The biceps brachii is a two-headed skeletal muscle situated in the anterior compartment of the upper arm, originating from the supraglenoid tubercle of the scapula (long head) and the coracoid process of the scapula (short head), and inserting via its tendon at the radial tuberosity and the bicipital aponeurosis into the fascia of the forearm flexors.[1] It is innervated by the musculocutaneous nerve (arising from spinal roots C5 and C6) and receives its primary blood supply from muscular branches of the brachial artery.[1][2]As a biarticular muscle crossing both the shoulder and elbow joints, the biceps brachii serves multiple key functions: it acts as a powerful supinator of the forearm (especially when the elbow is flexed), a flexor of the elbow joint (working synergistically with the brachialis and brachioradialis), and a minor contributor to shoulder flexion and stabilization of the glenohumeral joint during armelevation.[1][3] Its long head tendon, in particular, provides dynamic stability to the shoulder by resisting superior humeral head translation, particularly in the initial phases of armabduction or elevation up to 30 degrees.[3] The muscle's fusiform shape and parallel fiber arrangement enable forceful contractions essential for everyday activities like lifting, turning the palm upward (supination), and stabilizing the arm during overhead motions.[1][4]Embryologically, the biceps brachii develops from the myotomes of somites during the fifth week of gestation, forming part of a common muscular mass with the coracobrachialis and brachialis before differentiating into distinct structures.[1] Clinically, it is notable for conditions like biceps tendinopathy, where the long head tendon may become inflamed or torn, often treated via tenotomy or tenodesis to prevent cosmetic deformities such as the "Popeye" sign in older patients.[1] Variations, including supernumerary heads, occur with a prevalence ranging from approximately 8% to 20% across populations and can influence surgical approaches or athletic performance.[4]
Anatomy
Origin and insertion
The biceps brachii muscle originates proximally via two distinct heads on the scapula. The long head arises from the supraglenoid tubercle and the superior portion of the glenoid labrum.[1] The short head originates from the apex of the coracoid process of the scapula, sharing this attachment site with the coracobrachialis muscle.[5]From their origins, the two heads follow divergent paths before converging. The tendon of the long head is long and thin, measuring approximately 9 cm, and passes intracapsularly through the glenohumeral joint capsule before exiting and traveling within the intertubercular (bicipital) groove of the humerus, held in place by the transverse humeral ligament; this positioning allows it to contribute to shoulder joint stability by depressing the humeral head during arm elevation.[1] In contrast, the short head tendon courses anterior to the shoulder joint capsule and lies medial to the long head. The heads merge into a single fusiform muscle belly in the anterior compartment of the arm, approximately midway along the humerus.[6]Distally, the muscle tapers to form a common tendon that inserts primarily onto the bicipital tuberosity of the radius.[7] A flat, broad expansion known as the bicipital aponeurosis arises from this tendon and extends medially to blend with the deep fascia overlying the flexor muscles of the forearm, providing additional attachment and distributing force across the antebrachial structures.[5]In adults, the total length of the biceps brachii, from origin to insertion, measures approximately 25-30 cm, varying with arm length and individual anatomy.
