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Toe

A toe is one of the five digits located at the distal end of the human foot, serving as a critical component of the lower limb's anatomy for locomotion and stability.^[1] Each foot contains five toes, collectively accounting for 14 of the 26 bones in the foot through their phalangeal structure.^[2] The great toe, also known as the hallux, consists of two phalanges (proximal and distal), while the lesser toes (second through fifth) each have three phalanges (proximal, middle, and distal).^[1] These phalanges articulate via interphalangeal joints, and the proximal phalanges connect to the metatarsal bones of the forefoot through metatarsophalangeal (MTP) joints, which are hinge-like structures essential for flexion and extension.^[1] The toes are supported by a network of intrinsic and extrinsic muscles, tendons, and ligaments that enable precise movements.^[1] Key intrinsic muscles include the flexor digitorum brevis, which flexes the second through fifth toes; the lumbricals, which extend the interphalangeal joints while flexing the MTP joints; and the interossei muscles, responsible for abducting and adducting the toes.^[1] For the great toe, specialized muscles such as the abductor hallucis, flexor hallucis brevis, and adductor hallucis provide additional control for abduction, flexion, and adduction.^[1] The fifth toe is supported by the abductor digiti minimi and flexor digiti minimi brevis.^[1] These muscular attachments, combined with sesamoid bones in the great toe's MTP joint, enhance leverage and reduce friction during weight-bearing activities.^[1] Functionally, the toes play a pivotal role in bipedal gait, balance, and force absorption.^[2] The great toe, in particular, contributes significantly to propulsion during the toe-off phase of walking, bearing a significant portion of the body's weight at push-off, while the lesser toes aid in stability and shock absorption upon heel strike.^[1] This structure allows the foot to adapt to varied terrains, distribute pressure across the forefoot, and maintain postural equilibrium.^[1] Anatomical variations, such as polydactyly (extra toes) or syndactyly (fused toes), can occur but are less common, with the typical configuration optimized for human upright posture.^[1]

Anatomy

Bones and Joints

The human foot contains 14 phalanges, which form the skeletal structure of the toes. The hallux, or big toe, consists of two phalanges: a proximal phalanx and a distal phalanx. Each of the lesser toes (second through fifth) has three phalanges: proximal, middle, and distal.^[3] These phalanges are elongated bones with a base, shaft, and head, adapted for articulation and weight transmission during locomotion.^[4] The toes articulate via synovial joints that enable flexion and extension. The metatarsophalangeal (MTP) joints connect the proximal phalanges to the metatarsal bones at the base of each toe, while interphalangeal (IP) joints link the phalanges within each toe: a proximal IP joint between the proximal and middle phalanges (in lesser toes), and a distal IP joint between the middle and distal phalanges (or proximal and distal in the hallux).^[5] These joints are stabilized by collateral ligaments on the medial and lateral sides, which prevent excessive lateral deviation, and plantar plates—a fibrocartilaginous structure on the volar aspect—that reinforce the joint capsule and maintain alignment under load.^[5] The hallux differs from the lesser toes in its structure and function, featuring two sesamoid bones embedded in the tendon of the flexor hallucis brevis beneath the head of the first metatarsal at the MTP joint. These sesamoids, one medial and one lateral, enhance leverage and distribute pressure during push-off.^[6] In contrast, the lesser toes lack prominent sesamoids, relying more on the metatarsal heads for support.^[7] Biomechanically, the toe bones contribute to weight-bearing by forming the forefoot's transverse arch, with the metatarsal heads collectively comprising the "ball of the foot" that absorbs and propels forces during gait. The first metatarsal head, augmented by sesamoids, bears approximately 30-50% of the body's weight in the stance phase, facilitating efficient propulsion.^[8]

Muscles and Tendons

The toes are primarily moved by a combination of extrinsic muscles originating in the leg and intrinsic muscles within the foot, with long tendons transmitting force to the phalanges. Extrinsic muscles include the flexor digitorum longus, which originates from the posterior surface of the tibia and inserts via tendons into the distal phalanges of toes 2–5, enabling plantarflexion at the interphalangeal joints and contributing to overall toe flexion.^[9] Similarly, the extensor hallucis longus arises from the anterior fibula and interosseous membrane, inserting into the dorsal aspect of the distal phalanx of the great toe to produce dorsiflexion (extension) of the great toe at the metatarsophalangeal and interphalangeal joints.^[9] The extensor digitorum longus, originating from the lateral tibial condyle and fibula, extends toes 2–5 through tendons that divide into slips inserting at the middle and distal phalanges, facilitating dorsiflexion.^[9] These extrinsic tendons cross the ankle and metatarsophalangeal joints, allowing powerful but less precise movements compared to intrinsic actions.^[9] Intrinsic foot muscles provide fine control over toe positioning, particularly for abduction, adduction, and interphalangeal adjustments. The lumbricals (four in number) originate from the tendons of the flexor digitorum longus and insert into the extensor hood expansions of toes 2–5, flexing the metatarsophalangeal joints while extending the interphalangeal joints to support balanced toe posture.^[9] The dorsal interossei (three muscles) abduct toes 2–5 relative to the second toe axis, inserting into the proximal phalanges and extensor hood, whereas the plantar interossei (three muscles) adduct toes 3–5, aiding in inversion-like adjustments during weight-bearing.^[9] These intrinsic muscles enable subtle eversion and inversion per toe through coordinated abduction and adduction, essential for stability on uneven surfaces.^[9] Tendon sheaths and pulleys ensure efficient gliding of these tendons across the foot's joints, minimizing friction and preventing bowstringing. Synovial sheaths envelop the long flexor and extensor tendons, such as those of the flexor digitorum longus and extensor hallucis longus, providing lubrication via peritendinous fluid for smooth passage over the phalanges.^[10] Annular pulleys (A1–A4 in lesser toes, A1–A3 in the great toe) form thickened fibrous bands at the metatarsophalangeal, proximal interphalangeal, and distal interphalangeal joints, acting as reflection pulleys to guide flexor tendons during plantarflexion.^[11] The extensor hood mechanism, a fibroaponeurotic expansion over the dorsal toe, integrates extrinsic tendons like the extensor digitorum longus with intrinsic contributions from lumbricals and interossei, allowing coordinated dorsiflexion at the metatarsophalangeal joint and interphalangeal extension through central and lateral slips.^[12] This hood anchors to the joint capsules, distributing forces to prevent tendon subluxation during toe eversion or inversion.^[12]

