Cross-sectional changes of the distal carpal tunnel with simulated carpal bone rotation.

This study simulated the cross-sectional changes in the distal carpal tunnel resulting from inward rotations of the hamate and trapezium. Rotations which decreased the carpal arch width, increased the carpal arch area. For example, simultaneous rotation of 5 degrees around the hamate and trapezium centroids decreased the carpal arch width by 1.69 ± 0.17 mm and increased the carpal arch area by 6.83 ± 0.68 mm2. Although the bone arch area decreased, decompression of the median nerve would likely occur due to the adjacent location of the nerve near the transverse carpal ligament.

Morphometric analysis of the intercarpal ligaments of the equine proximal carpal bones during simulated flexion and extension of cadaver limbs.

Despite many reported cases of carpal lameness associated with intercarpal ligament injuries in horses, the morphometry, movement pattern and general intrinsic biomechanics of the carpus are largely unknown. Using osteoligamentous preparation of the carpus prepared from 14 equine cadaver forelimbs (aged 9.62 ± 4.25 years), locomotory simulations of flexion and extension movements of the carpal joint were carried out to observed carpal biomechanics and, thereafter, the limbs were further dissected to obtain morphometric measurements of the medial and lateral collateral ligaments (MLC and LCL); medial and lateral palmar intercarpal ligaments (MPICL and LPICL); intercarpal ligaments between radial (Cr) and intermediate (Ci) carpal bones (Cr-Ci ICL); and intercarpal ligaments between Ci and ulnar (Cu) carpal bones (Ci-Cu ICL). The Cr, Ci, Cu and Ca are held together by a series of intercarpal ligaments and move in unison lateropalmarly during flexion, and mediodorsally during extension with a distinguishable proximo-distal sliding movement (gliding) of Cr and Ci against each other during movement. The mean length of MCL (108.82 ± 9.64 mm) was significantly longer (p = 0.042) than LCL (104.43 ± 7.65 mm). The Cr-Ci ICL has a dorsopalmar depth of 37.58 ± 4.14 mm and a midpoint width of 12.05 ± 3.09 mm and its fibres ran diagonally from the medial side of the Ci in a proximo-palmar disto-dorsal direction (i.e. palmarodistally) to the lateral side of the Cr. The specialized movement of the Cr-Ci ICL, which appeared to be further facilitated by a longer MCL suggest a biomechanical function by which carpal damage may be minimized in the equine carpus.

Kinematics of the Feline Antebrachiocarpal Joint from Supination to Pronation.

OBJECTIVE: Cats rely on their forelimb mobility for everyday activities including climbing and grooming. Supination and pronation of the forelimb in cats are considered to primarily involve the antebrachium, rather than the carpus. Therefore, our null hypothesis was that there would be no movement of the carpal bones (radial carpal bone, ulnar carpal bone and accessory carpal bone) relative to the ulna during supination and pronation. STUDY DESIGN: Eight feline cadaveric forelimbs were rotated from supination to pronation in a jig and computed tomography was performed in the neutral, supinated and pronated positions. The individual carpal bones were segmented from computed tomography images of the supinated and pronated scans in each of the eight specimens. A feline ulna coordinate system was established and used to quantify the translations and rotations between bones of the proximal carpal row and antebrachium. RESULTS: After the carpus was rotated from the initial supinated position into pronation, there was significant translation (x, y and z axes) and rotation (x and y axes) of the proximal row of carpal bones based on absolute magnitude values. Given the differences in translations and rotations of the proximal row of carpal bones, our null hypothesis was rejected. CONCLUSION: The proximal row of carpal bones translate and rotate independently from the ulna in the cat during pronation of the antebrachium. This may have future implications in the diagnosis and management of feline carpal injuries involving the antebrachiocarpal joint.

Time of appearance of ossification centers in carpal bones. A radiological retrospective study on Saudi children.

OBJECTIVES: To find reference data for the time of appearance of ossification centers in carpal bones and the lower ends of the radius and ulna in the Saudi population. In addition, to check the sequence of appearance of carpal bones and the relation of this sequence to the appearance of distal epiphyses of the radius and ulna. Methods: A retrospective radiological study was carried out between 2012 to 2020 at King Fahad Hospital of the University, Al-Khobar, Saudi Arabia. A sample of 279 hand/wrist plain radiographs of Saudi children was analyzed. RESULTS: The first bones at the wrist region to appear in Saudi children are the capitate, hamate, and distal epiphysis of the radius, and these appear during the first year of life. The other bones develop subsequently at yearly intervals, and the last one to appear is the pisiform, which arises at the end of the first decade of life. CONCLUSION: The sequence of appearance of carpal bones in the Saudi population is similar to what is described in the literature. However, the time of appearance of some of these bones is earlier than that in other populations.

