A New Interpretation of the Crocodile Forelimb Morphological Features as Adaptation to Parasagittal Quadrupedal Locomotion on the Ground

We describe the crocodile forelimb features that distinguish them from other reptiles. Reduction of the clavicle and a change in the coracoid shape seem to be another way of maintaining the efficient step length, while the antebrachium and manus transformations create peculiar oblique manus position on the ground to promote the forelimb parasagittalization.     

Architectural Properties of Sloth Forelimb Muscles (Pilosa: Bradypodidae)

Tree sloths have reduced skeletal muscle mass, and yet they are able to perform suspensory behaviors that require both strength and fatigue resistance to suspend their body mass for extended periods of time. The muscle architecture of sloths is hypothesized to be modified in ways that will enhance force production to compensate for this reduction in limb muscle mass. Our objective is to test this hypothesis by quantifying architecture properties in the forelimb musculature of the brown-throated three-toed sloth (Bradypus variegatus: N = 4). We evaluated architecture from 52 forelimb muscles by measuring muscle moment arm (r _m), muscle mass (MM), belly length (ML), fascicle length (L_F), pennation angle (θ), and physiological cross-sectional area (PCSA), and these metrics were used to estimate isometric force, joint torque, and power. Overall, the musculature becomes progressively more pennate from the extrinsic to intrinsic regions of the forelimb, and the flexors are more well developed than the extensors as predicted. However, most muscles are indicative of a mechanical design for fast joint rotational velocity instead of large joint torque (i.e., strength), although certain large, parallel-fibered shoulder (e.g., m. latissimus dorsi) and elbow (e.g., m. brachioradialis) flexors are capable of producing appreciable torques by having elongated moment arms. This type of functional tradeoff between joint rotational velocity and mechanical advantage is further exemplified by muscle gearing in Bradypus that pairs synergistic muscles with opposing L_F/r _m ratios in each functional group. These properties are suggested to facilitate the slow, controlled movements in sloths. In addition, the carpal/digital flexors have variable architectural properties, but their collective PCSA and joint torque indicates the capability for maintaining grip force and carpal stability while distributing load from the manus to the shoulder. The observed specializations provide a basis for understanding sustained suspension in sloths.     

The role of the forelimb in prey capture in the Late Triassic reptile Megalancosaurus (Diapsida,Drepanosauromorpha)

The pectoral girdle and forelimb of the Late Triassic drepanosauromorph reptile Megalancosaurus are redescribed and their function reinterpreted. The whole skeleton of this diapsid is highly specialised for arboreal life, and also the peculiarities of the shoulder girdle and forelimb were interpreted as adaptations for a limb-based locomotion using gap-bridging to move from one support to another, as in chameleons. Re-examination of the pectoral girdle and forelimb revealed the presence of clavicles fused into a furcula-like structure, a saddle-shaped glenoid and a tight connection between the radius and ulna that strengthened the forearm but hindered pronation and supination movements at that joint. The new information plus a reconstruction of the pectoral and forelimb musculature suggests that the forelimb was also specialised for grasping and raking in addition to climbing and thus prey capture may have been an important function for the forelimb. The new functional interpretation fits well with the overall body architecture of Megalancosaurus’ skeleton, suggesting that this reptile was an ambush predator that may have assumed a stable tripodal position, secured by the hooked tail and hind limbs, freeing its forelimbs to catch prey by sudden extension of the arm and firm grasping with the pincer-like digits.     

Density of muscle spindle profiles in the intrinsic forelimb muscles of the dog

The concept of parallel muscle combinations, in which spindle density is significantly higher in small muscles compared to their larger counterparts in large-small muscle combinations acting across a joint, is supported by the results of this study regardless of the joint. Analysis of the canine data as well as previously published guinea pig forelimb and human pelvic limb data revealed no significant difference in spindle density between antigravity and non-antigravity muscles. Furthermore, a gradual increase in spindle density from proximal to distal on the limb was not found, although spindle density was significantly higher in the intrinsic manus or pes muscles compared to more proximal limb muscles in all three species. The significant differences in spindle densities in parallel muscle combinations and in manus/pes versus proximal muscles are discussed relative to their possible role in the control of locomotion.     