Blood Supply

The blood supply to the toes is provided by a network of arteries originating from the anterior and posterior tibial arteries, ensuring oxygenation and nutrient delivery to the phalanges, skin, and soft tissues. The dorsal aspect receives arterial supply primarily from the dorsalis pedis artery, a continuation of the anterior tibial artery, which travels along the dorsum of the foot and gives rise to the arcuate artery and the first dorsal metatarsal artery.^[13] The arcuate artery branches into dorsal metatarsal arteries that supply the second through fifth toes via dorsal digital arteries, while the first dorsal metatarsal artery specifically provides blood to the hallux (big toe) by dividing into medial and lateral branches that form digital arteries for the hallux and the adjacent side of the second toe.^[13] On the plantar surface, the posterior tibial artery divides into the medial and lateral plantar arteries, which form the plantar arch and give rise to plantar metatarsal arteries that branch into plantar digital arteries supplying the toes.^[13] These digital arteries from both dorsal and plantar sources anastomose around the toe joints, creating a rich collateral circulation that supports tissue viability and healing.^[13] Venous drainage from the toes occurs through dorsal and plantar digital veins that converge into the dorsal and plantar venous arches on the respective surfaces of the foot.^[14] The dorsal venous arch, located proximal to the metatarsal heads, drains medially into the great saphenous vein and laterally into the small saphenous vein, facilitating superficial venous return to the femoral and popliteal veins, respectively.^[14] Similarly, the plantar venous arch collects blood from the plantar digital veins and connects to the deep plantar veins, which join the tibial veins for return to the heart, aided by the plantar pump mechanism during weight-bearing.^[14] In the pulp of the toes, arteriovenous anastomoses (AVAs) provide specialized direct connections between arterioles and venules, bypassing the capillary bed to regulate blood flow and heat dissipation.^[15] These AVAs, abundant in the glabrous skin of the toes, enable rapid adjustments in peripheral blood volume for thermoregulation, particularly in response to environmental temperature changes within the thermoneutral zone.^[15]

Nerve Supply

The sensory innervation of the toes is provided primarily by branches of the tibial and common fibular (peroneal) nerves. On the plantar surface, the medial plantar nerve, a terminal branch of the tibial nerve, supplies sensation to the medial three toes (hallux, second, and third) and the medial aspect of the fourth toe, while the lateral plantar nerve innervates the lateral aspect of the fourth toe and the fifth toe.^[16] On the dorsal surface, the superficial fibular nerve provides sensory input to the dorsum of the second through fifth toes and the medial side of the hallux, whereas the deep fibular nerve innervates the skin in the first dorsal web space between the hallux and second toe.^[17]^[18] Motor innervation to the toe muscles arises from the same tibial and fibular nerve branches, targeting both intrinsic and extrinsic muscles. The medial plantar nerve supplies motor fibers to the intrinsic muscles including the abductor hallucis, flexor hallucis brevis, flexor digitorum brevis, and the first lumbrical muscle, facilitating flexion and abduction of the toes.^[16] The lateral plantar nerve innervates the remaining intrinsic muscles such as the quadratus plantae, abductor digiti minimi, adductor hallucis, interossei, and the second through fourth lumbricals, enabling adduction, abduction, and fine toe movements.^[9] Extrinsic muscles acting on the toes, like the flexor digitorum longus and extensor hallucis longus, receive innervation proximally from the tibial and deep fibular nerves, respectively, before their tendons reach the toes.^[19] The toes correspond to dermatomes L4 through S1 of the lumbosacral plexus. The L4 dermatome covers the medial aspect of the foot extending to the medial hallux, L5 supplies the dorsum of the foot and the first three toes, and the S1 dermatome innervates the lateral foot and fifth toe, with digital nerves branching to supply each phalanx.^[20]^[21] Proprioceptive feedback in the toe skin is mediated by specialized mechanoreceptors, including Meissner's corpuscles and Pacinian corpuscles. Meissner's corpuscles, located in the dermal papillae of glabrous skin on the toes, detect low-frequency vibrations (10-50 Hz) and light touch, contributing to tactile discrimination and positional awareness during movement.^[22] Pacinian corpuscles, situated deeper in the dermis and subcutaneous tissue, respond to high-frequency vibrations and rapid pressure changes, aiding in the detection of transient stimuli for proprioceptive integration.^[23]