Real-time three-dimensional MRI for the assessment of dynamic carpal instability.

BACKGROUND: Carpal instability is defined as a condition where wrist motion and/or loading creates mechanical dysfunction, resulting in weakness, pain and decreased function. When conventional methods do not identify the instability patterns, yet clinical signs of instability exist, the diagnosis of dynamic instability is often suggested to describe carpal derangement manifested only during the wrist's active motion or stress. We addressed the question: can advanced MRI techniques provide quantitative means to evaluate dynamic carpal instability and supplement standard static MRI acquisition? Our objectives were to (i) develop a real-time, three-dimensional MRI method to image the carpal joints during their active, uninterrupted motion; and (ii) demonstrate feasibility of the method for assessing metrics relevant to dynamic carpal instability, thus overcoming limitations of standard MRI. METHODS: Twenty wrists (bilateral wrists of ten healthy participants) were scanned during radial-ulnar deviation and clenched-fist maneuvers. Images resulting from two real-time MRI pulse sequences, four sparse data-acquisition schemes, and three constrained image reconstruction techniques were compared. Image quality was assessed via blinded scoring by three radiologists and quantitative imaging metrics. RESULTS: Real-time MRI data-acquisition employing sparse radial sampling with a gradient-recalled-echo acquisition and constrained iterative reconstruction appeared to provide a practical tradeoff between imaging speed (temporal resolution up to 135 ms per slice) and image quality. The method effectively reduced streaking artifacts arising from data undersampling and enabled the derivation of quantitative measures pertinent to evaluating dynamic carpal instability. CONCLUSION: This study demonstrates that real-time, three-dimensional MRI of the moving wrist is feasible and may be useful for the evaluation of dynamic carpal instability.

Predicting Carpal Bone Kinematics Using an Expanded Digital Database of Wrist Carpal Bone Anatomy and Kinematics.

The wrist can be considered a 2 degrees-of-freedom joint with all movements reflecting the combination of flexion-extension and radial-ulnar deviation. Wrist motions are accomplished by the kinematic reduction of the 42 degrees-of-freedom of the individual carpal bones. While previous studies have demonstrated the minimal motion of the scaphoid and lunate as the wrist moves along the dart-thrower's path or small relative motion between hamate-capitate-trapezoid, an understanding of the kinematics of the complete carpus across all wrist motions remains lacking. To address this, we assembled an open-source database of in vivo carpal motions and developed mathematical models of the carpal kinematics as a function of wrist motion. Quadratic surfaces were trained for each of the 42-carpal bone degrees-of-freedom and the goodness of fits were evaluated. Using the models, paths of wrist motion that generated minimal carpal rotations or translations were determined. Model predictions were best for flexion-extension, radial-ulnar deviation, and volar-dorsal translations for all carpal bones with R 2 > 0.8, while the estimates were least effective for supination-pronation with R 2 < 0.6. The wrist path of motion's analysis indicated that the distal row of carpal bones moves rigidly together (<3° motion), along the anatomical axis of wrist motion, while the bones in the proximal row undergo minimal motion when the wrist moves in a path oblique to the main axes. The open-source dataset along with its graphical user interface and mathematical models should facilitate clinical visualization and enable new studies of carpal kinematics and function. © 2019 Orthopaedic Research Society. Published by Wiley Periodicals, Inc. J Orthop Res 37:2661-2670, 2019.

Maximum loading of carpal bones during movements: a finite element study.

BACKGROUND: Maximum stresses show critical points on an object because failure may start from the area close to maximum stress points. However, there appears no study on maximum loading points of carpal bones. PURPOSE: To clarify the loading pattern of each carpal bone during wrist movements. METHODS: A finite element wrist model was designed using a three-dimensional reconstruction of computed tomographic images from the distal end of radius and ulna to the proximal third of metacarpals. Loading was performed in neutral, 45° of flexion and extension, 5° of radial and 25° of ulnar deviation, and maximum loading points were plotted. RESULTS: In each position except for extension, minimum loads were carried by triquetrum, while minimum loads were carried by capitatum in extension. Maximum loads were carried by trapezium in neutral and ulnar deviation and flexion but by scaphoideum in radial deviation and extension. CONCLUSION: Studies on maximum loading of each bone are a new approach and may help to improve the knowledge on wrist mechanics.

