Human ankle joint movements during walking are probably not determined by talar morphology.

Knowledge about the orientation of a representative ankle joint axis is limited to studies of tarsal morphology and of quasistatic movements. The aim of our study was therefore to determine the development of the axis orientation during walking. Intracortical bone pins were used to monitor the kinematics of the talus and tibia of five healthy volunteers. The finite helical axis was determined for moving windows of 10% stance phase and its orientation reported if the rotation about the axis was more than 2°. A representative axis for ankle dorsi- and plantarflexion was also estimated based on tarsal morphology. As reported by literature, the morphology-based axis was inclined more medially upwards for dorsiflexion than for plantarflexion. However, when a mean of the finite helical axis orientations was calculated for each walking trial for dorsiflexion (stance phase 15-25%) and for plantarflexion (stance phase 85-95%), the inclination was less medially upwards in dorsiflexion than in plantarflexion in four out of five participants. Thus, it appears that the inclination of a representative ankle joint axis for dynamic loading situations cannot be estimated from either morphology or quasi-static experiments. Future studies assessing muscle activity, ligament behaviour and articulating surfaces may help to identify the determining factors for the orientation of a representative ankle joint axis.

Multibody kinematics optimization with marker projection improves the accuracy of the humerus rotational kinematics.

Markers put on the arm undergo large soft tissue artefact (STA). Using markers on the forearm, multibody kinematics optimization (MKO) helps improve the accuracy of the arm kinematics especially its longitudinal rotation. However deleterious effect of STA may persist and affect other segment estimate. The objective was to present an innovative multibody kinematics optimization algorithm with projection of markers onto a requested axis of the local system of coordinates, to cancel their deleterious effect on this degree-of-freedom. Four subjects equipped with markers put on intracortical pins inserted into the humerus, on skin (scapula, arm and forearm) and subsequently on rigid cuffs (arm and forearm) performed analytic, daily-living, sports and range-of-motion tasks. Scapulohumeral kinematics was estimated using 1) pin markers (reference), 2) single-body optimization, 3) MKO, 4) MKO with projection of all arm markers and 5) MKO with projection of a selection of arm markers. Approaches 2-4 were applied to markers put on the skin and the cuff. The main findings were that multibody kinematics optimization improved the accuracy of 40-50% and the projection algorithm added an extra 20% when applied to cuff markers or a selection of skin markers (all but the medial epicondyle). Therefore, the projection algorithm performed better than multibody and single-body optimizations, especially when using markers put on a cuff. Error of humerus orientation was reduced by half to finally be less than 5°. To conclude, this innovative algorithm is a promising approach for estimating accurate upper-limb kinematics.

The effect of intracortical bone pin application on kinetics and tibiocalcaneal kinematics of walking gait.

Bone anchored markers using intracortical bone pins are one of the few available methods for analyzing skeletal motion during human gait in-vivo without errors induced by soft tissue artifacts. However, bone anchored markers require local anesthesia and may alter the motor control and motor output during gait. The purpose of this study was to examine the effect of local anesthesia and the use of bone anchored markers on typical gait analysis variables. Five subjects were analyzed in two different gait analysis sessions. In the first session, a protocol with skin markers was used. In the second session, bone anchored markers were added after local anesthesia was applied. For both sessions, three dimensional infrared kinematics of the calcaneus and tibia segments, ground reaction forces, and plantar pressure data were collected. 95% confidence intervals and boxplots were used to compare protocols and assess the data distribution and data variability for each subject. Although considerable variation was found between subjects, within-subject comparison of the two protocols revealed non-systematic effects on the target variables. Two of the five subjects walked at reduced gait speed during the bone pin session, which explained the between-session differences found in kinetic and kinematic variables. The remaining three subjects did not systematically alter their gait pattern between the two sessions. Results support the hypothesis that local anesthesia and the presence of bone pins still allow a valid gait pattern to be analyzed.

