The phalanges (singular: ''phalanx'' ) are

digital Digital usually refers to something using discrete digits, often binary digits. Technology and computing Hardware *Digital electronics, electronic circuits which operate using digital signals **Digital camera, which captures and stores digital i ...

bone A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provid ...

s in 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 "h ...

s and

feet 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 most

vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...

. In

primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...

s, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as

long bones The long bones are those that are longer than they are wide. They are one of five types of bones: long, short, flat, irregular and sesamoid. Long bones, especially the femur and tibia, are subjected to most of the load during daily activities and ...

Structure

The phalanges are the bones that make up the fingers of the hand and the toes of the foot. There are 56 phalanges in the human body, with fourteen on each hand and foot. Three phalanges are present on each finger and toe, with the exception of 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 thumb ...

and large toe, which possess only two. The middle and far phalanges of the fifth toes are often fused together (symphalangism). The phalanges of the

are commonly known as the finger bones. The phalanges of the foot differ from the hand in that they are often shorter and more compressed, especially in the proximal phalanges, those closest to the torso. A phalanx is named according to whether it is

proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

, middle, or

distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

and its associated finger or toe. The proximal phalanges are those that are closest to the hand or foot. In the hand, the prominent, knobby ends of the phalanges are known as

knuckle The knuckles are the joints of the fingers. The word is cognate to similar words in other Germanic languages, such as the Dutch "knokkel" (knuckle) or German "Knöchel" (ankle), i.e., ''Knöchlein'', the diminutive of the German word for bone (' ...

s. The proximal phalanges join with the

metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ...

s of the hand or

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 med ...

s of the foot at the

metacarpophalangeal joint The metacarpophalangeal joints (MCP) are situated between the metacarpal bones and the proximal phalanges of the fingers. These joints are of the condyloid kind, formed by the reception of the rounded heads of the metacarpal bones into shallow cav ...

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 ...

. The intermediate phalanx is not only intermediate in location, but usually also in size. The thumb and large toe do not possess a middle phalanx. The distal phalanges are the bones at the tips of the fingers or toes. The proximal, intermediate, and distal phalanges articulate with one another through

interphalangeal joints of hand The interphalangeal joints of the hand are the hinge joints between the phalanges of the fingers that provide flexion towards the palm of the hand. There are two sets in each finger (except in the thumb, which has only one joint): * "proximal int ...

and

interphalangeal joints of the foot The interphalangeal joints of the foot are between the phalanx bones of the toes in the feet. Since the great toe only has two phalanx bones (proximal and distal phalanges), it only has one interphalangeal joint, which is often abbreviated as the ...

Bone anatomy

Each phalanx consists of a central part, called the ''body'', and two extremities. * The ''body'' is flat on either side, concave on the palmar surface, and convex on the dorsal surface. Its sides are marked with rough areas giving attachment to fibrous sheaths of flexor tendons. It tapers from above downwards. * The ''proximal extremities'' of the bones of the first row present oval, concave articular surfaces, broader from side to side than from front to back. The proximal extremity of each of the bones of the second and third rows presents a double concavity separated by a median ridge. * The ''distal extremities'' are smaller than the proximal, and each ends in two

condyles A condyle (;Entry "condyle"
in
The phalanx of the thumb has a pronounced insertion for the

flexor pollicis longus The flexor pollicis longus (; FPL, Latin ''flexor'', bender; ''pollicis'', of the thumb; ''longus'', long) is a muscle in the forearm and hand that flexes the thumb. It lies in the same plane as the flexor digitorum profundus. This muscle is uniqu ...

(asymmetric towards the radial side), an ungual fossa, and a pair of unequal ungual spines (the ulnar being more prominent). This asymmetry is necessary to ensure that the thumb pulp is always facing the pulps of the other digits, an osteological configuration which provides the maximum contact surface with held objects. In the foot, the distal phalanges are flat on their dorsal surface. It is largest proximally and tapers to the distal end. The proximal part of the phalanx presents a broad base for articulation with the middle phalanx, and an expanded distal extremity for the support of the nail and end of the toe. The phalanx ends in a crescent-shaped rough cap of bone

epiphysis The epiphysis () is the rounded end of a long bone, at its joint with adjacent bone(s). Between the epiphysis and diaphysis (the long midsection of the long bone) lies the metaphysis, including the epiphyseal plate (growth plate). At the join ...