Composition
The biceps brachii muscle is a fusiform structure consisting of two distinct heads—the long head and the short head—that arise separately from the scapula and converge distally to form a single, spindle-shaped belly in the anterior upper arm. This bipennate-like arrangement, with fibers oriented primarily parallel to the muscle's long axis, facilitates efficient contraction for power generation. The muscle is predominantly composed of type II fast-twitch fibers, which account for approximately 60% of the fiber population in humans, enabling rapid force production suited to dynamic movements.[8]The proximal tendons exhibit distinct anatomical features: the long head tendon traverses an intra-articular path through the glenohumeral joint capsule, while the short head tendon follows an extra-articular course from the coracoid process. Distally, the tendon extends 2-3 cm from the muscle belly before flattening into a broad aponeurosis that inserts on the radial tuberosity and forearmfascia.[1][9]Architecturally, the biceps brachii features parallel fiber alignment within the belly, with a low pennation angle ranging from about 4° in elbow extension to 13° in flexion, which minimizes force loss and supports high contraction velocity. The physiological cross-sectional area averages 8.2 cm² (SD 3.4 cm²) in adults, reflecting its capacity for substantial force output relative to body size.[10][11]Connective tissue components include the epimysium encasing the entire muscle, perimysium organizing fiber bundles into fascicles, and endomysium sheathing individual fibers, all contributing to structural integrity and force transmission. The lacertus fibrosus, or bicipital aponeurosis, emerges from the distal tendon and distributes the muscle's contractile force to adjacent forearm structures, thereby reducing direct loading on the radial insertion site.[1]
Variations
The biceps brachii muscle exhibits several anatomical variations, most notably the presence of an accessory or third head, which occurs with an overall pooled prevalence of 9.6% (95% CI 8–11%) across studied populations.[12] This third head typically originates from the anteromedial aspect of the humerus, between the insertion sites of the coracobrachialis and brachialis muscles, and joins the common distal tendon of the biceps brachii to insert on the radial tuberosity.[13] Prevalence rates for this variation range from 3% to 20% depending on the study cohort, with the accessory head more commonly unilateral than bilateral.[14]Variations in the origins of the biceps heads include rare instances of fusion between the long and short heads, where the two proximal tendons merge early along their course, reported in fewer than 5% of dissections.[15] Complete absence of the short head is exceptionally uncommon, documented primarily through isolated case reports rather than population-level data, often resulting in a unipennate muscle structure.[16] In some cases, the long head may exhibit a humeral origin as part of the third head configuration, altering its proximal trajectory without affecting the short head.[17]Distal insertion anomalies of the biceps brachii include tendinous slips fusing with the brachialis muscle or extending to the ulna via connections to the bicipital aponeurosis, observed in approximately 5–10% of anatomical specimens.[18] These variations show gender differences, with females exhibiting higher rates of complex distal insertions, such as multiple tendon bands (type III morphology in up to 7.5% of cases, significantly more frequent in females than males).[19]Population-specific traits influence the incidence of these variations, particularly the third head, which appears more frequently in Asian cohorts (e.g., 18% in Japanese and 8% in Chinese populations) compared to Europeans (around 10%).[20] Left-right asymmetry is common, with accessory heads occurring unilaterally in about 50% of affected individuals, leading to bilateral symmetry in only half of cases.[13]
Innervation
The biceps brachii muscle receives its primary motor innervation from the musculocutaneous nerve, a terminal branch of the lateral cord of the brachial plexus derived from spinal roots C5-C7.[21] The musculocutaneous nerve pierces the coracobrachialis muscle near its insertion on the humerus and emits motor branches to the biceps brachii shortly thereafter, typically in the mid-arm region.[22] These branches supply both the long and short heads of the muscle, with anatomical studies identifying distinct patterns: in approximately 28% of cases, separate branches innervate the long head and short head individually, while a single branch or additional supply to the common belly occurs in the majority.[23]Intramuscular branching of these motor nerves forms characteristic motor endplate bands within the biceps brachii, appearing as an inverted V-shaped zone approximately 1 cm wide, located 7-11 cm superior to the olecranon process depending on the medial-to-lateral position.[24] This organization facilitates coordinated contraction across the muscle heads and is relevant for targeted interventions such as botulinum neurotoxin injections. Sensory innervation includes proprioceptive fibers conveyed via the same musculocutaneous nerve, primarily from C5-C6 roots, which monitor muscle length and tension through spindle afferents.[1]Anatomical anomalies in biceps brachii innervation are uncommon, occurring in 1-2% of cases, and may involve bifurcation of the musculocutaneous nerve or rare accessory contributions, such as indirect involvement from the accessory phrenic nerve through brachial plexus variations, potentially altering motor distribution to the coracobrachialis and biceps.[25]
