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Toes are the digits (fingers) of the
foot The foot ( : feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg made ...
of a
tetrapod 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 therapsid ...
.
Animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
species such as
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s that walk on their toes are described as being ''
digitigrade In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...
''.
Human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, an ...
s, and other animals that walk on the soles of their feet, are described as being ''
plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
''; ''
unguligrade Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, c ...
'' animals are those that walk on
hooves The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rum ...
at the tips of their toes.


Structure

There are normally five toes present on each human foot. Each toe consists of three
phalanx bone The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
s, the proximal, middle, and distal, with the exception of the big toe ( la, hallux). For a minority of people, the little toe also is missing a middle bone. The hallux only contains two phalanx bones, the proximal and distal. The joints between each phalanx are the interphalangeal joints. The proximal phalanx bone of each toe articulates with the
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the medi ...
bone of the foot at the
metatarsophalangeal joint The metatarsophalangeal joints (MTP joints), also informally known as toe knuckles, are the joints between the metatarsal bones of the foot and the proximal bones (proximal phalanges) of the toes. They are condyloid joints, meaning that an ellipt ...
. Each toe is surrounded by skin, and present on all five toes is a
toenail A nail is a claw-like plate found at the tip of the fingers and toes on most primates. Nails correspond to the claws found in other animals. Fingernails and toenails are made of a tough protective protein called alpha-keratin, which is a polymer. ...
. The toes are, from medial to lateral: * the first toe, also known as the hallux ("big toe", "great toe", "thumb toe"), the innermost toe; * the second toe, ("Index toe", "pointer toe"), * the third toe, ("middle toe"); * the fourth toe, ("fore toe"); and (fourth toe) * the fifth toe, ("baby toe", "little toe", "pinky toe", "small toe"), the outermost toe.


Muscles

Toe movement is generally flexion and extension (movement toward the sole or the back of the foot, resp.) via muscular
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
s that attach to the toes on the anterior and superior surfaces of the phalanx bones. With the exception of the hallux, toe movement is generally governed by action of the
flexor digitorum brevis The flexor digitorum brevis is a muscle which lies in the middle of the sole of the foot, immediately above the central part of the plantar aponeurosis, with which it is firmly united. Its deep surface is separated from the lateral plantar ves ...
and
extensor digitorum brevis The extensor digitorum brevis muscle (sometimes EDB) is a muscle on the upper surface of the foot that helps extend digits 2 through 4. Structure The muscle originates from the forepart of the upper and lateral surface of the calcaneus (in fro ...
muscles. These attach to the sides of the bones, making it impossible to move individual toes independently. Muscles between the toes on their top and bottom also help to abduct and adduct the toes. The hallux and little toe have unique muscles: * The hallux is primarily flexed by the
flexor hallucis longus muscle The flexor hallucis longus muscle (FHL) is one of the three deep muscles of the posterior compartment of the leg that attaches to the plantar surface of the distal phalanx of the great toe. The other deep muscles are the flexor digitorum longus a ...
, located in the deep posterior of the lower leg, via the
flexor hallucis longus The flexor hallucis longus muscle (FHL) is one of the three deep muscles of the posterior compartment of the leg that attaches to the plantar surface of the distal phalanx of the great toe. The other deep muscles are the flexor digitorum longus a ...
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
. Additional flexion control is provided by the
flexor hallucis brevis A flexor is a muscle that flexes a joint. In anatomy, flexion (from the Latin verb ''flectere'', to bend) is a joint movement that decreases the angle between the bones that converge at the joint. For example, one’s elbow joint flexes when o ...
. It is extended by the
abductor hallucis muscle The abductor hallucis muscle is an intrinsic muscle of the foot. It participates in the abduction and flexion of the great toe. Structure The abductor hallucis muscle is located in the medial border of the foot and contributes to form the prom ...
and the
adductor hallucis muscle The Adductor hallucis (Adductor obliquus hallucis) arises by two heads—oblique and transverse and is responsible for adducting the big toe. It has two heads, both are innervated by the lateral plantar nerve. Structure Oblique head The ''oblique ...
. * The little toe has a separate set of control muscles and tendon attachments, the
flexor A flexor is a muscle that flexes a joint. In anatomy, flexion (from the Latin verb ''flectere'', to bend) is a joint movement that decreases the angle between the bones that converge at the joint. For example, one’s elbow joint flexes when one ...
and abductor digiti minimi. Numerous other
foot muscles The foot ( : feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg made ...
contribute to fine motor control of the foot. The connective tendons between the minor toes account for the inability to actuate individual toes.