Interspecific variation in the relationship between clutch size, laying date and intensity of urbanization in four species of hole-nesting birds.

The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole-nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large-scale study showing a species-specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size.

Glenohumeral joint kinematics measured by intracortical pins, reflective markers, and computed tomography: A novel technique to assess acromiohumeral distance.

Combination of biplane fluoroscopy and CT-scan provides accurate 3D measurement of the acromiohumeral distance (AHD) during dynamic tasks. However, participants performed only two and six trials in previous experiments to respect the recommended radiation exposure per year. Our objective was to propose a technique to assess the AHD in 3D during dynamic tasks without this limitation. The AHD was computed from glenohumeral kinematics obtained using markers fitted to pins drilled into the scapula and the humerus combined with 3D bone geometry obtained using CT-scan. Four participants performed range-of-motion, daily-living, and sports activities. Sixty-six out of 158trials performed by each participant were analyzed. Two participants were not considered due to experimental issues. AHD decreased with arm elevation. Overall, the smallest AHD occurred in abduction (1.1mm (P1) and 1.2mm (P2)). The smallest AHD were 2.4mm (P1) and 3.1mm (P2) during ADL. It was 2.8mm (P1) and 1.1mm (P2) during sports activities. The humeral head greater and lesser tuberosities came the nearest to the acromion. The proposed technique increases the number of trials acquired during one experiment compared to previous. The identification of movements maximizing AHD is possible, which may provide benefits for shoulder rehabilitation.

Glenohumeral translations during range-of-motion movements, activities of daily living, and sports activities in healthy participants.

BACKGROUND: Glenohumeral translations have been mainly investigated during static poses while shoulder rehabilitation exercises, activities of daily living, and sports activities are dynamic. Our objective was to assess glenohumeral translations during shoulder rehabilitation exercises, activities of daily living, and sports activities to provide a preliminary analysis of glenohumeral arthrokinematics in a broad range of dynamic tasks. METHODS: Glenohumeral translations were computed from trajectories of markers fitted to intracortical pins inserted into the scapula and the humerus. Two participants (P1 and P2) performed full range-of-motion movements including maximum arm elevations and internal-external rotations rehabilitation exercises, six activities of daily living, and five sports activities. FINDINGS: During range-of-motion movements, maximum upward translation was 7.5mm (P1) and 4.7mm (P2). Upward translation during elevations was smaller with the arm internally (3.6mm (P1) and 2.9mm (P2)) than neutrally (4.2mm (P1) and 3.7mm (P2)) and externally rotated (4.3mm (P1) and 4.3mm (P2)). For activities of daily living and sports activities, only anterior translation during reach axilla for P1 and upward translation during ball throwing for P2 were larger than the translation measured during range-of-motion movements (108% and 114%, respectively). INTERPRETATION: While previous electromyography-based studies recommended external rotation during arm elevation to minimize upward translation, measures of glenohumeral translations suggest that internal rotation may be better. Similar amplitude of translation during ROM movement and sports activities suggests that large excursions of the humeral head may be caused not only by fast movements, but also by large amplitude movements.

Variation in clutch size in relation to nest size in birds.

Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Coupling between 3D displacements and rotations at the glenohumeral joint during dynamic tasks in healthy participants.