Blood supply
The biceps brachii muscle receives its arterial blood supply from multiple sources that ensure oxygenation and nutrient delivery across its proximal, main, and distal regions. The proximal portion, particularly the long head originating from the supraglenoid tubercle, is primarily supplied by branches of the suprascapular artery and the anterior circumflex humeral artery, which enter near the glenohumeral joint and provide vascularization to the intra-articular tendon segment.[26][27] In contrast, the short head, arising from the coracoid process, derives its proximal supply mainly from the anterior circumflex humeral artery, which courses deep to the muscle and delivers blood via ascending branches.[28]The primary vascularization of the muscle belly occurs through nutrient branches from the brachial artery and the profunda brachii artery, which penetrate the muscle along its length to form an extensive intramuscular network. These branches, often numbering up to eight, arise predominantly from the middle third of the brachial artery and distribute to both heads, supporting the bulk of metabolic demands during contraction.[1][29]Distally, at the tendon insertion on the radial tuberosity, blood supply is provided by the radial recurrent artery, which typically crosses volar to the tendon approximately 4 mm proximal to the insertion and forms anastomoses that nourish the enthesis. Venous drainage parallels this arterial system, with accompanying venae comitantes collecting blood from the muscle and tendon, ultimately draining into the brachial veins and then the axillary vein.[30][31]The biceps brachii demonstrates a high capillary density, especially in type I (slow-twitch) fibers, averaging 4.9 to 5.5 capillaries per fiber, which facilitates efficient oxygen delivery and contributes to the muscle's role in sustained endurance activities such as forearm supination. Arterioles branch frequently, typically every 1-2 mm along the muscle length, enhancing this microvascular architecture.[32]
Function
Elbow flexion
The biceps brachii contributes to elbow flexion by contracting to shorten its muscle fibers, which pulls the radial tuberosity toward the humerus through its distal insertion at the bicipital tuberosity of the radius.[1] This action flexes the forearm at the elbow joint, with the muscle spanning from its origins on the coracoid process and supraglenoid tubercle to its insertion via the bicipital aponeurosis.[1]In terms of force generation, the biceps brachii produces peak isometric torque of approximately 50-70 Nm in healthy adults during elbow flexion, with maximum values occurring at around 90° of flexion due to optimal moment arm length.[33] The muscle's length-tension relationship reaches its optimum at elbow flexion angles of 110-120°, where sarcomere overlap allows for maximal active force production.The biceps brachii functions as a synergist to the primary elbow flexor, the brachialis, and the secondary flexor brachioradialis, all of which contribute to net flexion torque at the elbow.[1] Its action is opposed by the antagonist triceps brachii, which extends the elbow.[1]Electromyographic studies show that the biceps brachii exhibits its highest activation levels, reaching 80-100% of maximum voluntary contraction (MVC), during isolated isometric elbow flexion tasks against resistance, particularly in supinated forearm positions.[35]
Forearm supination
The biceps brachii contributes to forearm supination by rotating the radius such that the palm faces upward, a motion facilitated by the spiral path of its distal tendon around the radial tuberosity. This insertion creates a cam effect, where contraction of the muscle generates supinatory torque as the tendon wraps around the radius, optimizing rotational force during elbow flexion.[36] The torque is maximal when the elbow is flexed, such as at 90°, as this position reduces the leverage of antagonist muscles like the pronator teres and aligns the biceps' line of pull for greater rotational efficiency.[37] With elbow extension, the effectiveness of supination diminishes because the moment arm shortens, altering the tendon's alignment relative to the radioulnar joint axis.[37]In terms of torque output, the biceps brachii can produce approximately 10-20 Nm of supinatory force at neutral forearm position with the elbow flexed, though values vary by individual factors like age and training status.[38] This capacity allows the biceps to interact synergistically with the supinator muscle to overcome pronator forces from muscles such as the pronator teres, enabling net supination even against resistance.[39] The biceps provides the primary supinatory power, often exceeding the supinator's contribution, particularly when the forearm starts from a pronated position.[40]Kinematically, the biceps brachii plays an essential role in fine motor tasks requiring precise supination, such as turning a key or screwing a lid, where controlled rotation is critical for hand positioning. Electromyographic (EMG) studies show higher biceps activation in supinated forearm positions during elbow flexion tasks between 60° and 90° of elbow flexion, reflecting heightened recruitment to sustain torque in mid-range positions.[41] This activation pattern synergizes with elbow flexion, enhancing overall supinatory efficiency without relying on isolated extension.[42]