Blood supply

The toes receive blood from the digital branches of the plantar metatarsal arteries and drain blood into the
dorsal venous arch The dorsal venous arch of the foot is a superficial vein that connects the small saphenous vein and the great saphenous vein. Anatomically, it is defined by where the dorsal veins of the first and fifth digit, respectively, meet the great saphen ...
of the foot.


Nerve supply

Sensation to the skin of the toes is provided by five nerves. The
superficial fibular nerve The superficial fibular nerve (also known as superficial peroneal nerve) innervates the fibularis longus and fibularis brevis muscles and the skin over the antero-lateral aspect of the leg along with the greater part of the dorsum of the foot (wit ...
supplies sensation to the top of the toes, except between the hallux and second toe, which is supplied by the
deep fibular nerve The deep fibular nerve (also known as deep peroneal nerve) begins at the bifurcation of the common fibular nerve between the fibula and upper part of the fibularis longus, passes infero-medially, deep to the extensor digitorum longus, to the ante ...
, and the outer surface of the fifth toe, supplied by the
sural nerve The sural nerve ''(L4-S1)'' is generally considered a pure cutaneous nerve of the posterolateral leg to the lateral ankle. The sural nerve originates from a combination of either the sural communicating branch and medial sural cutaneous nerve, o ...
. Sensation to the bottom of the toes is supplied by the
medial plantar nerve The medial plantar nerve (internal plantar nerve) is the larger of the two terminal divisions of the tibial nerve (medial and lateral plantar nerve), which accompanies the medial plantar artery. From its origin under the laciniate ligament it p ...
, which supplies sensation to the great toe and inner three-and-a-half toes, and the
lateral plantar nerve The lateral plantar nerve (external plantar nerve) is a branch of the tibial nerve, in turn a branch of the sciatic nerve and supplies the skin of the fifth toe and lateral half of the fourth, as well as most of the deep muscles, its distribution ...
, which supplies sensation to the little toe and half of the sensation of the fourth toe. In
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, an ...
s, the hallux is usually longer than the second toe, but in some individuals, it may not be the longest toe. There is an inherited trait in humans, where the dominant
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
causes a longer second toe ("
Morton's toe Morton's toe is the condition of having a first metatarsal which is short in relation to the second metatarsal (see diagram). It is a type of brachymetatarsia. This condition is the result of a premature closing of the first metatarsal's grow ...
" or "Greek foot") while the
homozygous Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. M ...
recessive
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
presents with the more common trait: a longer hallux. People with the rare genetic disease
fibrodysplasia ossificans progressiva Fibrodysplasia ossificans progressiva (; FOP; also called Münchmeyer disease and formerly called myositis ossificans progressiva or Stoneman disease) is an extremely rare connective tissue disease in which fibrous connective tissue such as ...
characteristically have a short hallux which appears to turn inward, or medially, in relation to the foot.


Variation

Humans usually have five toes on each foot. When more than five toes are present, this is known as
polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In human ...
. Other variants may include syndactyly or arachnodactyly. Forefoot shape, including toe shape, exhibits significant variation among people; these differences can be measured and have been statistically correlated with
ethnicity An ethnic group or an ethnicity is a grouping of people who identify with each other on the basis of shared attributes that distinguish them from other groups. Those attributes can include common sets of traditions, ancestry, language, history, ...
. Such deviations may affect comfort and fit for various shoe types. Research conducted for the U.S. Army indicated that larger feet may still have smaller arches, toe length, and toe-breadth.Freedman, A., Huntington, E.C., Davis, G.C., Magee, R.B., Milstead, V.M. and Kirkpatrick, C.M.. 1946. Foot Dimensions of Soldiers (Third Partial Report), Armored Medical Research Laboratory,
Fort Knox Fort Knox is a United States Army installation in Kentucky, south of Louisville and north of Elizabethtown. It is adjacent to the United States Bullion Depository, which is used to house a large portion of the United States' official gold re ...
, Kentucky.


