Control exerted by ligaments

The function of the ligaments as local controllers, independent of the central nervous system, in maintaining the integrity of the joint is demonstrated by modelling the human knee in the sagittal plane, and studying its anterior-posterior motion. In addition to the ligaments, the model includes the characteristic geometry of the joint surface and some muscle groups. The connecting reaction forces at the point of contact between the tibia and the femur are considered to be constraint forces due to three different surface motions--gliding, rolling and combined gliding and rolling. It is demonstrated that the ligamentous structure maintains these holonomic and nonholonomic constraints that describe the joint motion, and that stability of the knee joint is provided mainly by ligaments. Muscular structures further stabilize and contribute to joint movement. Computer simulation of rolling movement of the knee is presented to illustrate the importance of the ligaments for joint integrity and stability.     

Comparative development of thoracic intervertebral discs and intra-articular ligaments in the human, monkey, mouse, and cat

Developmental features of thoracic intervertrebral discs and their association in the adult with other vertebral structures were investigated in four species. The human anulus fibrosus, nucleus pulposus, and intra-articular ligaments were compared to those of the fetal rhesus monkey, mouse, and kitten. Photomicrographs of transverse sections of intervertebral discs document the presence of intra-articular ligaments in fetuses of these four species. Both transverse and sagittal sections of kittens were used to identify the intercapital ligament as it differentiated from the dorsal part of the intra-articular ligament. Relatively frequent dorsal herniation of the thoracic nucleus pulposus in humans may be due to the vestigial nature of the human intra-articular ligament. Quadrupeds have well-developed intra-articular ligaments, which explains anatomically the paucity of dorsal protrusions of the nucleus pulposus into the vertebral canal in the thoracic region of the cat and mouse when compared to the human. The intra-articular ligament was closely associated with the developing prenatal mammalian intervertebral disc in the four species studied, and this relationship and its surgical importance are described.     

A global verification study of a quasi-static knee model with multi-bundle ligaments

The ligaments of the knee consist of fiber bundles with variable orientations, lengths and mechanical properties. In concept, however, these structures were too often seen as homogeneous structures, which are either stretched or slack during knee motions. In previous studies, we proposed a new structural concept of the ligaments of the knee. In this concept, the ligaments were considered as multi-bundle structures, with nonuniform mechanical properties and zero force lengths. The purpose of the present study was to verify this new concept.

For this purpose, laxity characteristics of a human knee joint were compared as measured in an experiment and predicted in a model simulation study. In the experiment, the varus-valgus and anterior-posterior laxities of a knee-joint specimen containing the ligaments and the articular surfaces only, were determined. From this knee-joint, geometric and mechanical parameters were derived to supply the parameters for a three-dimensional quasi-static knee-joint model. These parameters included (i) the three-dimensional insertion points of bundles, defined in the four major knee ligaments, (ii) the mechanical properties of these ligament, as functions of their relative insertion orientations and (iii) three-dimensional representations of the articular surfaces. With this model the experiments were simulated. If knee-model predictions and experimental results agree, then the multi-bundle ligament models are validated, at least with respect to their functional role in anterior-posterior and varus-valgus loading of the joint.

The model described the laxity characteristics in AP-translation and VV-rotation of the cadaveric knee-joint specimen reasonably well. Both display the same patterns of laxity changes during knee flexion. Only if a varus moment of 8 N m was applied and if the tibia was posteriorly loaded, did the model predict a slightly higher laxity than that measured experimentally.

From the model-experiment comparisons it was concluded that the proposed structural representations of the ligaments and their mechanical property distributions seem to be valid for studying the anterior-posterior and varus-valgus laxity characteristics of the human knee-joint.     

Cutaneous ligaments of the human hand

A topographical study concerning the cutaneous ligaments of adult as well as human fetal hands was performed. In order to be able to preserve the cutaneous ligaments in their entirety two different methods, a careful dissection and a new histological technique, have been employed. The results of these methods are compared and the detailed topography of the different cutaneous ligaments is clarified. In addition to topographical details, the functional co-operation of the ligaments especially in regard to their clinical relevance is reported.     