Accessory roles
The long head of the biceps brachii contributes to shoulder stabilization by exerting a depressive force on the humeral head during arm elevation, counteracting superior translation and maintaining glenohumeral congruence. This action is mediated through the tendon's intra-articular course, which tensions to resist upward migration of the humeral head, particularly under dynamic loads such as overhead reaching. In healthy individuals, electromyographic studies indicate modest activation of the long head during flexion and abduction, with activity increasing proportionally to load demands (e.g., 11.6% maximum voluntary contraction at 90° flexion with added weight). In cases of rotator cuff pathology, this depressive role becomes more pronounced, as the biceps compensates for deficient cuff muscles to preserve joint centering.[43][44]Beyond direct stabilization, the biceps brachii aids in postural support of the upper limb by countering gravitational forces during mid-range elbow flexion, helping to sustain arm position without excessive energy expenditure from primary movers. This isometric contribution enhances overall upper limb endurance in activities like holding objects at waist level. Additionally, the muscle generates compressive forces across the humeroulnar joint, promoting joint integrity and load distribution during sustained postures. These effects are evident in biomechanical models showing coordinated activation with synergists to minimize shear stresses.[3]Reflexive mechanisms further underscore the biceps' accessory functions, with Golgi tendon organs embedded in its tendon providing autogenic inhibition feedback to regulate tension. This sensory input facilitates load sharing among elbow flexors, such as the brachialis and brachioradialis, by inhibiting excessive biceps activation during high-force tasks and redistributing effort to prevent fatigue or injury. Computational simulations demonstrate that integrating Golgi tendon organ signals with muscle spindle data reduces postural errors by up to 70% and accelerates motor responses by 50% in multi-muscle coordination.[45]Despite these supportive roles, the biceps brachii exhibits clear limitations in upper limb mechanics. It plays no significant part in shoulder adduction or extension, as its line of pull favors anterior translation and flexion instead. Furthermore, in full forearm pronation, the muscle deactivates substantially, diminishing its contributions to both stabilization and compression due to biomechanical disadvantage. While primary actions in elbow flexion and supination form the basis for these accessory effects, anatomical variations like tendon path anomalies can modulate stability outcomes.[3]
Clinical significance
Tendon disorders
Tendon disorders of the biceps brachii encompass a range of non-traumatic inflammatory and degenerative conditions affecting the proximal and distal tendons, often resulting from repetitive stress or underlying shoulder pathology. These disorders primarily involve the long head of the biceps tendon (LHBT) in its proximal portion and the distal tendon at its insertion on the radial tuberosity, leading to pain, functional limitations, and potential progression if untreated.[46][47]Bicipital tendinitis, also known as proximal biceps tendinitis, refers to inflammation of the LHBT within the bicipital groove of the humerus, typically arising from overuse in activities involving repetitive overhead motions such as throwing, swimming, or serving in racket sports.[48][49] This condition is characterized by microtears and synovitis in the tendon sheath, exacerbated by impingement from surrounding structures like the rotator cuff. Symptoms include a deep, throbbing ache in the anterior shoulder that may radiate distally toward the elbow, along with localized tenderness over the bicipital groove and mild weakness during shoulder flexion or elevation.[50][46] Pain is often worse at night or with resisted supination, and the condition predominantly affects active individuals in their 40s and 50s.[49]Distal biceps tendinosis involves degenerative changes, including collagen disorganization and mucoid degeneration, at the tendon's insertion on the radial tuberosity, commonly seen in manual laborers engaged in heavy lifting or repetitive gripping tasks that impose eccentric loads on the elbow.[47] Unlike acute injuries, this chronic process develops gradually from cumulative microtrauma, leading to tendon thickening and reduced elasticity without significant inflammation. Clinical presentation features insidious onset of anterior elbowpain, exacerbated by flexion or supination against resistance, accompanied by localized swelling and weakness in gripping activities, though full rupture is rare in this degenerative stage.