Primary motor cortex reorganization in a long-term monkey amputee

The primary motor cortex (M1) was mapped with intracortical microstimulation (ICMS) in a 15 year-old macaque whose right upper extremity was amputated at the shoulder joint prior to 2 years of age. Movements of the right shoulder girdle and stump were evoked by ICMS throughout the left M1 upper extremity region. The size of the left M1 upper extremity region contralateral to the amputated arm was not appreciably different from the size of the right upper extremity region contralateral to the intact arm. Long stimulus trains and/or higher stimulus currents were needed to evoke detectable movements at significantly more loci in the left than in the right M1 upper extremity region. These observations would be consistent with unmasking of a high threshold representation of shoulder musculature that normally exists throughout the central core of the upper extremity region, where it underlies a lower threshold representation of the distal forelimb. Alternatively, invasion of the de-efferented distal forelimb core by surrounding shoulder representation may have occurred. Differences between the limited M1 reorganization observed in the present study and the more extensive reorganization of S1 observed in other studies may reflect fundamental differences between M1 and S1, and/or differences in the extent of de-efferentation versus deafferentation.     

Forelimb Kinematics of Rats Using XROMM,with Implications for Small Eutherians and Their Fossil Relatives

The earliest eutherian mammals were small-bodied locomotor generalists with a forelimb morphology that strongly resembles that of extant rats. Understanding the kinematics of the humerus, radius, and ulna of extant rats can inform and constrain hypotheses concerning typical posture and mobility in early eutherian forelimbs. The locomotion of Rattus norvegicus has been extensively studied, but the three-dimensional kinematics of the bones themselves remains under-explored. Here, for the first time, we use markerless XROMM (Scientific Rotoscoping) to explore the three-dimensional long bone movements in Rattus norvegicus during a normal, symmetrical gait (walking). Our data show a basic kinematic profile that agrees with previous studies on rats and other small therians: rats maintain a crouched forelimb posture throughout the step cycle, and the ulna is confined to flexion/extension in a parasagittal plane. However, our three-dimensional data illuminate long-axis rotation (LAR) movements for both the humerus and the radius for the first time. Medial LAR of the humerus throughout stance maintains an adducted elbow with a caudally-facing olecranon process, which in turn maintains a cranially-directed manus orientation (pronation). The radius also shows significant LAR correlated with manus pronation and supination. Moreover, we report that elbow flexion and manus orientation are correlated in R. norvegicus: as the elbow angle becomes more acute, manus supination increases. Our data also suggest that manus pronation and orientation in R. norvegicus rely on a divided system of labor between the ulna and radius. Given that the radius follows the flexion and extension trajectory of the ulna, it must rotate at the elbow (on the capitulum) so that during the stance phase its distal end lies medial to ulna, ensuring that the manus remains pronated while the forelimb is supporting the body. We suggest that forelimb posture and kinematics in Juramaia, Eomaia, and other basal eutherians were grossly similar to those of rats, and that humerus and radius LAR may have always played a significant role in forelimb and manus posture in small eutherian mammals.     

A behavior-based inverse kinematics algorithm to predict arm prehension postures for computer-aided ergonomic evaluation.

In this paper, the computational problem of inverse kinematics of arm prehension movements was investigated. How motions of each joint involved in arm movements can be used to control the end-effector (hand) position and orientation was first examined. It is shown that the inverse kinematics problem due to the kinematic redundancy in joint space is ill-posed only at the control of hand orientation but not at the control of hand position. Based upon this analysis, a previously proposed inverse kinematics algorithm (Wang et Verriest, 1998a) to predict arm reach postures was extended to a seven-DOF arm model to predict arm prehension postures using a separate control of hand position and orientation. The algorithm can be either in rule-based form or by optimization through appropriate choice of weight coefficients. Compared to the algebraic inverse kinematics algorithm, the proposed algorithm can handle the non-linearity of joint limits in a straightforward way. In addition, no matrix inverse calculation is needed, thus avoiding the stability and convergence problems often occurring near a singularity of the Jacobian. Since an end-effector motion-oriented method is used to describe joint movements, observed behaviors of arm movements can be easily implemented in the algorithm. The proposed algorithm provides a general frame for arm postural control and can be used as an efficient postural manipulation tool for computer-aided ergonomic evaluation.     