Anatomical Variations

Anatomical variations in toes encompass a range of normal developmental differences that occur without associated pathology, influencing structure and proportions across individuals. These variations arise primarily from genetic and embryonic factors during limb formation, leading to diversity in digit number, fusion, length, and accessory skeletal elements. Such differences are common and typically asymptomatic, contributing to the wide spectrum of human foot morphology observed in populations worldwide. Polydactyly, characterized by the presence of supernumerary toes, represents a congenital variation where extra digits develop along the preaxial (medial, great toe side) or postaxial (lateral, fifth toe side) borders of the foot. Preaxial polydactyly often involves duplication of the great toe or hallux, while postaxial forms may add digits adjacent to the fifth toe, with prevalence rates varying by population but generally ranging from 1 in 1,000 live births globally. Syndactyly, another frequent congenital anomaly, involves partial or complete soft tissue or bony fusion between adjacent toes, most commonly affecting the second and third toes (toes 2-3), and occurs in approximately 1 in 2,000 to 3,000 births. These conditions stem from disruptions in the apoptotic processes that normally separate digits during embryogenesis, yet they are considered normal variants when isolated and non-debilitating. Variations in phalangeal and metatarsal lengths contribute significantly to toe proportions, with two primary foot types distinguished: the Egyptian foot, where the hallux (big toe) is the longest and subsequent toes taper gradually, and the Greek foot (also known as Morton's toe), featuring a longer second toe relative to the hallux. The Egyptian configuration predominates in many populations, while the Greek type appears in about 5-30% of individuals depending on ethnic groups, reflecting inherited skeletal ratios that influence weight distribution during gait. These length disparities arise from differential growth in the metatarsals and phalanges, with the second metatarsal often being longer in Greek feet, a trait linked to autosomal dominant inheritance patterns. Sesamoid bones, small ossicles embedded within tendons to reduce friction and enhance leverage, exhibit notable presence and positional variations in the toes, particularly in the hallux where paired medial and lateral sesamoids are typically found plantar to the first metatarsophalangeal joint. Absence or asymmetry of these sesamoids occurs in up to 10-15% of cases, with unilateral variations more common, and they may ossify variably with age or remain cartilaginous. Accessory bones, such as the os peroneum—an ossicle within the peroneus longus tendon near the cuboid—represent additional skeletal variants in the lateral foot that can indirectly affect toe alignment, present in 5-30% of individuals and often bilateral. Racial and genetic factors play a key role in the distribution of toe proportions and variations, with certain traits showing higher incidence in specific ethnic groups. For instance, postaxial polydactyly demonstrates elevated prevalence among individuals of African descent, reported at rates up to 10 times higher than in Caucasian populations, attributed to genetic loci influencing limb bud segmentation. Brachymetatarsia, a condition involving congenital shortening of one or more metatarsals (most often the fourth), leading to a shortened toe, has a genetic basis in autosomal dominant mutations and shows variable incidence across races, though isolated cases are documented more frequently in Asian and African cohorts in clinical series. These ethnic disparities highlight polygenic influences on toe morphology, underscoring the need for population-specific anatomical references in medical and forensic contexts.

Function

Role in Locomotion

In human locomotion, toes play a pivotal role during the toe-off phase of the gait cycle, where dorsiflexion of the metatarsophalangeal joints, particularly the big toe, engages the windlass mechanism of the plantar fascia to provide propulsion.^[24] This mechanism, first described by Hicks in 1954, involves the plantar aponeurosis winding around the metatarsal heads as the toes extend, thereby tightening the fascia, elevating the medial longitudinal arch, and stiffening the foot for efficient forward thrust.^[24] The big toe contributes disproportionately to this propulsion during walking by acting as the primary lever against the ground. During the stance phase, toe flexor muscles provide stabilization to maintain foot integrity and prevent collapse, such as excessive flattening that could lead to instability or foot drop tendencies under load.^[25] These muscles, including the flexor digitorum longus and flexor hallucis longus, contract to grip the ground and support the arch, correlating with longer single-limb support duration and reduced risk of gait deviations in older adults.^[25] Weakness in these flexors has been shown to shorten stride length and decrease walking speed, underscoring their role in controlled weight-bearing.^[25] Toe abduction and adduction movements, facilitated by the interossei and abductor hallucis muscles, enhance lateral balance during walking and running by adjusting foot positioning and increasing ground contact area for stability.^[26] Stronger toe muscles correlate with improved anterior-posterior and mediolateral stability, as they generate forces to counteract body sway and support rapid postural adjustments, particularly when leaning forward or on uneven terrain.^[26] This dynamic adjustment helps prevent falls by optimizing the center of pressure under the foot.^[26] In specialized activities like sprinting, intrinsic toe muscles remain active throughout the weight-bearing phase to maximize propulsion and ground reaction forces, enabling faster acceleration compared to walking.^[27] During climbing, toes adapt by gripping small holds, with the big toe providing primary leverage and stability through flexion, allowing precise force application on irregular surfaces for upward progression.^[28]