[47][51]Subluxation of the LHBT occurs when the tendon medially displaces from the bicipital groove, often secondary to rotator cuff tears that compromise the stabilizing pulley formed by the superior glenohumeral ligament and subscapularis tendon.[52] This instability is most frequently associated with subscapularis pathology, as the tendon's medial restraint is lost, allowing dynamic or static dislocation during shoulder motion. It arises in 20-30% of rotator cuff tear cases, with higher rates in full-thickness subscapularis defects.[53] Patients typically report snapping sensations, anterior shoulder pain, and a palpable tendon shift in the groove, particularly with abduction and external rotation, which can mimic or coexist with impingement syndromes.[52][54]Diagnosis of biceps tendon disorders relies on a combination of clinical tests and imaging to confirm pathology and rule out associated conditions. The Speed's test, performed by having the patient flex the shoulder forward against resistance with the elbow extended, elicits anterior shoulder pain indicative of LHBT irritation, showing moderate sensitivity (around 60-70%) for tendinitis.[49][55] Yergason's test, involving resisted supination with the elbow flexed at 90 degrees, reproduces pain or subluxation in the bicipital groove, with specificity up to 78% for proximal tendon issues.[56][55] Magnetic resonance imaging (MRI) provides definitive visualization, revealing tendon thickening greater than 7 mm, increased intrasubstance signal, or contour abnormalities as hallmarks of tendinosis or tendinitis, while also assessing for subluxation through dynamic sequences if needed.[46][57]
Rupture
A biceps tendon rupture involves the complete or partial tearing of the tendon connecting the biceps brachii muscle to the bone, most commonly at the proximal or distal attachment sites. Proximal ruptures, which account for 90-97% of all biceps tendon ruptures and primarily affect the long head tendon at the superior glenoid labrum, often occur spontaneously due to degenerative changes in older individuals.[58] In contrast, distal ruptures at the radial tuberosity represent only 3-10% of cases but are more likely to result from acute trauma, such as an eccentric overload during forceful elbow extension against resistance.[59] These injuries predominantly affect men, with proximal ruptures common in those over 60 years and distal ruptures in middle-aged individuals aged 40-60.[60]The incidence of distal biceps ruptures is estimated at 2-5.4 per 100,000 patient-years, while proximal ruptures occur more frequently but lack precise population-level data due to their often conservative management.[61] Risk factors for both types include advanced age, male sex, smoking, which impairs tendon vascularity and strength, chronic corticosteroid use that weakens connective tissue, and hyperlipidemia associated with degenerative tendon changes.[60] Additional contributors encompass obesity, overuse from repetitive heavy lifting, and rarely, conditions like diabetes or fluoroquinolone antibiotic use.[60]Symptoms typically manifest acutely with a sudden sharp pain in the anterior shoulder or elbow, often accompanied by an audible "pop" or snapping sensation at the moment of injury.[62] A characteristic "Popeye" deformity arises from proximal retraction of the muscle belly, creating a visible bulge in the upper arm, particularly prominent in long head proximal ruptures.[62] Bruising, swelling, and cramping with arm use follow, alongside functional deficits such as weakness in elbow flexion and forearm supination; untreated distal ruptures can result in 30-50% loss of supination strength, while proximal injuries cause milder flexion weakness of about 20%.[63]Diagnosis relies on clinical examination, including the hook test for distal integrity and assessment of the deformity, with ultrasound or MRI confirming the tear extent and retraction.[58]Treatment varies by rupture location and patient factors. Proximal long head ruptures are managed conservatively in most cases with a sling for 1-2 weeks, ice, nonsteroidal anti-inflammatory drugs, and physical therapy to restore range of motion and strength, yielding good functional outcomes without surgery for non-athletes.[62] Surgical intervention for proximal tears, such as tenodesis to the humerus, is reserved for young active individuals or those with persistent pain, cramping, or cosmetic concerns.[62] Distal ruptures, however, warrant prompt surgical reattachment to prevent permanent strength deficits, ideally within 2-3 weeks using techniques like the single-incision anterior (Henry) approach with suture anchors or the double-incision (Boyd-Anderson) method to minimize nerve injury risk.[63] Postoperative rehabilitation involves immobilization followed by progressive therapy, with surgical repair achieving over 90% restoration of strength and range of motion in most patients.[64]Conservative management of distal ruptures leads to acceptable results in low-demand patients but with notable supination weakness and fatigue.[63]
Training and rehabilitation