Finite element analysis for transverse carpal ligament tensile strain and carpal arch area.

Mechanics of carpal tunnel soft tissue, such as fat, muscle and transverse carpal ligament (TCL), around the median nerve may render the median nerve vulnerable to compression neuropathy. The purpose of this study was to understand the roles of carpal tunnel soft tissue mechanical properties and intratunnel pressure on the TCL tensile strain and carpal arch area (CAA) using finite element analysis (FEA). Manual segmentation of the thenar muscles, skin, fat, TCL, hamate bone, and trapezium bone in the transverse plane at distal carpal tunnel were obtained from B-mode ultrasound images of one cadaveric hand. Sensitivity analyses were conducted to examine the dependence of TCL tensile strain and CAA on TCL elastic modulus (0.125-10 MPa volar-dorsally; 1.375-110 MPa transversely), skin-fat and thenar muscle initial shear modulus (1.6-160 kPa for skin-fat; 0.425-42.5 kPa for muscle), and intratunnel pressure (60-480 mmHg). Predictions of TCL tensile strain under different intratunnel pressures were validated with the experimental data obtained on the same cadaveric hand. Results showed that skin, fat and muscles had little effect on the TCL tensile strain and CAA changes. However, TCL tensile strain and CAA increased with decreased elastic modulus of TCL and increased intratunnel pressure. The TCL tensile strain and CAA increased linearly with increased pressure while increased exponentially with decreased elastic modulus of TCL. Softening the TCL by decreasing the elastic modulus may be an alternative clinical approach to carpal tunnel expansion to accommodate elevated intratunnel pressure and alleviate median nerve compression neuropathy.

Kinematics of the anthropoid os centrale and the functional consequences of scaphoid-centrale fusion in African apes and hominins.

In most primates, the os centrale is interposed between the scaphoid, trapezoid, trapezium, and head of the capitate, thus constituting a component of the wrist's midcarpal complex. Scaphoid-centrale fusion is among the clearest morphological synapomorphies of African apes and hominins. Although it might facilitate knuckle-walking by increasing the rigidity and stability of the radial side of the wrist, the exact functional significance of scaphoid-centrale fusion is unclear. If fusion acts to produce a more rigid radial wrist that stabilizes the hand and limits shearing stresses, then in taxa with a free centrale, it should anchor ligaments that check extension and radial deviation, but exhibit motion independent of the scaphoid. Moreover, because the centrale sits between the scaphoid and capitate (a major stabilizing articulation), scaphoid-centrale mobility should correlate with scaphocapitate mobility in extension and radial deviation. To test these hypotheses, the centrale's ligamentous binding was investigated via dissection in Pongo and Papio, and the kinematics of the centrale were quantified in a cadaveric sample of anthropoids (Pongo sp., Ateles geoffroyi, Colobus guereza, Macaca mulatta, and Papio anubis) using a computed-tomography-based method to track wrist-bone motion. Results indicate that the centrale rotates freely relative to the scaphoid in all taxa. However, centrale mobility is only correlated with scaphocapitate mobility during extension in Pongo-possibly due to differences in overall wrist configuration between apes and monkeys. If an extant ape-like wrist characterized early ancestors of African apes and hominins, then scaphoid-centrale fusion would have increased midcarpal rigidity in extension relative to the primitive condition. Although biomechanically consistent with a knuckle-walking hominin ancestor, this assumes that the trait evolved specifically for that biological role, which must be squared with contradictory interpretations of extant and fossil hominoid morphology. Regardless of its original adaptive significance, scaphoid-centrale fusion likely presented a constraint on early hominin midcarpal mobility.

Subject-specific finite element analysis of the carpal tunnel cross-sectional to examine tunnel area changes in response to carpal arch loading.