Sensory and Balance Functions

The toes play a critical role in tactile sensation through specialized mechanoreceptors in the glabrous skin of their pulp and soles, enabling detection of ground textures during contact. Merkel cell-neurite complexes, slowly adapting type I receptors, provide sustained feedback on surface features such as roughness and edges, facilitating precise perception of terrain variations underfoot.^[22] Meissner's corpuscles, rapidly adapting type II receptors densely distributed in the dermal papillae of the toes, detect low-frequency vibrations (10-50 Hz) and subtle skin deformations, contributing to the identification of slip or texture changes on uneven ground.^[29] These receptors, innervated by Aβ low-threshold mechanoreceptive afferents, enhance the toes' ability to sense fine spatial details, which is essential for adaptive responses to environmental surfaces without relying solely on visual cues.^[22] Proprioceptors within the toe joints and tendons further support position sense, particularly during dynamic weight shifts in bipedal posture. Ruffini endings and Pacinian corpuscles in the joint capsules of the metatarsophalangeal and interphalangeal joints detect static and dynamic changes in toe alignment, providing continuous input on angular positions to the central nervous system. Golgi tendon organs embedded in the flexor and extensor tendons of the toes monitor tension variations during load transfer, signaling adjustments needed for maintaining foot arch integrity amid weight redistribution.^[22] This proprioceptive feedback from the toes allows for subconscious corrections in toe flexion or extension, optimizing stability as body weight oscillates forward over the forefoot.^[30] The toes contribute significantly to postural stability by modulating forefoot pressure distribution during quiet standing, where approximately 40% of body weight is borne by the forefoot region including the toes.^[31] This pressure sensing, mediated by mechanoreceptors and joint proprioceptors, helps distribute load across the metatarsal heads and phalanges, reducing sway and enhancing equilibrium by providing real-time data on center-of-pressure shifts.^[32] Studies demonstrate that alterations in toe positioning, such as extension, increase forefoot loading and improve postural control metrics like reduced anterior-posterior sway, underscoring the toes' role in fine-tuning base-of-support stability.^[33] Toe-derived somatosensory inputs integrate with vestibular and visual systems to orchestrate overall balance, forming a multisensory framework for postural control. Foot mechanoreceptors and proprioceptors relay ground reaction forces to the brainstem and cortex, where they converge with vestibular signals from the inner ear (detecting head acceleration) and visual cues (orienting to the environment) via the vestibulospinal and reticulospinal tracts.^[34] This integration, evaluated through sensory organization tests, prioritizes reliable inputs—such as toe pressure when vision is absent—ensuring adaptive postural adjustments and minimizing fall risk in varying conditions.^[35] The somatosensory contributions from the toes thus complement vestibular and visual processing, enabling robust equilibrium through weighted sensory reweighting.^[36]