Strengthening the biceps brachii typically involves resistance exercises targeting elbow flexion and forearm supination, with common variations including barbell curls, dumbbell curls, concentration curls, and hammer curls to emphasize the brachialis.[65] These exercises promote balanced activation, as electromyographic (EMG) studies show higher biceps brachii excitation during supinated grips in standard curls and increased brachialis involvement with neutral grips in hammer curls.[42] Guidelines recommend 3 sets of 8-12 repetitions at 70-80% of one-repetition maximum (1RM) to optimize hypertrophy and strength gains, performed 2-3 times per week with progressive overload.[66]Resistance training induces physiological adaptations such as muscle hypertrophy, where the cross-sectional area of the biceps brachii can increase by approximately 10-20% over 6-12 months of consistent programming, depending on volume and individual factors.[67] EMG-guided protocols ensure balanced activation across muscle heads, with variations like preacher curls showing superior long-head recruitment compared to standard curls.[68]Rehabilitation following biceps injuries, such as ruptures, employs progressive loading to restore function while minimizing re-injury risk. Initial phases (4-6 weeks post-rupture or repair) focus on isometrics and gentle range-of-motion exercises for the elbow and wrist, advancing to eccentric contractions and light curls by 6-12 weeks.[69] Full return to sport or heavy activity typically occurs in 3-6 months, guided by pain-free strength milestones and functional testing.[70]Precautions during training and rehabilitation include avoiding heavy loads in early phases to prevent re-injury and integrating rotator cuff and periscapular stabilization exercises to support shoulder health, as biceps function intersects with glenohumeral stability.[69]
History and terminology
Etymology
The term "biceps" originates from Latin, where it combines the prefix "bi-" meaning "two" or "double" with "ceps," a variant of "caput" meaning "head," literally translating to "two-headed." This nomenclature specifically refers to the muscle's anatomical structure, characterized by two distinct origins or heads that converge into a single tendon.[71] The word was adopted into anatomical terminology during the Renaissance to denote muscles with dual proximal attachments, distinguishing them from single-headed counterparts like the triceps brachii.[72]The term "biceps" appears in Leonardo da Vinci's anatomical drawings between 1505 and 1510, and its first documented use in printed anatomical literature is in the work of Andreas Vesalius, the Flemish anatomist, in his seminal 1543 text De humani corporis fabrica libri septem. Vesalius employed the term to describe the biceps brachii as a flexor of the elbow, emphasizing its two-headed configuration in detailed dissections and illustrations, which marked a shift from medieval reliance on ancient texts toward empirical observation.[73] This Latin term built upon earlier classical foundations; in the 2nd centuryCE, the Greek physician Galen recognized the arm's flexor muscle as two-headed in his anatomical writings, such as De usu partium, though without the precise Latin coinage that Vesalius standardized during the Renaissance revival of anatomy.[74]By the 19th century, "biceps" had transcended strictly anatomical usage, becoming a cultural symbol of physical prowess and arm strength in the burgeoning fitness and bodybuilding movements of Victorian Britain and America. This evolution coincided with the popularization of "muscular Christianity" and public displays of strength in circuses and gymnasiums, where flexed biceps represented ideals of masculinity and vigor.[75]
Grammar and nomenclature
In English, the term "biceps" functions as both a singular and plural noun, allowing constructions such as "the biceps flexes the elbow" for singular reference or "the biceps are visible during flexion" for plural, reflecting its adoption from Latin where the proper plural "bicipites" is rarely used and considered nonstandard.[76][77] The form "bicepses" occasionally appears as a plural but is less common than retaining "biceps" for multiple instances.[78]According to the International Anatomical Terminology established by the Federative International Programme for Anatomical Terminology (FIPAT) under the International Federation of Associations of Anatomists (IFAA) in 2019, the official Latin designation is musculus biceps brachii, with the English equivalent simply "biceps brachii." In medical and scientific literature, it is commonly abbreviated as "BB" for brevity, as seen in studies on muscle morphology and function.[6][79]Synonyms for the muscle include "biceps humeri" and descriptive phrases such as "two-headed muscle of the arm," emphasizing its dual origins. Historical texts occasionally employed terms like "cubital muscle" to refer to flexors in the elbow region, though this is less specific to the biceps brachii today.[80]In colloquial English, "biceps" frequently denotes the visible bulge or peak in the anterior upper arm produced during flexion, rather than the complete musculotendinous unit spanning from the shoulder to the elbow.[1] This usage highlights its cultural association with physical strength and aesthetics in fitness contexts.