Avian wing proportions and flight styles: first step towards predicting the flight modes of mesozoic birds

We investigated the relationship between wing element proportions and flight mode in a dataset of living avian species to provide a framework for making basic estimates of the range of flight styles evolved by Mesozoic birds. Our results show that feather length (f(prim)) and total arm length (ta) (sum of the humerus, ulna and manus length) ratios differ significantly between four flight style groups defined and widely used for living birds and as a result are predictive for fossils. This was confirmed using multivariate ordination analyses, with four wing elements (humerus, ulna/radius, manus, primary feathers), that discriminate the four broad flight styles within living birds. Among the variables tested, manus length is closely correlated with wing size, yet is the poorest predictor for flight style, suggesting that the shape of the bones in the hand wing is most important in determining flight style. Wing bone thickness (shape) must vary with wing beat strength, with weaker forces requiring less bone. Finally, we show that by incorporating data from Mesozoic birds, multivariate ordination analyses can be used to predict the flight styles of fossils.     

Anatomy of the hand and arm in Daubentonia madagascariensis : a functional and phylogenetic outlook

The aye-aye (Daubentonia madagascariensis) is easily the most enigmatic of living primates. It sports a unique combination of derived characters, including continuously growing incisors, functional claws, the largest hand of any primate and a highly modified middle finger. The specialised middle finger is no longer used in locomotion and serves as a probe-like instrument for investigating, locating and extracting xylophagous (wood-boring) larvae as well as other food items. Its phalanges can be moved both at great speed and independently of each other. The present study reports on dissections of the forelimbs of two individuals of D. madagascariensis and one specimen each of Lemur catta and Cercopithecus cephus. Derived characters of the forelimb musculature in Daubentonia are interpreted within the context of its distinct locomotor and foraging adaptations. The primary adaptations underlying speed and mobility in the third manual digit of Daubentonia are found in the intrinsic hand musculature and notably in the arrangement of the dorsal aponeurosis. Implications for the interpretation of suggested convergences between the aye-aye, the diprotodont marsupial Dactylopsila palpator and the early Tertiary apatemyid genus Heterohyus are discussed.     

Evolution of the mammalian middle ear.

The structure and evolution of the mandible, suspensorium, and stapes of mammal-like reptiles and early mammals are examined in an attempt to determine how, why, and when in phylogeny the precursors of the mammalian tympanic bone, malleus, and incus (postdentary jaw elements and quadrate) came to function in the reception of air-borne sound. The following conclusions are reached: It is possible that at no stage in mammalian phylogeny was there a middle ear similar to that of "typical" living reptiles, with a postquadrate tympanic membrane contracted by an extrastapes. The aquamosal sulcus of cynodonts and other therapsids, usually thought to have housed a long external acoustic meatus, possibly held a depressor mandibulae muscle. In therapsids an air-filled chamber (recessus mandibularis of Westoll) extended deep to the reflected lamina and into the depression (external fossa) on the outer aspect of the angular element. A similar chamber was present in sphenacodontids but pterygoideus musculature occupied the small external fossa. The thin tissues superficial to the recessus mandibularis served as eardrum. Primitively, vibrations reached the stapes mainly via the anterior hyoid cornu, but in dicynodonts, therocephalians, and cynodants vibrations passed mainly or exclusively from mandible to quadrate to stapes and the reflected lamina was a component of the eardrum. In the therapsid phase of mammalian phylogeny, auditory adaptation was an important aspect of jaw evolution. Auditory efficiency, and sensitivity to higher sound frequencies were enhanced by diminution and loosening of the postdentary elements and quadrate, along with transference of musculature from postdentary elements to the dentary. These changes were made possible by associated modifications, including posterior expansion of the dentary. Establishment of a dentary-squamosal articulation permitted continuation of these trends, leading to the definitive mammalian condition, with no major change in auditory mechanism except that in most mammals (not monotremes) the angular, as tympanic, eventually bcame a non-vibrating structure.     