Clinical Significance

Common Disorders and Injuries

Ingrown toenails, also known as onychocryptosis, occur when the edge of the toenail grows into the surrounding skin, most commonly affecting the big toe. The primary etiology involves improper nail trimming, such as cutting nails too short or in a curved manner rather than straight across, which allows the nail to embed into the soft tissue. Other contributing factors include wearing tight or ill-fitting shoes that compress the toes and predispose the nail to abnormal growth. Symptoms typically include localized pain and tenderness along the nail edge, inflamed and swollen skin, and potential secondary bacterial infection characterized by pus, increased redness, and warmth. General management for mild cases involves soaking the affected foot in warm water several times daily to reduce swelling, gently lifting the nail edge with cotton to promote proper growth, and trimming nails straight across while avoiding cutting too short. If infection develops, topical or oral antibiotics may be necessary, and persistent cases require professional intervention to prevent recurrence.^[37] Toe fractures are common injuries that can be classified as traumatic or stress-related, each arising from distinct mechanisms and often impacting mobility. Traumatic fractures result from acute events, such as a direct blow from dropping a heavy object on the foot or stubbing the toe forcefully, leading to immediate pain, swelling, bruising, and possible deformity if the bone is displaced. In contrast, stress fractures develop gradually from repetitive mechanical loading that exceeds the bone's repair capacity, commonly seen in athletes or individuals suddenly increasing activity levels, with symptoms including insidious onset of pain and mild swelling that worsens with weight-bearing. A notable example is the dancer's fracture, a spiral or avulsion injury at the base of the fifth metatarsal, which indirectly affects the little toe through pain and instability; it typically occurs when the foot rolls inward during activities like ballet, causing sharp pain at the outer foot edge. Management for both types emphasizes rest, ice application, elevation, and non-weight-bearing support via crutches or a walking boot, with taping the injured toe to an adjacent one for stability in minor cases; healing generally occurs within 6-8 weeks without surgical intervention unless displacement is severe.^[38] Infections of the toes frequently involve fungal or bacterial pathogens, with athlete's foot (tinea pedis) being a prevalent fungal condition that primarily affects the interdigital spaces. Caused by dermatophyte fungi thriving in warm, moist environments, tinea pedis spreads through direct contact with infected skin or contaminated surfaces like locker room floors, leading to symptoms such as intense itching, red or scaly rash, peeling or cracking skin between the toes, and occasional blisters or stinging sensations. Risk factors include heavy foot sweating, prolonged wear of occlusive footwear, and walking barefoot in public damp areas, which facilitate fungal proliferation. Bacterial cellulitis, often a complication of minor toe trauma or fungal infections, involves bacterial invasion of the skin and subcutaneous tissues, presenting with rapid-onset redness, swelling, warmth, and pain that may extend beyond the initial site. Diabetes significantly heightens the risk for cellulitis in the toes due to peripheral neuropathy reducing sensation and poor vascular supply impairing immune response, potentially leading to deeper infections like osteomyelitis if untreated. Initial management for tinea pedis includes over-the-counter antifungal creams applied twice daily for 2-4 weeks alongside keeping the feet dry and using moisture-absorbing powders, while cellulitis requires systemic antibiotics for 7-14 days, wound care, and elevation to control spread, with diabetic patients needing prompt evaluation to prevent amputation.^[39]^[40] Gout, particularly in its acute form known as podagra, manifests as intense inflammation in the big toe joint due to monosodium urate crystal deposition. Elevated serum uric acid levels, resulting from overproduction or underexcretion of uric acid—often linked to purine-rich diets, alcohol consumption, or conditions like obesity and hypertension—lead to crystal formation in the synovial fluid, triggering a sudden inflammatory response. Symptoms of podagra include excruciating pain peaking within 4-12 hours, often awakening the individual at night, accompanied by extreme joint tenderness, swelling, redness, and warmth, rendering even light touch unbearable. Risk factors encompass male sex, age over 30, family history, and comorbidities such as diabetes or kidney disease, with attacks potentially recurring if hyperuricemia persists. General management involves rest, ice, elevation, and anti-inflammatory medications like colchicine or nonsteroidal anti-inflammatory drugs to alleviate acute symptoms, alongside long-term strategies to lower uric acid through dietary modifications and urate-lowering therapies.^[41]

Deformities

Toe deformities involve structural misalignments of the toes, often resulting from imbalances in muscle, tendon, and joint forces, leading to progressive bending or deviation that can impair mobility and cause pain. These conditions primarily affect the interphalangeal (IP) joints and metatarsophalangeal (MTP) joint, where flexion contractures develop due to dominance of flexor tendons over extensors.^[42]^[43] Hammertoe is characterized by a flexion contracture at the proximal interphalangeal (PIP) joint, causing the toe—typically the second, third, or fourth—to bend downward in a hammer-like shape, with the distal interphalangeal (DIP) joint remaining neutral or hyperextended. Claw toe, a related deformity, extends this pattern with hyperextension at the MTP joint and flexion at both the PIP and DIP joints, often affecting multiple toes. Both arise from muscle imbalances where extrinsic flexors overpower weakened intrinsic muscles, frequently exacerbated by tight or high-heeled shoes that force toes into prolonged bent positions.^[42]^[44]^[43] Mallet toe presents as an isolated flexion deformity at the DIP joint, usually limited to one toe such as the second or third, without significant involvement of the PIP or MTP joints. This condition often stems from direct pressure from ill-fitting footwear, leading to tightening of the flexor digitorum longus tendon.^[42]^[44] Hallux valgus, commonly known as a bunion, involves lateral deviation of the big toe at the first MTP joint, with medial protrusion of the first metatarsal head, often associated with metatarsus primus varus where the first metatarsal angles inward. The precise etiology is multifactorial, but it commonly progresses from inherited foot structures, joint hypermobility, or biomechanical stresses, worsened by narrow shoes.^[45]^[46] Common risk factors across these deformities include genetic predisposition, such as inherited ligament laxity or foot shape; improper footwear like high heels or pointed toes; and systemic conditions like rheumatoid arthritis, which inflame joints and promote instability. Women face higher risk due to footwear choices and hormonal influences on connective tissue. Progression typically begins as a flexible misalignment, allowing some correction through positioning, but advances to rigid contractures as tendons shorten, joint capsules tighten, and secondary issues like corns or calluses form from friction—potentially leading to chronic pain and altered gait if unmanaged.^[44]^[45]^[46]^[42]