Historical development
The anatomical understanding of the biceps brachii muscle originated in ancient times with Claudius Galen's descriptions around 175 AD in On the Usefulness of the Parts of the Body, where he identified it as the primary flexor of the elbow joint, emphasizing its role in arm movement; however, his accounts were constrained by limited access to human cadavers, relying primarily on dissections of animals like apes and oxen, which introduced inaccuracies in depicting human-specific structures such as the precise origins and insertions.[81]Renaissance anatomists marked a pivotal shift toward empirical observation through human dissection. Andreas Vesalius, in his seminal 1543 work De humani corporis fabrica, provided the first accurate illustrations of the biceps brachii's dual heads—the long head originating from the supraglenoid tubercle and the short head from the coracoid process—challenging Galenic errors and establishing a foundation for precise muscular anatomy based on direct examination.[82]Leonardo da Vinci had earlier recognized the biceps' role in forearm supination in his anatomical drawings from 1505-1510, though this insight was not widely disseminated until later anatomists like William Cheselden in 1713 and Jacob Winslow in 1732 rediscovered and confirmed it, with Guillaume Duchenne providing a detailed account in 1867.[81]In the 19th century, Theodor Kocher advanced clinical applications in the 1870s by pioneering surgical techniques for biceps ruptures, including tenodesis and reattachment methods via his posterolateral elbow approach, which improved outcomes for traumatic injuries and laid groundwork for modern repairs.[83]The 20th and 21st centuries brought quantitative and imaging innovations. Electromyography (EMG) studies in the 1940s, such as those by Inman and colleagues, quantified biceps activation patterns during dynamic upper limb movements, revealing peak activity in flexion and supination with measurable electrical signals that confirmed its biarticular contributions.[84] More recently, MRI advancements in the 2000s enabled three-dimensional visualization of anatomical variations, as in Blemker et al.'s 2005 finite-element modeling, which highlighted nonuniform strain distributions and intraspecific differences in head architecture and tendon paths using high-resolution scans.[85]
Comparative anatomy
In primates
In non-human primates, the biceps brachii exhibits structural variations that reflect adaptations to arboreal locomotion and suspension, contrasting with the human emphasis on forearm supination. In great apes such as chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla), the muscle features longer fascicle lengths, typically ranging from 10.5 to 20.3 cm in chimpanzees, compared to human values of approximately 15 cm, enabling greater excursion for brachiation and climbing.[86] These longer muscle bellies, combined with relatively larger overall mass (e.g., 62–191 g per head in chimpanzees), support sustained overhead arm positions during suspensory behaviors.[86] The insertion primarily occurs on the radial tuberosity, with the bicipital aponeurosis providing additional attachment to the forearmfascia, which may extend more broadly in non-human primates including toward the ulnar side.[87]Among New World monkeys (Platyrrhini), such as capuchins (Cebus apella), the biceps brachii shows a reduced short head relative to the long head, with the short head originating more proximally from the coracoid process or humerus, emphasizing the long head's role in suspension and grasping during arboreal travel.[88] This configuration results in a smaller overall physiological cross-sectional area, estimated at 5–10 cm² based on body size scaling from dissections, compared to 20–30 cm² in humans, reflecting lower force demands for rapid, leaping locomotion rather than prolonged loading.[89]Fiber composition in capuchin biceps includes a mix of slow-twitch oxidative (SO, ~40%) and fast-twitch oxidative-glycolytic (FOG, ~35%) fibers, supporting both postural maintenance and phasic contractions for foraging and tool manipulation.[90]Evolutionary shifts in the hominid lineage enhanced the biceps brachii's capacity for supination torque, facilitating precise tool use and manipulation. In early hominids, increased integration of forearm muscles, including the biceps, with the shoulder-arm module improved rotational efficiency, as evidenced by reduced musculoskeletal modularity in Homo sapiens (5 modules) compared to chimpanzees (11 modules), allowing finer control during dexterous activities.[91]Neanderthal (Homo neanderthalensis) variants demonstrate particularly robust insertions, with 126–138% greater biceps tension capacity than modern humans, inferred from proximal ulna morphology in fossils like Shanidar 3 and Kebara 2 (dated ~40,000–70,000 years ago), suggesting adaptations for high-leverage throwing and scraping tasks.[92]Functionally, non-human primates rely more heavily on the biceps brachii for shoulder flexion and stabilization during overhead postures than for isolated elbow flexion, with electromyographic patterns in rhesus monkeys (Macaca mulatta) showing preferred torque directions shifted 22° toward shoulder-extension/elbow-flexion combinations to counter gravitational loads in suspension.[93] In contrast, humans emphasize the muscle's supinatory role, particularly when the elbow is flexed, as the biceps generates peak torque in pronated-to-supinated transitions essential for tool handling, with minimal shoulder flexion contribution beyond abduction support.[93] These differences underscore the biceps' evolutionary repurposing from locomotor to manipulative functions in hominids.[91]