— the apical tuft (or ungual tuberosity/process) which covers a larger portion of the phalanx on the volar side than on the dorsal side. Two lateral ungual spines project proximally from the apical tuft. Near the base of the shaft are two lateral tubercles. Between these a V-shaped ridge extending proximally serves for the insertion of the

. Another ridge at the base serves for the insertion of the extensor

aponeurosis An aponeurosis (; plural: ''aponeuroses'') is a type or a variant of the deep fascia, in the form of a sheet of pearly-white fibrous tissue that attaches sheet-like muscles needing a wide area of attachment. Their primary function is to join musc ...

. The flexor insertion is sided by two fossae — the ungual fossa distally and the proximopalmar fossa proximally.

Development

The number of phalanges in animals is often expressed as a "phalangeal formula" that indicates the numbers of phalanges in digits, beginning from the innermost medial or proximal. For example, humans have a 2-3-3-3-3 formula for the

, meaning that the thumb has two phalanges, whilst the other fingers each have three. In the distal phalanges of the hand the centres for the bodies appear at the distal extremities of the phalanges, instead of at the middle of the bodies, as in the other phalanges. Moreover, of all the bones of the hand, the distal phalanges are the first to ossify.

Function

The distal phalanges of

ungulate 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, cam ...

s carry and shape nails and

claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...

s and these in primates are referred to as the ''

ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...

phalanges''.

History of phalanges

Etymology

The term phalanx or phalanges refers to an ancient Greek army formation in which soldiers stand side by side, several rows deep, like an arrangement of fingers or toes.

In animals

Most land

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

s including humans have a 2-3-3-3-3 formula in both the

s (or

paw A paw is the soft foot-like part of a mammal, generally a quadruped, that has claws. Common characteristics The paw is characterised by thin, pigmented, keratinised, hairless epidermis covering subcutaneous collagenous and adipose tissue, ...

s) and

. Primitive

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...

s usually had the formula 2-3-4-4-5, and this pattern, with some modification, remained in many later reptiles and in the

mammal-like reptile Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term ''mammal-like reptile'' had been used, and pelycosaur was considered an order, but this is no ...

s. The phalangeal formula in the flippers of

cetaceans Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...

(marine mammals) varies widely due to hyperphalangy (the increase in number of

phalanx bones 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. ...

in the digits). In

humpback whales The humpback whale (''Megaptera novaeangliae'') is a species of baleen whale. It is a rorqual (a member of the family Balaenopteridae) and is the only species in the genus ''Megaptera''. Adults range in length from and weigh up to . The hump ...

, for example, the phalangeal formula is 0/2/7/7/3; in

pilot whales Pilot whales are cetaceans belonging to the genus ''Globicephala''. The two extant species are the long-finned pilot whale (''G. melas'') and the short-finned pilot whale (''G. macrorhynchus''). The two are not readily distinguishable at sea, ...

the formula is 1/10/7/2/1. In vertebrates, proximal phalanges have a similar placement in the corresponding limbs, be they

wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...

fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...

. In many species, they are the longest and thickest phalanx ("finger" bone). The middle phalanx also a corresponding place in their limbs, whether they be

hoof 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 rumin ...

. The distal phalanges are cone-shaped in most mammals, including most primates, but relatively wide and flat in humans.