Surgical and Reconstructive Procedures

Surgical and reconstructive procedures for toe conditions primarily address structural deformities, injuries, and severe infections through targeted interventions that restore alignment, function, and stability. Bunionectomy, also known as hallux valgus correction, involves osteotomy of the first metatarsal bone combined with soft tissue realignment to reposition the big toe and alleviate pain from the bunion deformity.^[47] This procedure typically includes a distal soft tissue release to balance the joint capsule and ligaments, enhancing the corrective effect and preventing recurrence.^[48] For mild to moderate cases, a chevron osteotomy is commonly employed, where an incision is made over the big toe joint to cut and realign the bone, often secured with screws or pins for stability.^[49] Outcomes generally show pain relief and improved footwear tolerance for the majority of patients, though recovery may involve 6-12 weeks of limited weight-bearing.^[47] Tendon transfers represent a key technique for correcting flexible claw toe deformities, where the toe exhibits hyperextension at the metatarsophalangeal joint and flexion at the proximal interphalangeal joint. In the flexor-to-extensor rerouting procedure, the flexor digitorum longus or hallucis longus tendon is harvested and transferred to the extensor hood at the base of the proximal phalanx to counteract the imbalance and promote proper toe alignment.^[50] This minimally invasive approach is particularly effective for dynamic deformities, often performed alongside tenotomy or capsulotomy for optimal results.^[51] Postoperative outcomes show effective correction in most cases with low complication rates, allowing patients to resume normal activities within 4-6 weeks.^[52] Amputation of toes is reserved for severe cases where conservative measures fail, with primary indications including uncontrolled infection, extensive tissue necrosis from trauma or vascular insufficiency, and non-viable bone due to osteomyelitis.^[53] Procedures are performed at varying levels depending on the extent of involvement: partial phalangeal amputation for distal tip gangrene, interphalangeal disarticulation for mid-toe involvement, or ray amputation (toe plus metatarsal) for proximal spread to preserve foot function.^[54] In diabetic patients, early intervention at the phalangeal level can prevent progression to higher amputations, with healing achievable in many cases when vascular status is adequate.^[53] Reconstructive options post-injury focus on restoring joint integrity and preventing chronic instability, with arthrodesis (joint fusion) serving as a reliable method for toe interphalangeal or metatarsophalangeal joints damaged by trauma. This involves removing articular cartilage and fixating the bones with screws, plates, or wires to achieve bony union, thereby eliminating painful motion while maintaining weight-bearing capacity.^[55] For the first metatarsophalangeal joint, fusion is the gold standard in post-traumatic arthritis or instability, achieving union in over 90% of cases and significant pain reduction.^[56] Recovery typically spans 8-12 weeks, with patients progressing to full weight-bearing in supportive footwear once radiographic fusion is confirmed.^[57]

Development and Evolution

Embryological Development

The embryological development of the toes originates from the lower limb buds, which emerge during the fourth week of gestation as protrusions from the lateral plate mesoderm, covered by ectoderm. These buds rapidly elongate under the influence of the apical ectodermal ridge (AER), a thickened ectodermal structure at the distal margin that secretes fibroblast growth factors (FGFs), such as FGF8, to promote proximo-distal outgrowth and patterning of the limb elements, including the future toes.^[58]^[59] The AER interacts with the underlying progress zone of mesenchyme to ensure sequential development from proximal to distal structures.^[60] By the sixth week, the flattened limb paddle develops paddle-like digital rays, the anlagen of the five toes, initially connected by interdigital tissue. Digit separation occurs between weeks 7 and 8 through interdigital necrosis, a localized programmed cell death (apoptosis) in the mesenchymal webbing, sculpting individual toes while preserving the rays.^[58]^[61] This process is regulated by signaling molecules like bone morphogenetic proteins (BMPs), which trigger cell death in the interdigital regions.^[62] Phalangeal segmentation within each toe arises from chondrification centers that form around week 6, where mesenchymal condensations differentiate into cartilage templates for the proximal, middle, and distal phalanges.^[61] These centers segment through differential growth and joint formation, with the hallux (first toe) differentiating distinctly by retaining only two phalanges, a pattern established by week 7 via anterior-posterior signaling gradients independent of Sonic hedgehog (SHH) for the first digit.^[58]^[63] Exposure to teratogens during the critical window of weeks 4-8 can disrupt toe formation; for instance, thalidomide inhibits angiogenesis and interferes with cereblon-mediated protein degradation, leading to limb reduction defects such as phocomelia and alterations in digit number or structure.^[64]^[65]

Evolutionary Aspects

The evolution of the human toe reflects a profound transition from prehensile appendages adapted for arboreal locomotion in early primate ancestors to rigid, weight-bearing structures optimized for terrestrial bipedalism in hominins. In arboreal primates, toes functioned primarily for grasping branches, with long, curved phalanges and opposable digits enabling secure arboreal travel. This grasping capability persisted in early hominins but gradually diminished as bipedal locomotion became habitual, shifting the foot's role to propulsion and shock absorption during upright walking. Fossil evidence indicates this mosaic evolution began around 6 million years ago, with early hominins retaining ape-like toe features while developing initial bipedal adaptations.^[66] A key milestone occurred with the loss of opposability in the lesser toes (digits 2–5), which aligned parallel to the metatarsals and shortened, reducing their grasping function to facilitate efficient weight transfer during bipedal strides. This change is evident by approximately 4.4 million years ago in Ardipithecus ramidus, whose foot phalanges show reduced curvature compared to modern apes, indicating early commitment to terrestrial locomotion. In contrast, Australopithecus species (circa 4–2 million years ago) further solidified these adaptations, with fossilized foot bones from A. afarensis displaying straight, non-opposable lesser toes that supported a more stable platform for walking, as seen in the Laetoli footprints. These modifications enhanced energy efficiency in bipedalism by creating a lever-like forefoot.^[67]^[68]^[66] The big toe (hallux) underwent a more protracted transformation, evolving from a divergent, partially opposable structure to a fully adducted, aligned digit crucial for push-off in modern human gait. In Ardipithecus ramidus, the hallux remained widely abducted, retaining prehensile potential for climbing while the midfoot exhibited increased rigidity absent in apes, allowing limited bipedal propulsion. This partial grasping persisted into early Australopithecus but was largely lost by around 2.3–1.2 million years ago in early Homo species, where joint morphology shows complete adduction and robust alignment with the other toes, enabling forceful toe-off and longitudinal arch support. Modern human toes, by comparison, are short, straight, and non-grasping, contributing to the foot's overall stiffness and efficiency in bipedal endurance running and walking.^[67]^[68]^[68]