In other mammals
In carnivores such as dogs and cats, the biceps brachii is typically a single-headed muscle originating primarily from the coracoid process of the scapula and inserting onto the radial tuberosity of the radius, facilitating powerful elbow flexion essential for predatory behaviors like grasping and subduing prey.[94] This configuration supports rapid, forceful movements, with the muscle predominantly composed of fast-twitch glycolytic fibers to enable quick contractions and short tendons that enhance efficiency during bursts of activity.[95] Innervation is provided by the musculocutaneous nerve (C6-C7 spinal segments), with variations including an axillary loop in cats that connects to the median nerve, unlike the more straightforward branching in dogs.[96]In rodents like rats, the biceps brachii represents a scaled-down version of the muscle, with a dominant origin from the coracoid process contributing to its compact structure, alongside a smaller long head from the supraglenoid tubercle, and insertion primarily on the radius to support forelimb flexion during locomotion and burrowing.[97] Fiber composition features slow-twitch oxidative fibers restricted to deep regions for sustained activity, while fast-twitch glycolytic fibers predominate in intermediate and superficial layers for agile movements.[98] This muscle is frequently utilized in laboratory models for studying muscle regeneration and nerve repair, as demonstrated in rat brachial plexus injury experiments where biceps function recovery informs therapeutic strategies for peripheral nerve damage.Among non-equine ungulates such as deer and other ruminants, the biceps brachii exhibits an elongated form adapted to the grazing posture, with origins from the supraglenoid tubercle and coracoid process extending via a long tendon that inserts in fusion with the brachialis muscle onto the proximal radius and ulna for stable elbow flexion during quadrupedal support.[99] Its vascular supply derives from branches of the subclavian artery transitioning to the axillary artery, ensuring robust perfusion in large-bodied species.[100]Notable variations occur in aquatic mammals, where the biceps brachii is absent in extant cetaceans like whales, having been lost independently in odontocete and mysticete lineages, with other flexor muscles assuming its roles in the streamlined, paddle-like forelimbs.[101] This evolutionary reduction contrasts with its presence in terrestrial and semi-aquatic ancestors, highlighting adaptations to fully aquatic locomotion.
Functional adaptations
In mammals, the biceps brachii has undergone evolutionary shifts from primarily supporting quadrupedal locomotion in early forms to facilitating manipulative behaviors in primates. Fossil evidence indicates a notable increase in overall forelimb robusticity, including biceps size, in Homo erectus around 1.8 million years ago, correlating with enhanced tool use and bipedal efficiency.[102]In horses, the biceps brachii functions as an accessory to the deep digital flexor tendon via its lacertus fibrosus, storing elastic energy to facilitate passive protraction and stability during high-speed gaits like the gallop. This adaptation aids limb extension by countering elbow extension forces from the triceps during the stance phase, enabling efficient weight support and forward propulsion in behavioral contexts such as fleeing or racing. Rupture of the biceps tendon is rare but results in significant forelimb lameness, often confirmed via ultrasonography and bursoscopy. Electromyographic (EMG) studies reveal peak biceps activity during the early stance and late swing phases of the canter, with amplitudes significantly higher than in walking, underscoring its role in dynamic locomotor control.[103][104][105]Felines exhibit biceps adaptations suited to explosive predatory behaviors, where the muscle enables rapid elbow flexion for pouncing on prey, with the short head contributing dominantly to forearm supination and power generation during short bursts. This configuration supports vertical climbing, allowing cats to scale trees or walls by gripping and pulling with precise forelimb control, integrating with digital flexors for stability on irregular surfaces.[106][107]Aquatic mammals like seals demonstrate modified biceps brachii suited to swimming propulsion rather than terrestrial supination. In pinnipeds such as phocid seals, the muscle exhibits reduced size relative to terrestrial counterparts, becoming integrated with pectoral muscles like the pectoralis and deltoideus to drive foreflipper retraction and depression during underwater strokes. This adaptation prioritizes hydrodynamic efficiency in diving and foraging behaviors, generating lift-based thrust with minimal drag.[108]