Primates

The morphology of the distal phalanges of human thumbs closely reflects an adaptation for a refined precision grip with pad-to-pad contact. This has traditionally been associated with the advent of stone tool-making. However, the intrinsic hand proportions of

australopiths Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' (cladistically including the genera ''Homo'', '' Paranthropus'', and ''Kenyanthropus''), and it typically includ ...

and the resemblance between human hands and the short hands of

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

apes, suggest that human hand proportions are largely

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

(as found in ancestral species) — in contrast to the derived elongated hand pattern and poorly developed thumb musculature of other extant

hominoid Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister g ...

s. In

Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While th ...

s, the apical tufts were expanded and more robust than in modern and early upper Paleolithic humans. A proposal that Neanderthal distal phalanges was an adaptation to colder climate (than in Africa) is not supported by a recent comparison showing that in

hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...

s, cold-adapted populations possessed smaller apical tufts than do warm-adapted populations. In non-human, living primates the apical tufts vary in size, but they are never larger than in humans. Enlarged apical tufts, to the extent they actually reflect expanded digital pulps, may have played a significant role in enhancing friction between the hand and held objects during Neolithic toolmaking. Among non-human primates

phylogenesis Phylogenesis (from Greek φῦλον ''phylon'' "tribe" + γένεσις ''genesis'' "origin") is the biological process by which a taxon (of any rank) appears. The science that studies these processes is called phylogenetics. These terms may be ...

and style of locomotion appear to play a role in apical tuft size. Suspensory primates and

New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboidea ( ...

s have the smallest apical tufts, while terrestrial quadrupeds and

Strepsirrhines Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Colle ...

have the largest. A study of the fingertip morphology of four small-bodied New World monkey species, indicated a correlation between increasing small-branch foraging and reduced flexor and extensor tubercles in distal phalanges and broadened distal parts of distal phalanges, coupled with expanded apical pads and developed epidermal ridges. This suggests that widened distal phalanges were developed in arboreal primates, rather than in quadrupedal terrestrial primates.

Cetaceans

Whales exhibit hyperphalangy. Hyperphalangy is an increase in the number of phalanges beyond the

mammal condition of three phalanges-per-digit. Hyperphalangy was present among extinct

marine reptiles Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during the ...

ichthyosaurs Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...

plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...

, and

mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...

-- but not other marine mammals, leaving whales as the only

marine mammals Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their relia ...

to develop this characteristic. The evolutionary process continued over time, and a very derived form of hyperphalangy, with six or more phalanges per digit, evolved convergently in rorqual whales and

oceanic dolphins Oceanic dolphins or Delphinidae are a widely distributed family of dolphins that live in the sea. Close to forty extant species are recognised. They include several big species whose common names contain "whale" rather than "dolphin", such as the ...

, and was likely associated with another wave of signaling within the interdigital tissues.

Other mammals

s (hoofed mammals) the forelimb is optimized for speed and endurance by a combination of length of stride and rapid step; the proximal forelimb segments are short with large muscles, while the distal segments are elongated with less musculature. In two of the major groups of ungulates, odd-toed and even-toed ungulates, what remain of the "hands" — the

and phalangeal bones — are elongated to the extent that they serve little use beyond locomotion. The

giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...

, the largest even-toed ungulate, has large terminal phalanges and fused metacarpal bones able to absorb the stress from running. The

sloth Sloths are a group of Neotropical xenarthran mammals constituting the suborder Folivora, including the extant arboreal tree sloths and extinct terrestrial ground sloths. Noted for their slowness of movement, tree sloths spend most of their li ...

spends its life hanging upside-down from branches, and has highly specialized third and fourth digits for the purpose. They have short and squat proximal phalanges with much longer terminal phalanges. They have

vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...

second and fifth metacarpals, and their palm extends to the distal

interphalangeal joints Interphalangeal joint may refer to: *Interphalangeal articulations of hand The interphalangeal joints of the hand are the hinge joints between the phalanges of the fingers that provide flexion towards the palm of the hand. There are two sets in ...

. The arboreal specialization of these terminal phalanges makes it impossible for the sloth to walk on the ground where the animal has to drag its body with its claws.

Additional images

File:Q11.jpg, Phalanges. File:Carpus2.jpg, Phalanges. File:Unilateral extra phalangeal crease.jpg, Unilateral extra phalangeal crease File:814 Radiograph of Hand.jpg, Joints of the hand in an X-ray image File:Finger phalanges movement.png, Movement of the three finger phalanges; Distal, middle and proximal

References

External links

MedTerms.com Medical Dictionary
{{Authority control Long bones Vertebrate anatomy