BACKGROUND: Manipulating the carpal arch width (i.e. distance between hamate and trapezium bones) has been suggested as a means to increase carpal tunnel cross-sectional area and alleviate median nerve compression. The purpose of this study was to develop a finite element model of the carpal tunnel and to determine an optimal force direction to maximize area. METHODS: A planar geometric model of carpal bones at hamate level was reconstructed from MRI with inter-carpal joint spaces filled with a linear elastic surrogate tissue. Experimental data with discrete carpal tunnel pressures (50, 100, 150, and 200mmHg) and corresponding carpal bone movements were used to obtain material property of surrogate tissue by inverse finite element analysis. The resulting model was used to simulate changes of carpal arch widths and areas with directional variations of a unit force applied at the hook of hamate. FINDINGS: Inverse finite element model predicted the experimental area data within 1.5% error. Simulation of force applications showed that carpal arch width and area were dependent on the direction of force application, and minimal arch width and maximal area occurred at 138° (i.e. volar-radial direction) with respect to the hamate-to-trapezium axis. At this force direction, the width changed to 24.4mm from its initial 25.1mm (3% decrease), and the area changed to 301.6mm2 from 290.3mm2 (4% increase). INTERPRETATION: The findings of the current study guide biomechanical manipulation to gain tunnel area increase, potentially helping reduce carpal tunnel pressure and relieve symptoms of compression median neuropathy.

Decoupling the Wrist: A Cadaveric Experiment Examining Wrist Kinematics Following Midcarpal Fusion and Scaphoid Excision.

At the wrist, kinematic coupling (the relationship between flexion-extension and radial-ulnar deviation) facilitates function. Although the midcarpal joint is critical for kinematic coupling, many surgeries, such as 4-corner fusion (4CF) and scaphoidexcision 4-corner fusion (SE4CF), modify the midcarpal joint. This study examines how 4CF and SE4CF influence kinematic coupling by quantifying wrist axes of rotation. Wrist axes of rotation were quantified in 8 cadaveric specimens using an optimization algorithm, which fit a 2-revolute joint model to experimental data. In each specimen, data measuring the motion of the third metacarpal relative to the radius was collected for 3 conditions (nonimpaired, 4CF, SE4CF). The calculated axes of rotation were compared using spherical statistics. The angle between the axes of rotation was used to assess coupling, as the nonimpaired wrist has skew axes (ie, angle between axes approximately 60°). Following 4CF and SE4CF, the axes are closer to orthogonal than those of the nonimpaired wrist. The mean angle (±95% confidence interval) between the axes was 92.6° ± 25.2° and 99.8° ± 22.0° for 4CF and SE4CF, respectively. The axes of rotation defined in this study can be used to define joint models, which will facilitate more accurate computational and experimental studies of these procedures.

Locomotor Hand Postures, Carpal Kinematics During Wrist Extension, and Associated Morphology in Anthropoid Primates.

The biomechanics of wrist extension (or dorsiflexion) are important for understanding functional adaptation of the primate hand because extension mobility varies with habitual locomotor hand posture and facilitates certain manipulative tasks. Here, intercarpal kinematics are employed to investigate mechanisms underlying wrist extension in a sample of anthropoids representing various arboreal and terrestrial locomotor modes. Carpal kinematics are studied using computed-tomography of cadaveric forelimbs, and these data are combined with a morphometric analysis of biomechanically-informative anatomical features. The results indicate that stiff-wristed knuckle-walking chimpanzees and digitigrade baboons are characterized by low ranges of motion (ROMs) at the radiocarpal and midcarpal complexes. Palmigrade-capable monkeys have high extension ROMs at both the radiocarpus and midcarpus, while palmigrade-capable orangutans achieve wrist extension through moderate radiocarpal ROMs and high midcarpal ROMs. Morphometrics demonstrate that a more projecting dorsal ridge of the distal radius corresponds with low-to-moderate radioscaphoid mobility in the apes, but that baboons resemble palmigrade-capable monkeys in this metric. Thus, the dorsal ridge of the radius alone is not a good indicator of wrist mobility and hand posture. However, the extent of the lunate's articular arc on the dorsum of the capitate head is correlated with midcarpal mobility across taxa. These findings suggest that although a precise relationship between wrist extension ROM and morphology is difficult to define, the presence of a pronounced dorsal ridge combined with an abbreviated dorsal capitate articular arc reflects limited overall dorsiflexion with attendant constraints on the adoption of palmigrade hand postures. Anat Rec, 300:382-401, 2017. © 2016 Wiley Periodicals, Inc.

Carpal Kinematics and Kinetics.