Forearm elongation in gibbons: Hypothesis and preliminary results

The characteristic long forelimbs of gibbons are a consequence of elongation of all segments (hand, forearm, and arm) but especially the forearm. While the hand is limited in size by its prehensile role, both the arm and the forearm do not seem to be directly limited in a similar manner. Why is the elongation concentrated specifically in the forearm? One hypothesis, originally applied to cursorial ungulates, relates to the need to enhance angular velocity during the swing phase at a minimum cost by distributing the heaviest forelimb loads proximally near the shoulder joint. This appears to be a plausible explanation despite complicating factors associated with competing functions of the forelimb during the support phase of arm-swinging and the fact that the animal behaviorally adjusts inertial properties of its forelimb by flexing it while reaching for the support. In order to test this hypothesis, the forelimb was modeled as a series of uniform cylinders from which the radii of gyration (k) were calculated. After converting into relative values (%k), they were compared (1) interspecifically, among two hylobatids, an orangutan, common chimpanzee, and gorilla, and (2) hypothetically, among imaginary gibbons with brachial indices extending beyond the range found in hylobatids. The results of both tests were equivocal, suggesting that forelimb mass distribution may not be the dominant factor influencing elongation of the forearms in gibbons. Even though gibbons are unique in their great dependence upon arm-swinging, efficient generation of high angular velocities during the swing phase does not appear to be the overriding function beyond others related to support, propulsion, and prehension.     

The primary feather lengths of early birds with respect to avian wing shape evolution

We examine the relationships between primary feather length (f(prim)) and total arm length (ta) (sum of humerus, ulna and manus lengths) in Mesozoic fossil birds to address one aspect of avian wing shape evolution. Analyses show that there are significant differences in the composition of the wing between the known lineages of basal birds and that mean f(prim) (relative to ta length) is significantly shorter in Archaeopteryx and enantiornithines than it is in Confuciusornithidae and in living birds. Based on outgroup comparisons with nonavian theropods that preserve forelimb primary feathers, we show that the possession of a relatively shorter f(prim) (relative to ta length) must be the primitive condition for Aves. There is also a clear phylogenetic trend in relative primary feather length throughout bird evolution: our analyses demonstrate that the f(prim)/ta ratio increases among successive lineages of Mesozoic birds towards the crown of the tree ('modern birds'; Neornithes). Variance in this ratio also coincides with the enormous evolutionary radiation at the base of Neornithes. Because the f(prim)/ta ratio is linked to flight mode and performance in living birds, further comparisons of wing proportions among Mesozoic avians will prove informative and certainly imply that the aerial locomotion of the Early Cretaceous Confuciusornis was very different to other extinct and living birds.     

东北小鲵中枢神经系统形态学与组织学初步研究

本文应用脊椎动物神经标本制作法和HE染色法,对东北小鲵中枢神经系统的外部形态和组织学结构进行了初步研究,描述了东北小鲵神经系统形态和组织学结构的特点,并与无尾两栖类和爬行类相对比,探讨了有尾两栖类的进化地位。结果表明:与无尾两栖类(如蛙)相比,东北小鲵中枢神经系统中,大脑半球较小,结构较为原始,小脑结构简单,是两栖类中较为原始的类群。此外,东北小鲵开始具有了臂神经丛和骶神经丛,但没有爬行类的发达,可作为两栖类向爬行类进化的证据之一。     

On the musculature of the forelimb of the crab-eating monkey

In this paper, the musculature of the forelimb of the crab-eating monkey was described by anatomical observation of twenty limbs and compared with that of a few other primates, especially of man andMacaca. As for the musculature of the forelimb, that of the crab-eating monkey was naturally very similar to other species of theMacaca group, except for slight differences of individual muscles.     