Somatotopy of digital nerve projections to the cuneate nucleus in the monkey

Somatotopic arrangements of cells and fibers within the dorsal columns and the dorsal column nuclei have been mapped most precisely by electrophysiological recording methods. This study uses an anatomical approach to evaluate the precision of individual digital nerve projections to the cuneate nucleus (CN) in young macaque monkeys. Digital nerves supplying about one-half the palmar skin of a digit were surgically exposed, cut, and treated with wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) on 3 successive days. After 2 additional days, animals were killed and medullas were recovered for study of serial sections reacted to display axons labeled by transganglionic transport of label. Labeled afferent fibers from each digit were found within a circumscribed columnar zone extending through the caudal CN and rostrally throughout the pars rotunda of CN. At caudal levels, diffuse projections reach the dorsal edge of the CN; more rostrally, they shift into deeper parts of the nucleus and are heaviest along its ventral and medial edges at levels near the obex. Fibers from the thumb (digit 1) project lateral (and ventral) to those from digit 2, and projections from digit 3 are medial to those from 2. Each digital projection field is closely adjacent to that from the adjacent digit. Few fibers extend to the rostral CN. Projection fields of homologous digits are quite symmetrical on the two sides. Although there do seem to be some differences in the somatotopic arrangement of digital input in macaques compared to other nonprimate mammals studied previously, these observations (precisely organized, circumscribed fields for separate digits) define a system well designed for transmission of data encoding spatial relationships.     

Somatotopy of Digital Nerve Projections to the Cuneate Nucleus in the Monkey

Somatotopic arrangements of cells and fibers within the dorsal columns and the dorsal column nuclei have been mapped most precisely by electrophysiological recording methods. This study uses an anatomical approach to evaluate the precision of individual digital nerve projections to the cuneate nucleus (CN) in young macaque monkeys. Digital nerves supplying about one-half the palmar skin of a digit were surgically exposed, cut, and treated with wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) on 3 successive days. After 2 additional days, animals were killed and medullas were recovered for study of serial sections reacted to display axons labeled by transganglionic transport of label. Labeled afferent fibers from each digit were found within a circumscribed columnar zone extending through the caudal CN and rostrally throughout the pars rotunda of CN. At caudal levels, diffuse projections reach the dorsal edge of the CN; more rostrally, they shift into deeper parts of the nucleus and are heaviest along its ventral and medial edges at levels near the obex. Fibers from the thumb (digit 1) project lateral (and ventral) to those from digit 2, and projections from digit 3 are medial to those from 2. Each digital projection field is closely adjacent to that from the adjacent digit. Few fibers extend to the rostral CN. Projection fields of homologous digits are quite symmetrical on the two sides. Although there do seem to be some differences in the somatotopic arrangement of digital input in macaques compared to other nonprimate mammals studied previously, these observations (precisely organized, circumscribed fields for separate digits) define a system well designed for transmission of data encoding spatial relationships.     

Patterning mechanisms controlling digit development

Vertebrate digits are essential structures for movement,feeding and communication.Specialized regions of the developing limb bud including the zone of polarizing activity(ZPA),the apical ectodermal ridge(AER),and the non-ridge ectoderm regulate the patterning of digits.Although a series of signaling molecules have been characterized as patterning signals from the organizing centers,the delicate cellular and molecular mechanisms that interpret how these patterning signals control the detailed digit anatomy remain unclear.Recent studies from model organisms and human hand malformations provide new insights into the mechanisms regulating this process.Here,we review the current understanding of the genetic networks governing digit morphogenesis.     