The complex interaction of the carpal bones, their intrinsic and extrinsic ligaments, and the forces in the normal wrist continue to be studied. Factors that influence kinematics, such as carpal bone morphology and clinical laxity, continue to be identified. As imaging technology improves, so does our ability to better understand and identify these factors. In this review, we describe advances in our understanding of carpal kinematics and kinetics. We use scapholunate ligament tears as an example of the disconnect that exists between our knowledge of carpal instability and limitations in current reconstruction techniques.

Quantifying thumb opposition kinematics using dynamic computed tomography.

Current motion capture techniques all have shortcomings when applied to the 3D quantitative evaluation of thumb base motion. Dynamic CT might overcome these shortcomings but, so far, robustness of this technique in more than one specimen has not yet been demonstrated. The aim of the current study is to further evaluate the use of dynamic CT for quantification of thumb motion in a larger cadaveric study using a protocol which is feasible in a clinical context. A dynamic CT scan was acquired from six cadaveric human forearms, while a motion simulator imposed thumb opposition. After image acquisition and segmentation, carpal bone motion was quantified using helical axes. To enable comparisons between specimens, intersection points of the instantaneous helical axis with an anatomically defined plane were determined. Precision of the dynamic CT method, measured as variation in distances between silicon nitride beads between frames of a dynamic scan, was 0.43mm (+/-0.09mm) when fixed to the skin and 0.13mm (+/-0.04mm) when embedded into the bone. Absolute deviation between known and measured distances were not larger than 0.34mm. We could demonstrate and quantify that thumb opposition is associated with motion at the trapeziometacarpal and scaphotrapezotrapezoidal joints. High consistency in motion patterns between specimen were found, while the radiation dose was limited. We conclude that dynamic CT can be used to visualize and quantify 3D thumb kinematics, making it a promising method to explore kinematics in vivo.

The Functional Anatomy of the Carpometacarpal Complex in Anthropoids and Its Implications for the Evolution of the Hominoid Hand.

Previously, we described several features of the carpometacarpal joints in extant large-bodied apes that are likely adaptations to the functional demands of vertical climbing and suspension. We observed that all hominids, including modern humans and the 4.4-million-year-old hominid Ardipithecus ramidus, lacked these features. Here, we assess the uniqueness of these features in a large sample of monkey, ape, and human hands. These new data provide additional insights into the functional adaptations and evolution of the anthropoid hand. Our survey highlights a series of anatomical adaptations that restrict motion between the second and third metacarpals (MC2 and MC3) and their associated carpals in extant apes, achieved via joint reorganization and novel energy dissipation mechanisms. Their hamate-MC4 and -MC5 joint surface morphologies suggest limited mobility, at least in Pan. Gibbons and spider monkeys have several characters (angled MC3, complex capitate-MC3 joint topography, variably present capitate-MC3 ligaments) that suggest functional convergence in response to suspensory locomotion. Baboons have carpometacarpal morphology suggesting flexion/extension at these joints beyond that observed in most other Old World monkeys, probably as an energy dissipating mechanism minimizing collision forces during terrestrial locomotion. All hominids lack these specializations of the extant great apes, suggesting that vertical climbing was never a central feature of our ancestral locomotor repertoire. Furthermore, the reinforced carpometacarpus of vertically climbing African apes was likely appropriated for knuckle-walking in concert with other novel potential energy dissipating mechanisms. The most parsimonious explanation of the structural similarity of these carpometacarpal specializations in great apes is that they evolved independently.

Functional kinematics of the wrist.

The purpose of this article is to review past and present concepts concerning functional kinematics of the healthy and injured wrist. To provide a context for students of the wrist, we describe the progression of techniques for measuring carpal kinematics over the past century and discuss how this has influenced today's understanding of functional kinematics. Next, we provide an overview of recent developments and highlight the clinical relevance of these findings. We use these findings and recent evidence that supports the importance of coupled motion in early rehabilitation of radiocarpal injuries to develop the argument that coupled motion during functional activities is a clinically relevant outcome; therefore, clinicians should develop a framework for its dynamic assessment. This should enable a tailored and individualized approach to the treatment of carpal injuries.

Influence of Exercise and Intra-articular Site on Canals in Articular Calcified Cartilage of Equine Third Carpal Bones.