The interplay between speed,kinetics, and hand postures during primate terrestrial locomotion

Nonprimate terrestrial mammals may use digitigrade postures to help moderate distal limb joint moments and metapodial stresses that may arise during high‐speed locomotion with high‐ground reaction forces (GRF). This study evaluates the relationships between speed, GRFs, and distal forelimb kinematics in order to evaluate if primates also adopt digitigrade hand postures during terrestrial locomotion for these same reasons. Three cercopithecine monkey species (Papio anubis, Macaca mulatta, Erythrocebus patas) were videotaped moving unrestrained along a horizontal runway instrumented with a force platform. Three‐dimensional forelimb kinematics and GRFs were measured when the vertical force component reached its peak. Hand posture was measured as the angle between the metacarpal segment and the ground (MGA). As predicted, digitigrade hand postures (larger MGA) are associated with shorter GRF moment arms and lower wrist joint moments. Contrary to expectations, individuals used more palmigrade‐like (i.e. less digitigrade) hand postures (smaller MGA) when the forelimb was subjected to higher forces (at faster speeds) resulting in potentially larger wrist joint moments. Accordingly, these primates may not use their ability to alter their hand postures to reduce rising joint moments at faster speeds. Digitigrady at slow speeds may improve the mechanical advantage of antigravity muscles crossing the wrist joint. At faster speeds, greater palmigrady is likely caused by joint collapse, but this posture may be suited to distribute higher GRFs over a larger surface area to lower stresses throughout the hand. Thus, a digitigrade hand posture is not a cursorial (i.e. high speed) adaptation in primates and differs from that of other mammals. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

The skull and the palaeoecological significance of Labidosaurus hamatus, a captorhinid reptile from the Lower Permian of Texas

The cranial skeleton of the large captorhinid reptile Labidosaurus hamatus , known only from the Lower Permian of Texas, is described on the basis of new, undescribed specimens. Labidosaurus is distinguished from other captorhinids by the more extreme sloping of the ventral (alveolar) margin of the premaxilla, a low dorsum sellae of the parabasisphenoid, a reduced prootic, a narrow stapes, and a relatively small foramen intermandibularis medius. Despite the presence of a single row of teeth in each jaw, the skull of Labidosaurus resembles most closely those of moradisaurines, the large multiple-tooth-rowed captorhinids of the latest Early and Middle Permian. A phylogenetic analysis confirms that the single-tooth-rowed L. hamatus is related most closely to moradisaurines within Captorhinidae, a relationship that supports the hypothesis of a diphyletic origin for multiple rows of marginal teeth in captorhinids (in the genus Captorhinus and in the clade Moradisaurinae). In view of the close relationship between L. hamatus and moradisaurines, which are regarded to have been herbivorous, L. hamatus is a critical taxon for studies of the evolution of herbivory in early tetrapods. L. hamatus shares several trademark features of herbivorous adaptation with moradisaurines, which suggest that this captorhinid species was omnivorous. As such, it represents a transitional taxon between faunivorous basal reptiles and the herbivorous moradisaurines.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 237–262.     

Electromyographic activity in the Rhesus monkey forelimb muscles during a goal directed movement and locomotion before, during and after spaceflight