BACKGROUND: Glenohumeral displacements assessment would help to design shoulder prostheses with physiological arthrokinematics and to establish more biofidelic musculoskeletal models. Though displacements were documented during static tasks, there is little information on their 3D coupling with glenohumeral angle during dynamic tasks. Our objective was to characterize the 3D glenohumeral displacement-rotation couplings during dynamic arm elevations and rotations. METHODS: Glenohumeral displacements were measured from trajectories of reflective markers fitted on intracortical pins inserted into the scapula and humerus. Bone geometry was recorded using CT-scan. Only four participants were recruited to the experiment due to its invasiveness. Participants performed dynamic arm abduction, flexion and axial rotations. Linear regressions were performed between glenohumeral displacements and rotations. The pin of the scapula of one participant moved, his data were removed from analysis, and results are based on three participants. FINDINGS: The measurement error of glenohumeral kinematics was less than 0.15mm and 0.2°. Maximum glenohumeral displacements were measured along the longitudinal direction and reached up to +12.4mm for one participant. Significant couplings were reported especially between longitudinal displacement and rotation in abduction (adjusted R(2) up to 0.94). INTERPRETATION: The proposed method provides the potential to investigate glenohumeral kinematics during all kinds of movements. A linear increase of upward displacement during dynamic arm elevation was measured, which contrasts with results based on a series of static poses. The systematic investigation of glenohumeral displacements under dynamic condition may help to provide relevant recommendation for the design of shoulder prosthetic components and musculoskeletal models.

A two-year radiostereometric follow-up of the first generation Birmingham mid head resection arthroplasty.

During the first decade of the 21st century, metal-on-metal hip resurfacing became one of the main treatment options for younger, more active patients with osteoarthritis. However, as a result of the reported failure rate of both total hip replacement (THR) and resurfacing in patients with considerable loss of bone stock in the femoral head (e.g., in extensive avascular necrosis), other solutions have been sought for these patients. The short-stemmed Birmingham Mid Head Resection prosthesis (BMHR) combines a metal-on-metal articulation and a femoral neck preserving feature. In this study, radiostereometric analysis (RSA) was used to study migration of the BMHR femoral component in 13 hips. Translations and rotations were measured up to two years. Relative values showed no statistically significant migration. Absolute values demonstrated settling in occurring between zero and two months postoperatively in all directions studied. From two months to two years no significant migration occurred except for rotation around the x-axis of the femoral segment (p = 0.049). After initial settling-in, absolute values were low, indicating that there was no evidence of early migration or loosening of the components.

Effect of an antipronation foot orthosis on ankle and subtalar kinematics.

INTRODUCTION/PURPOSE: The aim of this study was to describe the effect of an antipronation foot orthosis on motion of the heel relative to the leg and explore the individual contributions of the ankle and subtalar joints to this effect. METHODS: Five subjects were investigated using invasive intracortical pins to track the movement of the tibia, talus, and calcaneus during walking with and without a foot orthosis. RESULTS: The antipronation foot orthosis produced small and unsystematic reductions in eversion and abduction of the heel relative to the leg at various times during stance. Changes in calcaneus-tibia motion were comparable with those described in the literature (1°-3°). Changes at both the ankle and subtalar joints contributed to this orthotic effect. However, the nature and scale of changes were highly variable between subjects. Peak angular position, range of motion, and angular velocity in frontal and transverse planes were affected to different degrees in different subjects. In some cases, changes occurred mainly at the ankle; in other cases, changes occurred mainly at the subtalar joint. CONCLUSION: The changes in ankle and subtalar kinematics in response to the foot orthosis contradict existing orthotic paradigms that assume that changes occur only at the subtalar joint. The kinematic changes due to the orthosis are indicative of a strong interaction between the often common function of the ankle and subtalar joints.

The influences of impact interface, muscle activity, and knee angle on impact forces and tibial and femoral accelerations occurring after external impacts.

The purpose of this study was to quantify relative contributions of impact interface, muscle activity, and knee angle to the magnitudes of tibial and femoral accelerations occurring after external impacts. Impacts were initiated with a pneumatically driven impacter under the heels of four volunteers. Impact forces were quantified with a force sensor. Segmental accelerations were measured with bone mounted accelerometers. Experimental interventions were hard and soft shock interfaces, different knee angles (0 degrees, 20 degrees, 40 degrees knee flexion), and muscular preactivation (0%, 30%, 60% of maximal voluntary contraction) of gastrocnemii, hamstrings, and quadriceps. Greater knee flexion led to lower impact forces and higher tibial accelerations. Increased muscular activation led to higher forces and lower tibial accelerations. The softer of the two shock interfaces under study reduced both parameters. The effects on accelerations and forces through the activation and knee angle changes were greater than the effect of interface variations. The hardness of the two shock interfaces explained less than 10% of the variance of accelerations and impact forces, whereas knee angle changes explained 25-29%, and preactivation changes explained 35-48% of the variances. It can be concluded that muscle force and knee joint angle have greater effects in comparison with interface hardness on the severity of shocks on the lower leg.

