to the cells, affecting also their gene expression level.
Impact of hormones
Other studies suggest that hormones
A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and beh ...
play also an important role in vocal fold maturation. Hormones are molecules secreted into the blood stream to be delivered at different targeted sites. They usually promote growth, differentiation and functionality in different organs or tissues. Their effect is due to their ability to bind to intracellular receptors, modulating the gene expression, and subsequently regulating protein synthesis. The interaction between the endocrine system and tissues such as breast, brain, testicles, heart, bones, etc., is being extensively studied. It has clearly been seen that the larynx is somewhat affected by hormonal changes, but surprisingly, very few studies are working on elucidating this relationship. The effect of hormonal changes in voice is clearly seen when hearing male and female voices, or when listening to a teenage voice changing during puberty. Actually, it is believed that the number of hormonal receptors in the pre-pubertal phase is higher than in any other age.[ Menstruation has also been seen to influence the voice. In fact, singers are encouraged by their instructors not to perform during their pre-menstrual period, because of a drop in their voice quality.][
Vocal fold phonatory functions are known to change from birth to old age. The most significant changes occur in development between birth and puberty, and in old age.] Hirano et al. previously described several structural changes associated with aging, in the vocal fold tissue. Some of those changes are: a shortening of the membranous vocal fold in males, a thickening of the vocal fold mucosa and cover in females, and a development of edema in the superficial lamina propria layer in both sexes. Hammond et al. observed that the hyaluronic acid content in the vocal fold lamina propria was significantly higher in males than in females.[ Although all those studies did show that there are clear structural and functional changes seen in the human vocal cords which are associated with gender and age, none really fully elucidated the underlying cause of those changes. In fact, only a few recent studies started to look at the presence and role of hormone receptors in the vocal cords. Newman et al. found that hormone receptors are indeed present in the vocal cords, and show a statistical distribution difference with respect to age and gender.][ They have identified the presence of ]androgen
An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This inc ...
, estrogen
Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
, and progesterone
Progesterone (P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the m ...
receptors in epithelial cell
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
s, granular cells and fibroblast
A fibroblast is a type of cell (biology), biological cell that synthesizes the extracellular matrix and collagen, produces the structural framework (Stroma (tissue), stroma) for animal Tissue (biology), tissues, and plays a critical role in wound ...
s of the vocal cords, suggesting that some of the structural changes seen in the vocal cords could be due to hormonal influences.[ In this specific study, androgen and progesterone receptors were found more commonly in males than in females. In others studies, it has been suggested that the estrogen/androgen ratio be partly responsible for the voice changes observed at menopause. As previously said, Hammond et al. showed than the hyaluronic acid content was higher in male than in female vocal cords. Bentley et al. demonstrated that sex skin swelling seen in monkey was due to an increase in hyaluronic acid content, which was in fact mediated by estrogen receptors in dermal fibroblasts. An increase in collagen biosynthesis mediated by the estrogen receptors of dermal fibroblasts was also observed. A connection between hormone levels, and ]extracellular matrix
In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide stru ...
distribution in the vocal cords depending on age and gender could be made. More particularly a connection between higher hormone levels and higher hyaluronic acid content in males could exist in the human vocal fold tissue. Although a relationship between hormone levels and extracellular matrix biosynthesis in vocal fold can be established, the details of this relationship, and the mechanisms of the influence has not been elucidated yet.
Old age
There is a thinning in the superficial layer of the lamina propria in old age. In aging, the vocal fold undergoes considerable sex-specific changes. In the female larynx, the vocal fold cover thickens with aging. The superficial layer of the lamina propria loses density as it becomes more edematous. The intermediate layer of the lamina propria tends to atrophy only in men. The deep layer of the lamina propria of the male vocal fold thickens because of increased collagen deposits. The vocalis muscle atrophies in both men and women. However, the majority of elderly patients with voice disorders have disease processes associated with aging rather than physiologic aging alone.
Function
Oscillation
The larynx is a major (but not the only) source of sound in speech
Speech is a human vocal communication using language. Each language uses Phonetics, phonetic combinations of vowel and consonant sounds that form the sound of its words (that is, all English words sound different from all French words, even if ...
, generating sound through the rhythmic opening and closing of the vocal folds. To oscillate, the vocal folds are brought near enough together such that air pressure builds up beneath the larynx. The folds are pushed apart by this increased subglottal pressure, with the inferior part of each fold leading the superior part. Such a wave-like motion causes a transfer of energy from the airflow to the fold tissues. Under the correct conditions, the energy transferred to the tissues is large enough to overcome losses by dissipation and the oscillation pattern will sustain itself. In essence, sound is generated in the larynx by chopping up a steady flow of air into little puffs of sound waves.