The third carpal bone (C3) responds to exercise by adaptive modeling of bone and articular calcified cartilage along the dorsal load path. Canals penetrating articular calcified cartilage, thought to contain vascular tissue, are reported in numerous species. Their significance remains unclear. Our objective was to determine if the number of canals was significantly different in strenuously exercised and control young horses and in a site of intermittent high loading compared to sites sustaining lower habitual loads. Volumetric bone mineral density in the radial facet of C3 of strenuously exercised and gently exercised (control) 19-month-old thoroughbred horses (n= 6/group) was determined by peripheral quantitative computed tomography. The hyaline cartilage was corroded to expose the surface of articular calcified cartilage. The number of canals penetrating the articular calcified cartilage surface in en face scanning electron microscopy images was compared in 4 regions. Volumetric bone mineral density of C3 was significantly greater (P= .004) in strenuously exercised horses. There were 2 morphologically distinct groups of canals and significantly fewer (P= .006) large canals in the dorsal than in the palmar aspect of C3 in control but not in exercised horses. Roughly circular depressions in the articular calcified cartilage surface around apparently forming canals were visible in some samples and have not been previously described in the literature. The canals may be evidence of chondroclastic activity reaching the interface of hyaline and calcified cartilage. Further work is needed to elucidate the relationships between presence of canals and the responses to exercise and to joint disease.

The effects of partial carpal fusions on wrist range of motion.

UNLABELLED: The objective of this investigation was to evaluate the effects of different partial wrist fusions on wrist motion. A total of 20 cadaveric wrists were tested in the intact state and after undergoing either a four-corner fusion or 2- and 3-bone fusion. The moment-rotation behaviour was measured in 24 directions of wrist motion about the forearm axis. The 2- and 3-bone fusion groups demonstrated increased radial deviation and pure flexion. Pure flexion was decreased in the four-corner fusion group. Radial extension and pure extension were decreased in all treatments compared with normal range of motion. Increasing the number of carpal bones within the fusion construct did not alter the functional axis of the wrist. Essentially equivalent motion is possible with 2-bone, 3-bone and four-corner fusions, with the exceptions of pure flexion and radial deviation. This data may influence surgeons when choosing between treatment methods. LEVEL OF EVIDENCE: N/A.

The effect of lunate morphology on the 3-dimensional kinematics of the carpus.

PURPOSE: To assess carpal kinematics in various ranges of motion in 3 dimensions with respect to lunate morphology. METHODS: Eight cadaveric wrists (4 type I lunates, 4 type II lunates) were mounted into a customized platform that allowed controlled motion with 6 degrees of freedom. The wrists were moved through flexion-extension (15°-15°) and radioulnar deviation (RUD; 20°-20°). The relative motion of the radius, carpus, and third metacarpal were recorded using optical motion capture methods. RESULTS: Clear patterns of carpal motion were identified. Significantly greater motion occurred at the radiocarpal joint during flexion-extension of type I wrist than a type II wrist. The relative contributions of the midcarpal and radiocarpal articulations to movement of the wrist differed between the radial, the central, and the ulnar columns. During wrist flexion and extension, these contributions were determined by the lunate morphology, whereas during RUD, they were determined by the direction of wrist motion. The midcarpal articulations were relatively restricted during flexion and extension of a type II wrist. However, during RUD, the midcarpal joint of the central column became the dominant articulation. CONCLUSIONS: This study describes the effect of lunate morphology on 3-dimensional carpal kinematics during wrist flexion and extension. Despite the limited size of the motion arcs tested, the results represent an advance on the current understanding of this topic. CLINICAL RELEVANCE: Differences in carpal kinematics may explain the effect of lunate morphology on pathological changes within the carpus. Differences in carpal kinematics due to lunate morphology may have implications for the management of certain wrist conditions.

Homologies and homeotic transformation of the theropod 'semilunate' carpal.

The homology of the 'semilunate' carpal, an important structure linking non-avian and avian dinosaurs, has been controversial. Here we describe the morphology of some theropod wrists, demonstrating that the 'semilunate' carpal is not formed by the same carpal elements in all theropods possessing this feature and that the involvement of the lateralmost distal carpal in forming the 'semilunate' carpal of birds is an inheritance from their non-avian theropod ancestors. Optimization of relevant morphological features indicates that these features evolved in an incremental way and the 'semilunate' structure underwent a lateral shift in position during theropod evolution, possibly as a result of selection for foldable wings in birds and their close theropod relatives. We propose that homeotic transformation was involved in the evolution of the 'semilunate' carpal. In combination with developmental data on avian wing digits, this suggests that homeosis played a significant role in theropod hand evolution in general.