Evolution of digit identity in the three-toed Italian skink Chalcides chalcides: a new case of digit identity frame shift

SUMMARY Digit identity in the avian wing is a classical example of conflicting anatomical and embryological evidence regarding digit homology. Anatomical in conjunction with phylogenetic evidence supports the hypothesis that the three remaining digits in the bird wing are digits 1, 2, and 3. At the same time, various lines of embryological evidence support the notion that these digits develop in positions that normally produce digits 2, 3, and 4. In recent years, gene expression as well as experimental evidence was published that supports the hypothesis that this discrepancy arose from a digit identity shift in the evolution of the bird wing. A similar but less well-known controversy has been ongoing since the late 19th century regarding the identity of the digits of the three-toed Italian skink, Chalcides chalcides . Comparative anatomy identifies these digits as 1, 2, and 3, while embryological evidence suggests their derivation from embryological positions 2, 3, and 4. Here we re-examine this evidence and add gene expression data to determine the identity of the three digits of C. chalcides . The data confirm that the adult and the embryological evidence for digit identity are in conflict, and the expression of Hoxd11 suggests that digits 1, 2, and 3 develop in positions 2, 3, and 4. We conclude that in C. chalcides , and likely in its close relatives, a digit identity frame shift has occurred, similar to the one in avian evolution. This result suggests that changes in of digit identity might be a more frequent consequence of digit reduction than previously assumed.     

Birds have dinosaur wings: The molecular evidence

Within developmental biology, the digits of the wing of birds are considered on embryological grounds to be digits 2, 3 and 4. In contrast, within paleontology, wing digits are named 1, 2, 3 as a result of phylogenetic analysis of fossil taxa indicating that birds descended from theropod dinosaurs that had lost digits 4 and 5. It has been argued that the development of the wing does not support the conclusion that birds are theropods, and that birds must have descended from ancestors that had lost digits 1 and 5. Here we use highly conserved gene expression patterns in the developing limbs of mouse and chicken, including the chicken talpid(2)mutant and polydactylous Silkie breed (Silkie mutant), to aid the assessment of digital identity in the wing. Digit 1 in developing limbs does not express Hoxd12, but expresses Hoxd13. All other digits express both Hoxd12and Hoxd13. We found this signature expression pattern identifies the anteriormost digit of the wing as digit 1, in accordance with the hypothesis these digits are 1, 2 and 3, as in theropod dinosaurs. Our evidence contradicts the long-standing argument that the development of the wing does not support the hypothesis that birds are living dinosaurs.     

Grasping kinematics from the perspective of the individual digits: a modelling study

Grasping is a prototype of human motor coordination. Nevertheless, it is not known what determines the typical movement patterns of grasping. One way to approach this issue is by building models. We developed a model based on the movements of the individual digits. In our model the following objectives were taken into account for each digit: move smoothly to the preselected goal position on the object without hitting other surfaces, arrive at about the same time as the other digit and never move too far from the other digit. These objectives were implemented by regarding the tips of the digits as point masses with a spring between them, each attracted to its goal position and repelled from objects' surfaces. Their movements were damped. Using a single set of parameters, our model can reproduce a wider variety of experimental findings than any previous model of grasping. Apart from reproducing known effects (even the angles under which digits approach trapezoidal objects' surfaces, which no other model can explain), our model predicted that the increase in maximum grip aperture with object size should be greater for blocks than for cylinders. A survey of the literature shows that this is indeed how humans behave. The model can also adequately predict how single digit pointing movements are made. This supports the idea that grasping kinematics follow from the movements of the individual digits.     

Transplantation of an osteoarthrotendinous allograft with autogenous soft-tissue coverage for thumb reconstruction

A fresh-frozen thumb osteoarthrotendinous allograft and autogenous coverage were used to reconstruct a thumb. Immunosuppressants were not used. The components of the composite allograft are present and functioning 1 year post-operatively. Host cells have replaced and are replacing bone and tendinous structures. The "survival" of this osteoarthrotendinous allograft may have important implications in the treatment of patients with previous digital amputations, congenital absence of digits, and amputated digits that have failed replantation or are not replantable because of severely damaged vessels.     