Function

The human foot consists of multiple bones and soft tissues which support the weight of the upright human. Specifically, the toes assist the human while
walking Walking (also known as ambulation) is one of the main gaits of terrestrial locomotion among legged animals. Walking is typically slower than running and other gaits. Walking is defined by an 'inverted pendulum' gait in which the body vaults ov ...
,Janey Hughes, Peter Clark, & Leslie Klenerman. ''The Importance of the Toes in Walking.'' The Journal of Bone and Joint Surgery, Vol. 72-B, No. 2. March, 1990

/ref> providing balance, weight-bearing, and thrust during
gait Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency. ...
.


Clinical significance

A sprain or strain to the small interphalangeal joints of the toe is commonly called a stubbed toe. A sprain or strain where the toe joins to the foot is called
turf toe A metatarsophalangeal joint sprain is an injury to the connective tissue between the foot and a toe (at a metatarsophalangeal joint, one of the joints in the ball of the foot). When the big toe is involved, it is known as "turf toe". Causes Turf ...
. Long-term use of improperly sized shoes can cause misalignment of toes, as well as other orthopedic problems.
Morton's neuroma Morton's neuroma is a benign neuroma of an intermetatarsal plantar nerve, most commonly of the second and third intermetatarsal spaces (between the second/third and third/fourth metatarsal heads; the first is of the big toe), which results in t ...
commonly results in pain and numbness between the third and fourth toes of the sufferer, due to it affecting the nerve between the third and fourth metatarsal bones. The big toe is also the most common locus of
ingrown nail An ingrown nail, also known as onychocryptosis from el, ὄνυξ () 'nail' and () 'hidden', is a common form of nail disease. It is an often painful condition in which the nail grows so that it cuts into one or both sides of the paronychium ...
s, and its proximal phalanx joint is the most common locus for
gout Gout ( ) is a form of inflammatory arthritis characterized by recurrent attacks of a red, tender, hot and swollen joint, caused by deposition of monosodium urate monohydrate crystals. Pain typically comes on rapidly, reaching maximal intensi ...
attacks.


Deformity

Deformities of the foot include
hammer toe A hammer toe or contracted toe is a deformity of the muscles and ligaments of the proximal interphalangeal joint of the second, third, fourth, or fifth toe causing it to be bent, resembling a hammer. In the early stage a flexible hammertoe is ...
, trigger toe, and claw toe. Hammer toe can be described as an abnormal contraction or “buckling” of a toe. This is done by a partial or complete dislocation of one of the joints, which form the toes. Since the toes are deformed further, these may press against a shoe and cause pain. Deformities of the foot can also be caused by
rheumatoid arthritis Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and hands are involv ...
and
diabetes mellitus Diabetes, also known as diabetes mellitus, is a group of metabolic disorders characterized by a high blood sugar level (hyperglycemia) over a prolonged period of time. Symptoms often include frequent urination, increased thirst and increased ap ...
. Deformities may predispose to
ulcer An ulcer is a discontinuity or break in a bodily membrane that impedes normal function of the affected organ. According to Robbins's pathology, "ulcer is the breach of the continuity of skin, epithelium or mucous membrane caused by sloughing o ...
s and pain, especially when shoe-wearing. A common problem involving the big toe is the formation of
bunion A bunion, also known as hallux valgus, is a deformity of the joint connecting the big toe to the foot. The big toe often bends towards the other toes and the joint becomes red and painful. The onset of bunions is typically gradual. Complicatio ...
s. These are structural deformities of the bones and the
joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw ...
between the foot and big toe, and may be painful. Similar deformity involving the
fifth toe Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plant ...
is described as
tailor's bunion Tailor's bunion is a condition caused as a result of inflammation of the fifth metatarsal bone at the base of the little toe. It is usually characterized by inflammation, pain and redness of the little toe. Often a tailor's bunion is caused by a ...
or bunionette. In
polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In human ...
(which can also affect the
finger A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers (Pentadactyly). Chambers 1 ...
s) one or more extra toes are present.


In reconstruction

A favourable option for the reconstruction of missing adjacent
finger A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers (Pentadactyly). Chambers 1 ...
s/multiple digit amputations, i.e. such as a metacarpal hand reconstruction, is to have a combined second and third toe transplantation. Third and fourth toe transplantation to the hand in replacing lost fingers is also a viable option.


History


Etymology


The
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlers in the mid-5th c ...
term for ''toe'' is (plural ). This is a contraction of , and derives from Proto-Germanic (cognates: Old Norse , Old Frisian , Middle Dutch , Dutch (perhaps originally a plural), Old High German , German ), perhaps originally meaning 'fingers' as well (many
Indo-European language The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Dut ...
s use one word to mean both 'fingers' and 'toes', e.g. ''digit''), and thus from PIE root — 'to show'.