Etymology and History

Origins of Terminology

The English word "toe" originates from the Old English "tā," referring to a digit of the foot or a finger-like projection, which traces back to the Proto-Germanic *taihwō(n), meaning a toe or similar appendage.^[69] This term evolved through Middle English "to," maintaining its association with the protruding parts of the foot, akin to cognates in Old Norse "tá" and Old High German "zēha."^[70] The term "hallux," used specifically for the big toe, derives from Late Latin "hallux" or "allus," denoting the great toe, of unknown origin but possibly influenced by Greek terms implying a prominent structure.^[71] Introduced into modern anatomical nomenclature in the early 19th century, it emphasizes the toe's size and positional significance in the foot.^[72] In anatomical convention, toes are designated as digits 1 through 5, with the hallux as digit 1 (medialmost) and the little toe as digit 5 (lateralmost), a numbering system originating from standard human morphology descriptions to facilitate consistent medical and scientific reference.^[73] Culturally, informal terms include "pinky toe" for the fifth digit due to its small size, and slang like "dogs" for feet or toes collectively, emerging in early 20th-century American English from rhyming slang ("dog's meat" for "feet") to describe sore or tired extremities.^[74]^[75]

Historical Perspectives

Ancient Egyptian practices demonstrate early attention to toe preservation and treatment. During mummification, embalmers meticulously preserved foot structures, including toes, using natron and resins to dehydrate and protect tissues, allowing modern examinations to reveal pathologies such as deformities and even prosthetic toes crafted from wood or linen for the afterlife. Recent studies, including biomechanical analyses as of 2017, confirm these prosthetics were functional, aiding gait.^[76]^[77] The Ebers Papyrus, dating to approximately 1550 BCE, documents remedies for toe ailments, including salves with red ochre, natron, and linseed applied to painful areas, as well as treatments for conditions resembling gout using catfish extracts mixed with other ingredients.^[78]^[79] These texts highlight a blend of empirical observation and magical incantations in addressing toe pain and inflammation.^[80] In the 5th century BCE, Greek physician Hippocrates provided one of the earliest detailed clinical descriptions of gout, particularly its manifestation as podagra—an acute inflammation of the big toe joint—characterizing it as "the unwalkable disease" due to severe pain that rendered patients immobile.^[81] His observations in the Hippocratic Corpus emphasized environmental and dietary factors, such as rich foods and seasonal changes, influencing toe joint afflictions, laying foundational principles for later rheumatology. The Renaissance marked a pivotal advancement in anatomical understanding of toes through Andreas Vesalius' seminal work, De Humani Corporis Fabrica (1543), which featured precise woodcut illustrations of the lower limb musculature, including the intrinsic and extrinsic muscles controlling toe flexion and extension.^[82] Vesalius' dissections corrected Galenic errors and depicted toe structures in layered views, from skeletal to muscular systems, influencing subsequent anatomical studies. In the 19th and 20th centuries, toe-related pathologies gained further recognition with Thomas G. Morton's 1876 description of metatarsalgia, now known as Morton's neuroma, a painful thickening of tissue around a nerve between the toes, often the third and fourth.^[83] Post-World War II, podiatry emerged as a modern specialized field, driven by the need to treat soldiers' foot injuries; in the mid-20th century, particularly from the 1960s, U.S. practitioners fully transitioned from chiropody to earning Doctor of Podiatric Medicine degrees, with formalized curricula emphasizing surgical and conservative toe interventions.^[84]