Organization of connective tissue patterns by dermal fibroblasts in the regenerating axolotl limb

A set of tendons, aponeurotic sheets and retinaculae, which transduce muscle action from proximal limb levels to flexion and extension of the digits, is found in limbs of many vertebrates. This set of structures, here termed the digit tendon complex, is described for the axolotl forelimb. We show that the complex forms autonomously in muscleless axolotl limb regenerates produced from a cuff of unirradiated dermis surrounding an irradiated limb stump, and persists for up to a year after amputation. The pattern of other connective tissue structures, including the skeleton, is also normal. Fibroblast condensations that may represent sets of these cells normally associated with muscles in the extensor and flexor compartments of the carpal region also form in muscleless limbs. The results are discussed in terms of the importance of the dermis in pattern regulation, selforganization of connective tissues in general and autonomous development of the digit tendon complex in particular.     

Segmental 2:4 digit ratio. Unilateral, bilateral and hand-type differences in men

Most studies of the 2:4 digit ratio (DR) of the human hand have analyzed its possibilities for use as an indicator of the functional, psychological, and, with rare exception, morphological features of the body. Each of the functional flexor segments (each phalanx) contributes to the common variability of the total digital length. The aim of this study was to determine patterns in the relationships between each segmental digit ratio (SDR) and the overall 2:4 DR. We studied the variability of the SDRs in comparison with the general 2:4 DR of 202 young males aged 16–21 years. The length of the functional segments of the digits was measured between the flexor skin furrows on the palmar surface. The change in the SDR means that values can be ranked as follows: proximal SDR (SDR-P) > DR > distal SDR (SDR-D) > medial SDR (SDR-M). Bilateral differences are inherent in the segmental SDR-P. Most quantitative regularities inherent in the total 2:4 DR are also intrinsic to the 2:4 DR of the distal segment of the digit; in particular, this phenomenon is displayed in hands of the ulnar (male) type. Overall, the values of the distal rather than the other functional flexor segments of the second and fourth digits are similar in magnitude to the general 2:4 DR. At the same time, the greatest correlation and hence, the largest contribution to the total variability in the general DR is from the variability of the SDR-P values. Our data demonstrate the importance of various digit segment sizes in determining their overall length and the derivative ratios.     

Deformation and stress distribution of the human foot after plantar ligaments release: A cadaveric study and finite element analysis

The majority of foot deformities are related to arch collapse or instability, especially the longitudinal arch. Although the relationship between the plantar fascia and arch height has been previously investigated, the stress distribution remains unclear. The aim of this study was to explore the role of the plantar ligaments in foot arch biomechanics. We constructed a geometrical detailed three-dimensional (3-D) finite element (FE) model of the human foot and ankle from computer tomography images. The model comprised the majority of joints in the foot as well as bone segments, major ligaments, and plantar soft tissue. Release of the plantar fascia and other ligaments was simulated to evaluate the corresponding biomechanical effects on load distribution of the bony and ligamentous structures. These intrinsic ligaments of the foot arch were sectioned to simulate different pathologic situations of injury to the plantar ligaments, and to explore bone segment displacement and stress distribution. The validity of the 3-D FE model was verified by comparing results with experimentally measured data via the displacement and von Mise stress of each bone segment. Plantar fascia release decreased arch height, but did not cause total collapse of the foot arch. The longitudinal foot arch was lost when all the four major plantar ligaments were sectioned simultaneously. Plantar fascia release was compromised by increased strain applied to the plantar ligaments and intensified stress in the midfoot and metatarsal bones. Load redistribution among the centralized metatarsal bones and focal stress relief at the calcaneal insertion were predicted. The 3-D FE model indicated that plantar fascia release may provide relief of focal stress and associated heel pain. However, these operative procedures may pose a risk to arch stability and clinically may produce dorsolateral midfoot pain. The initial strategy for treating plantar fasciitis should be non-operative.     