In classical Latin ,Hyrtl, J. (1880). ''Onomatologia Anatomica. Geschichte und Kritik der anatomischen Sprache der Gegenwart.'' Wien: Wilhelm Braumüller. K.K. Hof- und Universitätsbuchhändler. p. 248–249
online
at Biodiversity Library.
Triepel, H. (1908). Memorial on the anatomical nomenclature of the anatomical society. In A. Rose (Ed.), ''Medical Greek. Collection of papers on medical onomatology and a grammatical guide to learn modern Greek'' (pp. 176–93). New York: Peri Hellados publication office. ,Lewis, C.T. & Short, C. (1879). ''A Latin dictionary founded on Andrews' edition of Freund's Latin dictionary.'' Oxford: Clarendon Press. and , with genitive and , are used to refer to the big toe. The form (genitive, ) currently in use is however a
blend word In linguistics, a blend (sometimes called blend word, lexical blend, portmanteau or portmanteau word) is a word formed from parts of two or more other words. At least one of these parts is not a morph (the realization of a morpheme) but instead ...
of the aforementioned forms.Triepel, H. (1910). ''Die anatomischen Namen. Ihre Ableitung und Aussprache. Mit einem Anhang: Biographische Notizen.''(Dritte Auflage). Wiesbaden: Verlag J.F. Bergmann. Compare ''pollex'', the equivalent term for the
thumb The thumb is the first digit of the hand, next to the index finger. When a person is standing in the medical anatomical position (where the palm is facing to the front), the thumb is the outermost digit. The Medical Latin English noun for thum ...
.


Evolution

Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...
traces the standard
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
five-toed schema from
fish fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as s ...
s via
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arborea ...
ancestors.


Other animals

In birds with
anisodactyl In biology, dactyly is the arrangement of digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. It comes from the Greek word δακτυλος (''dáktylos'') = "finger". Sometimes the ending "-dactylia" is use ...
or
heterodactyl In biology, dactyly is the arrangement of digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. It comes from the Greek word δακτυλος (''dáktylos'') = "finger". Sometimes the ending "-dactylia" is use ...
feet, the hallux is opposed or directed backwards and allows for grasping and perching. While the thumb is often mentioned as one of the signature characteristics in humans, this manual digit remains partially primitive and is actually present in all primates. In humans, the most
derived Derive may refer to: *Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguation ...
digital feature is the hallux.


See also

*
Ainhum Ainhum (from Portuguese, pronounced ), also known as dactylolysis spontanea, is a painful constriction of the base of the fifth toe frequently followed by bilateral spontaneous autoamputation a few years later. Signs and symptoms The groove be ...
*
Finger A finger is a limb of the body and a type of digit, an organ of manipulation and sensation found in the hands of most of the Tetrapods, so also with humans and other primates. Most land vertebrates have five fingers (Pentadactyly). Chambers 1 ...
, for the
hand A hand is a prehensile, multi-fingered appendage located at the end of the forearm or forelimb of primates such as humans, chimpanzees, monkeys, and lemurs. A few other vertebrates such as the koala (which has two opposable thumbs on each " ...
digits *
Morton's toe Morton's toe is the condition of having a first metatarsal which is short in relation to the second metatarsal (see diagram). It is a type of brachymetatarsia. This condition is the result of a premature closing of the first metatarsal's grow ...
*
Nail (anatomy) A nail is a claw-like plate found at the tip of the fingers and toes on most primates. Nails correspond to the claws found in other animals. Fingernails and toenails are made of a tough protective protein called alpha-keratin, which is a polymer. ...
*
Polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In human ...
and Syndactyly * Runner's toe * Toe cleavage *
Toe tag A toe tag is a piece of cardboard attached with string to the big toe of a deceased individual in a morgue. It is used for identification purposes, allowing the mortician, coroner, law enforcement, and others involved in the death process to co ...
* Toecuffs *
Webbed toes Webbed toes is the informal and common name for syndactyly affecting the feet—the fusion of two or more digits of the feet. This is normal in many birds, such as ducks; amphibians, such as frogs; and some mammals, such as kangaroos. In humans i ...


References

{{Authority control Foot Podiatry *