The perceived pitch of a person's voice is determined by a number of different factors, most importantly the fundamental frequency
The fundamental frequency, often referred to simply as the ''fundamental'', is defined as the lowest frequency of a periodic waveform. In music, the fundamental is the musical pitch of a note that is perceived as the lowest partial present. In ...
of the sound generated by the larynx. The fundamental frequency is influenced by the length, size, and tension of the vocal folds. This frequency averages about 125 Hz in an adult male, 210 Hz in adult females, and over 300 Hz in children. Depth-kymography Depth-kymography is the 3D display of the human vocal fold vibrations by measuring their vertical and horizontal movements simultaneously, using a specially designed 3D endoscope. This 3D imaging method and the scientific term "Depth-Kymography" wer ...
is an imaging method to visualize the complex horizontal and vertical movements of vocal folds.
The vocal folds generate a sound rich in harmonic
A harmonic is a wave with a frequency that is a positive integer multiple of the ''fundamental frequency'', the frequency of the original periodic signal, such as a sinusoidal wave. The original signal is also called the ''1st harmonic'', the ...
s. The harmonics are produced by collisions of the vocal folds with themselves, by recirculation of some of the air back through the trachea, or both. Some singers can isolate some of those harmonics in a way that is perceived as singing
Singing is the act of creating musical sounds with the voice. A person who sings is called a singer, artist or vocalist (in jazz and/or popular music). Singers perform music (arias, recitatives, songs, etc.) that can be sung with or without ...
in more than one pitch at the same time—a technique called overtone singing
Overtone singing – also known as overtone chanting, harmonic singing, polyphonic overtone singing, and diphonic singing – is a set of singing techniques in which the vocalist manipulates the resonances of the vocal tract, in order to arous ...
or throat singing such as in the tradition of Tuvan throat singing
Tuvan throat singing, the main technique of which is known as ''khoomei'' ( tyv, хөөмей, xöömej, mn, хөөмий; ᠬᠦᠭᠡᠮᠡᠢ, khöömii, russian: хоомей, Chinese: 呼麦, pinyin: ''hūmài''), includes a type of overt ...
.
Clinical significance
Lesions
The majority of vocal fold lesions primarily arise in the cover of the folds. Since the basal lamina secures the epithelium to the superficial layer of the lamina propria with anchoring fibers, this is a common site for injury. If a person has a phonotrauma or habitual vocal hyperfunction, also known as pressed phonation, the proteins in the basal lamina can shear, causing vocal fold injury, usually seen as nodules or polyps, which increase the mass and thickness of the cover. The squamous cell
Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
epithelium of the anterior glottis are also a frequent site of laryngeal cancer caused by smoking.
Reinke's edema
A voice pathology called Reinke's edema, swelling due to abnormal accumulation of fluid, occurs in the superficial lamina propria or Reinke's space. This causes the vocal fold mucosa to appear floppy with excessive movement of the cover that has been described as looking like a loose sock. The greater mass of the vocal folds due to increased fluid lowers the fundamental frequency
The fundamental frequency, often referred to simply as the ''fundamental'', is defined as the lowest frequency of a periodic waveform. In music, the fundamental is the musical pitch of a note that is perceived as the lowest partial present. In ...
during phonation.
Wound healing
Wound healing is a natural regeneration process of dermal and epidermal tissue involving a sequence of biochemical events. These events are complex and can be categorized into three stages: inflammation, proliferation and tissue remodeling. The study on vocal fold wound healing is not as extensive as that on animal models due to the limited availability of human vocal folds. Vocal fold injuries can have a number of causes including chronic overuse, chemical, thermal and mechanical trauma such as smoking, laryngeal cancer, and surgery. Other benign pathological phenomena like polyps, vocal fold nodule
Vocal cord nodules are bilaterally symmetrical benign white masses that form at the midpoint of the vocal folds. Although diagnosis involves a physical examination of the head and neck, as well as perceptual voice measures, visualization of the v ...
s and edema will also introduce disordered phonation.
Any injury to human vocal folds elicits a wound healing process characterized by disorganized collagen deposition and, eventually, formation of scar tissue. Verdolini and her group sought to detect and describe acute tissue response of injured rabbit vocal cord model. They quantified the expression of two biochemical markers: interleukin 1
The Interleukin-1 family (IL-1 family) is a group of 11 cytokines that plays a central role in the regulation of immune and inflammatory responses to infections or sterile insults.
Discovery
Discovery of these cytokines began with studies on t ...
and prostaglandin E2, which are associated with acute wound healing. They found the secretions of these inflammatory mediators were significantly elevated when collected from injured vocal cords versus normal vocal cords. This result was consistent with their previous study about the function of IL-1 and PGE-2 in wound healing. Investigation about the temporal and magnitude of inflammatory response in the vocal cords may benefit for elucidating subsequent pathological events in vocal fold wounding,[ which is good for clinician to develop therapeutic targets to minimize scar formation. In the proliferative phase of vocal cord wound healing, if the production of hyaluronic acid and collagen is not balanced, which means the hyaluronic acid level is lower than normal, the fibrosis of collagen cannot be regulated. Consequently, regenerative-type wound healing turns to be the formation of scar.][ Scarring may lead to the deformity of vocal fold edge, the disruption of lipopolysaccharides viscosity and stiffness. Patients suffering from vocal fold scar complain about increased phonatory effort, vocal fatigue, breathlessness, and ]dysphonia
A hoarse voice, also known as dysphonia or hoarseness, is when the voice involuntarily sounds breathy, raspy, or strained, or is softer in volume or lower in pitch. A hoarse voice, can be associated with a feeling of unease or scratchiness in the ...