Comparative Anatomy

Toes in Non-Human Animals

In mammals, toe structures exhibit significant diversity adapted to various locomotor needs. Carnivores, such as cats in the family Felidae, possess clawed toes where the claws are retractile, allowing them to be extended and retracted via a specialized mechanism involving elastic ligaments and the deep digital flexor tendon; this structure keeps the claws sharp and protected when not in use.^[85] In contrast, ungulates display hoofed toes, with even-toed ungulates (Artiodactyla) like cattle and deer featuring cloven hooves formed by the fusion of digits II and III, while odd-toed ungulates (Perissodactyla) such as horses have a single weight-bearing toe (digit III) encased in a solid hoof.^[86]^[87]^[88] Vestigial toes are common in some mammals, serving as non-weight-bearing remnants. In dogs (Canis familiaris), dewclaws represent these reduced digits, typically located on the medial aspect of the forelimbs (digit I) and sometimes hindlimbs, lacking full muscular and tendinous attachments for ground contact and thus not contributing to primary locomotion.^[89]^[90] Birds generally have anisodactyl or zygodactyl foot configurations, with the latter featuring a reversed fourth toe (digit IV) pointing backward alongside the hallux (digit I), while digits II and III point forward; this arrangement is prevalent in orders like Psittaciformes (parrots) and Strigiformes (owls), enabling enhanced grasping.^[91]^[92] The zygodactyl pattern has evolved independently multiple times, as evidenced by developmental studies showing backward rotation of digit IV during embryogenesis.^[93] Reptiles display a range of toe morphologies, with most lizards (Squamata: Lacertilia) retaining pentadactyl limbs—four limbs each with five toes (digits I–V) terminating in claws—reflecting the ancestral tetrapod condition.^[94] In snakes (Squamata: Serpentes), evolutionary limb reduction has led to the complete loss of external toes and limbs in most species, though vestigial hindlimb rudiments persist internally as scaled protrusions in some taxa like boas, resulting from modifications in gene regulatory networks during development.^[95]^[96]

Adaptations Across Species

In primates, prehensile toes enable enhanced grasping capabilities essential for arboreal locomotion, including climbing and brachiation. The opposable hallux (big toe) in species such as chimpanzees and gorillas allows the foot to function similarly to a hand, facilitating secure grips on branches during vertical climbing and suspension.^[97] This adaptation is particularly evident in great apes, where the foot's flexibility and muscle arrangement support propulsive forces during tree traversal, with studies showing that hindlimb grasping contributes significantly to stability in dynamic arboreal environments.^[98] Opposability in the big toe persists as a primitive trait across many primate lineages, aiding in load distribution and preventing slippage on irregular surfaces.^[99] Webbed toes represent a key hydrodynamic adaptation in semi-aquatic and fully aquatic species, increasing surface area for propulsion during swimming. In otters, such as the North American river otter (Lontra canadensis), fully webbed hind feet act as efficient paddles, enhancing thrust and maneuverability in water while allowing terrestrial mobility through partial retraction of the webbing.^[100] Similarly, sea otters (Enhydra lutris) exhibit densely webbed hind paws that generate greater propulsive efficiency, supporting sustained foraging dives and rapid directional changes in marine habitats.^[101] Among amphibians, webbed feet in anurans like frogs optimize swimming performance by creating vortex rings for forward propulsion, with the degree of webbing varying to balance aquatic efficiency against terrestrial needs.^[102] This convergent evolution underscores the webbing's role in reducing drag and amplifying force application in fluid media.^[103] Arboreal reptiles like geckos demonstrate specialized padded toes for adhesion on vertical and inverted surfaces, crucial for navigating foliage and bark. The subdigital pads of geckos are covered in millions of microscopic setae—branched, hair-like structures—that exploit van der Waals forces to achieve reversible attachment, enabling them to climb smooth substrates without residue.^[104] These setae enhance frictional adhesion during dynamic locomotion, with adhesion strength increasing at higher sliding speeds to support rapid traversal of uneven arboreal terrains.^[105] The hierarchical structure of setae, from macroscale pads to nanoscale spatulae, allows directional control, where toe angling maximizes contact and detachment for efficient movement.^[106] Cursorial adaptations in ungulates, exemplified by horses (Equus spp.), involve elongation of central toes and reduction of lateral digits to optimize speed and endurance on open plains. In modern equids, the single elongated third digit forms a hoof that concentrates force for efficient ground contact, minimizing energy loss during high-speed galloping.^[107] This monodactyly evolved from multi-toed ancestors through progressive reduction, driven by selective pressures for cursorial locomotion, where longer toes and fused metacarpals enhance stride length and stability at velocities exceeding 50 km/h.^[108] Fossil evidence indicates that side toe vestiges in early equids provided auxiliary support on soft substrates, but their elimination streamlined limb mechanics for sustained terrestrial pursuit.^[109]

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In this study, we discovered that gecko setae stick more strongly the faster they slide, and do not wear out after 30 000 cycles.
[105]
Repeated Origin and Loss of Adhesive Toepads in Geckos
The key component of the adhesive apparatus in lizards is the presence of setae, microscopic hair-like outgrowths of the superficial layer of the subdigital ...<|separator|>
[106]
Mechanics of evolutionary digit reduction in fossil horses (Equidae)
Aug 23, 2017 · Digit reduction is a major trend that characterizes horse evolution, but its causes and consequences have rarely been quantitatively tested.
[107]
Evolution of a Single Toe in Horses: Causes, Consequences, and ...
May 24, 2019 · Many artiodactyl taxa evolved elongated limbs and reduced digits throughout the Cenozoic (Janis and Wilhelm 1993), and they have been ...
[108]
The Evolution of Equid Monodactyly: A Review Including a New ...
Sondaar (1968) claims that the medial and lateral digits were reduced in size, but Parker et al. (2018) report a toe reduction index less than most other North ...