Circumventing the polydactyly 'constraint': the mole's 'thumb'

Talpid moles across all northern continents exhibit a remarkably large, sickle-like radial sesamoid bone anterior to their five digits, always coupled with a smaller tibial sesamoid bone. A possible developmental mechanism behind this phenomenon was revealed using molecular markers during limb development in the Iberian mole (Talpa occidentalis) and a shrew (Cryptotis parva), as shrews represent the closest relatives of moles but do not show these conspicuous elements. The mole's radial sesamoid develops later than true digits, as shown by Sox9, and extends into the digit area, developing in relation to an Msx2-domain at the anterior border of the digital plate. Fgf8 expression, marking the apical ectodermal ridge, is comparable in both species. Developmental peculiarities facilitated the inclusion of the mole's radial sesamoid into the digit series; talpid moles circumvent the almost universal pentadactyly constraint by recruiting wrist sesamoids into their digital region using a novel developmental pathway and timing.     

Collateral ligaments of the distal sesamoid bone in the digit of Equus: re-evaluating midstance function

The distal forelimb of the horse has a complex array of ligaments that play a critical role in determining function of the digit and are often associated with the initiation of foot pathologies. The collateral ligaments of the distal sesamoid bone (CLDS) play an important role in digit stabilization near the end of foot contact and there is also limited evidence to suggest that the CLDS stabilize the proximal interphalangeal joint (PIPJ) during weight bearing. By virtue of their anatomical attachments where the ligaments pass dorsal to the axis of rotation of the PIPJ, it is reasonable to assume that the CLDS prevent flexion of the PIPJ during weight bearing or midstance in a moving horse. To test this functional hypothesis, forelimb specimens from three mixed-breed horses were loaded in compression in a materials testing frame. Limb loading was applied with the CLDS intact and following transection. Average PIPJ angle and metacarpophalangeal joint (MCPJ) angle at maximum load (approximately 3000 N) were calculated from angular changes of proximal and middle phalanges and the third metacarpal, which were compared between intact and transected trials. PIPJ angles were found to be the same (175 degrees) at maximum load for intact and transected trials. The proximal and middle phalanges rotated together remaining aligned, regardless of the CLDS condition. Contrary to expectation, however, the combined proximal and middle phalanges unit rotates less relative to the third metacarpal under load after transection, indicating less digit extension at the metacarpophalangeal (fetlock) joint without the influence of CLDS. Since the mechanical properties of the fetlock joint are unchanged by CLDS transection, observed proximal and middle phalanx motion is dependent on increased rotation of the distal phalanx after transection. The original hypothesis was not supported and the results suggest that at midstance the CLDS function primarily to stabilize the articulation of the middle phalanx about the distal phalanx to limit distal interphalangeal joint extension during weight bearing. Establishing the functional role of the CLDS may help to better understand the biomechanical consequences of ligament injuries and diseases of the pastern.     

Biomechanical properties of human lumbar spine ligaments.

Biomechanical properties of the six major lumbar spine ligaments were determined from 38 fresh human cadaveric subjects for direct incorporation into mathematical and finite element models. Anterior and posterior longitudinal ligaments, joint capsules, ligamentum flavum, interspinous, and supraspinous ligaments were evaluated. Using the results from in situ isolation tests, individual force-deflection responses from 132 samples were transformed with a normalization procedure into mean force-deflection properties to describe the nonlinear characteristics. Ligament responses based on the mechanical characteristics as well as anatomical considerations, were grouped into T12-L2, L2-L4, and L4-S1 levels maintaining individuality and nonlinearity. A total of 18 data curves are presented. Geometrical measurements of original length and cross-sectional area for these six major ligaments were determined using cryomicrotomy techniques. Derived parameters including failure stress and strain were computed using the strength and geometry information. These properties for the lumbar spinal ligaments which are based on identical definitions used in mechanical testing and geometrical assay will permit more realistic and consistent inputs for analytical models.