.[ Vocal fold scar is one of the most challenging problems for otolaryngologists because it is hard to be diagnosed at germinal stage and the function necessity of vocal cords is delicate.
]
Terminology
The vocal folds are commonly referred to as ''vocal cords'', and less commonly as ''vocal flaps'' or ''vocal bands''. The term ''vocal cords'' was coined by the French anatomist Antoine Ferrein
Antoine Ferrein (October 25, 1693 – February 28, 1769) was a French anatomist who was a native of Frespech, which today is a commune in the arrondissement of Villeneuve-sur-Lot. He was a professor at the Collège Royal in Paris, and in 174 ...
in 1741. In his violin analogy of the human voice
The human voice consists of sound made by a human being using the vocal tract, including talking, singing, laughing, crying, screaming, shouting, humming or yelling. The human voice frequency is specifically a part of human sound production ...
, he postulated that the moving air acted like a bow on ''cordes vocales''. The alternative spelling in English is ''vocal chords'', possibly due to the musical connotations or to confusion with the geometrical definition of the word '' chord''. While both spellings have historical precedents, standard American spelling is ''cords''. According to the Oxford English Corpus
The Oxford English Corpus (OEC) is a text corpus of 21st-century English, used by the makers of the ''Oxford English Dictionary'' and by Oxford University Press' language research programme. It is the largest corpus of its kind, containing nearly ...
, a database of 21st-century texts that contains everything from academic journal articles to unedited writing and blog entries, contemporary writers opt for the nonstandard ''chords'' instead of ''cords'' 49% of the time. The ''cords'' spelling is also standard in the United Kingdom and Australia.
In phonetics
Phonetics is a branch of linguistics that studies how humans produce and perceive sounds, or in the case of sign languages, the equivalent aspects of sign. Linguists who specialize in studying the physical properties of speech are phoneticians. ...
, ''vocal folds'' is preferred over ''vocal cords'', on the grounds that it is more accurate and illustrative.
See also
* Adam's apple
* Electroglottograph
The electroglottograph, or EGG, (also referred to as a laryngograph) is a device used for the noninvasive measurement of the degree of contact between the vibrating vocal folds during voice production. Though it is difficult to verify the assum ...
* Vocal register
A vocal register is a range of tones in the human voice produced by a particular vibratory pattern of the vocal folds. These registers include modal voice (or normal voice), vocal fry, falsetto, and the whistle register.
Registers originate in ...
* Falsetto
''Falsetto'' (, ; Italian diminutive of , "false") is the vocal register occupying the frequency range just above the modal voice register and overlapping with it by approximately one octave.
It is produced by the vibration of the ligamentous ed ...
* Vocal fry
The human voice consists of sound made by a human being using the vocal tract, including talking, singing, laughing, crying, screaming, shouting, humming or yelling. The human voice frequency is specifically a part of human sound productio ...
* Vocal cord dysfunction
Vocal cord dysfunction (VCD) is a pathology affecting the vocal folds (commonly referred to as the vocal cords) characterized by full or partial vocal fold closure causing difficulty and distress during respiration, especially during inhalation.
...
* Vocology Vocology is the science and practice of vocal habilitation, or vocal training and therapy.Titze IR. (1996). What is vocology? Logopedics Phoniatrics Vocology, 21:5-6. Its concerns include the nature of speech and language pathology, the defects of ...
* Articulatory phonetics
The field of articulatory phonetics is a subfield of phonetics that studies articulation and ways that humans produce speech. Articulatory phoneticians explain how humans produce speech sounds via the interaction of different physiological struc ...
* Laryngospasm
Laryngospasm is an uncontrolled or involuntary muscular contraction (spasm) of the vocal folds. The condition typically lasts less than 60 seconds, but in cases partial blocking it may last 20 to 30 minutes and hinder inspiration, while exhala ...
Additional images
Image:Endolarynx vocal folds.jpg, Vocal folds.
Image:Gray954.png, Coronal section of larynx and upper part of trachea.
Image:Gray955.png, The entrance to the larynx, viewed from behind.
Image:Gray960.png, Muscles of the larynx, seen from above.
References
External links
National Center for Voice and Speech's official website
* Lewcock, Ronald, et al. "Acoustics: The Voice." In
Grove Music Online
' (by subscription)/ http://www.oxfordmusiconline.com/subscriber/article/grove/music/00134pg6
{{DEFAULTSORT:Vocal Folds
Human voice
Human head and neck
Opera terminology
Phonetics