The aim of the present study was to analyse the effects of microgravity on i) the achievement of goal-directed arm movements and ii) the quadrupedal non-human primate locomotion. A reaching movement in weightlessness would require less muscle contraction since there is no need to oppose gravity. Consequently the electromyographic (EMG) activity of the monkey forelimb muscles should be changed during or after spaceflight. EMG activity of the biceps and triceps muscles during goal-directed arm movements were studied in Rhesus monkeys before, during and after 14 days of spaceflight and flight simulation at normal gravity. The EMG activity was also recorded during treadmill locomotion before and after spaceflight. When performing arm motor tasks, the delay values of the EMG bursts were unchanged during the flight. On the contrary, mean EMG was significantly decreased during the flight comparatively to the pre- and post-flight values, which were very similar. Compared with flight animals, the control ground monkey showed no change in the burst durations and mean EMG. After spaceflight, quadrupedal locomotion was modified. The animals had some difficulty in moving, and abnormal steps were numerous. The integrated area of triceps bursts was increased for the stance phase during locomotion. Taken together these data showed that spaceflight induces a dual adaptative process: first, the discharge of the motor pools of the forelimb musculature was modified during exposure to microgravity, and then upon return to Earth, monkeys changed their new motor strategy and re-adapt to normal gravity.     

东北小鲵中枢神经系统形态学与组织学初步研究（图版Ⅵ，上）

本文应用脊椎动物神经标本制作法和HE染色法,对东北小鲵中枢神经系统的外部形态和组织学结构进行了初步研究,描述了东北小鲵神经系统形态和组织学结构的特点,并与无尾两栖类和爬行类相对比,探讨了有尾两栖类的进化地位。结果表明：与无尾两栖类（如蛙）相比,东北小鲵中枢神经系统中,大脑半球较小,结构较为原始,小脑结构简单,是两栖类中较为原始的类群。此外,东北小鲵开始具有了臂神经丛和骶神经丛,但没有爬行类的发达,可作为两柄类向爬行类进化的证据之一。     

Cyclic coordinate descent: A robotics algorithm for protein loop closure

In protein structure prediction, it is often the case that a protein segment must be adjusted to connect two fixed segments. This occurs during loop structure prediction in homology modeling as well as in ab initio structure prediction. Several algorithms for this purpose are based on the inverse Jacobian of the distance constraints with respect to dihedral angle degrees of freedom. These algorithms are sometimes unstable and fail to converge. We present an algorithm developed originally for inverse kinematics applications in robotics. In robotics, an end effector in the form of a robot hand must reach for an object in space by altering adjustable joint angles and arm lengths. In loop prediction, dihedral angles must be adjusted to move the C-terminal residue of a segment to superimpose on a fixed anchor residue in the protein structure. The algorithm, referred to as cyclic coordinate descent or CCD, involves adjusting one dihedral angle at a time to minimize the sum of the squared distances between three backbone atoms of the moving C-terminal anchor and the corresponding atoms in the fixed C-terminal anchor. The result is an equation in one variable for the proposed change in each dihedral. The algorithm proceeds iteratively through all of the adjustable dihedral angles from the N-terminal to the C-terminal end of the loop. CCD is suitable as a component of loop prediction methods that generate large numbers of trial structures. It succeeds in closing loops in a large test set 99.79% of the time, and fails occasionally only for short, highly extended loops. It is very fast, closing loops of length 8 in 0.037 sec on average.     

Ichnology of an Extant Belly-Dragging Lizard—Analogies to Early Reptile Locomotion?

A recently described Erpetopus trackway bearing unusual claw and belly-drag marks ostensibly indicates an obligatory sprawled posture and belly-walk in the locomotion of small captorhinids. Here, the ichnology of the blue-tongued skink (Tiliqua scincoides) is investigated in order to identify features of a trackway produced by a lizard in continuous belly-walk. Comparisons between T. scincoides and Erpetopus tracks tested whether the locomotory pattern observed for T. scincoides resembles that of small captorhinid track makers. Characteristic features of the T. scincoides track include: (1) a belly-dragging mark, (2) claw scratch marks produced during the early stance phase, and (3) claw drag marks produced by the forelimb during the swing phase. Trackway parameters did not correlate with track maker velocity, rendering inference of velocity for belly-dragging track makers problematic. This result was probably caused by increased substrate influence on locomotor speed because of belly contact with the ground. The track characteristics of T. scincoides match those recently described for Erpetopus and thus corroborate the notion of a similar pattern of locomotion for small captorhinids.