Does a specific MR imaging protocol with a supine-lying subject replicate tarsal kinematics seen during upright standing?

Magnetic resonance (MR) imaging is becoming increasingly important in the study of foot biomechanics. Specific devices have been constructed to load and position the foot while the subject is lying supine in the scanner. The present study examines the efficacy of such a newly developed device in replicating tarsal kinematics seen during the more commonly studied standing loading conditions. The results showed that although knee flexion and the externally applied load were carefully controlled, subtalar and talo-navicular joint rotations while lying during MR imaging and when standing (measured opto-electrically with markers attached to intracortical pins) did not match, nor were they systematically shifted. Thus, the proposed MR protocol cannot replicate tarsal kinematics seen during upright standing. It is concluded that specific foot loading conditions have to be considered when tarsal kinematics are evaluated. Improved replication of tarsal kinematics in different postures should comprehensively consider muscle activity, a fixed hip position, and a well-defined point of load application.

Ankle and subtalar kinematics measured with intracortical pins during the stance phase of walking.

BACKGROUND: The absence of external landmarks on the talus has rendered the description of ankle and subtalar joint kinematics difficult. Abnormal motion at these joints has, however, been implied in the etiology of an array of lower extremity overuse injuries. METHODS: Intracortical pins were inserted under local anesthesia in the tibia, talus, and calcaneus with external marker clusters traced by a video motion analysis system. Kinematic data were collected during walking trials on a flat surface for three subjects. Gait pattern was monitored by comparison of ground reaction force curves during stance phase with and without the pins inserted. RESULTS: Results were presented in terms of helical axis orientation for both joints and the component rotations about these axes. Large intersubject differences were seen in both ankle and subtalar joint helical axis orientation. Maximum rotations over the complete stance phase for the ankle and subtalar joints respectively were: eversion/inversion, 6.3 degrees and 8.3 degrees; dorsiflexion/plantarflexion, 18.7 degrees and 3.7 degrees; and abduction/adduction, 5.0 degrees and 6.1 degrees. CONCLUSIONS: The majority of ankle eversion/inversion occurred at the subtalar joint; however, the ankle component cannot be ignored. Abduction/adduction range of motion at the subtalar joint was surprisingly high, indicating that this component motion during walking is not purely attributable to the ankle joint. Future research should include greater subject numbers in order to present more universally applicable results. CLINICAL RELEVANCE: The in vivo kinematics of the talus during weightbearing activity are poorly understood. The description of this motion may assist in the structuring of clinical rehabilitation and in the design and insertion of ankle joint prostheses.

Differences in ankle-joint complex motion during the stance phase of walking as measured by superficial and bone-anchored markers.

Bones of the ankle-joint complex are difficult to represent with superficial markers and an invasive approach is required to quantify skin movement artifact. Three-dimensional coordinates during gait were first calculated from sets of three superficial markers located on both the lower leg and heel. Intracortical pins equipped with external marker arrays were subsequently inserted in the tibia, talus and calcaneus for further trials. Tibiocalcaneal and talocalcaneal joint helical axis component rotations were calculated. Intersubject motion patterns were comparable. Root mean square (RMS) differences were chosen to describe discrepancies between temporal intracortical pin and superficial marker curves. Results for the tibiocalcaneal rotations inversion/eversion, plantarflexion/ dorsiflexion and abduction/adduction were 2.5 degrees, 1.7 degrees and 2.8 degrees respectively. Inversion/eversion about the talocalcaneal joint showed an RMS difference of 2.1 degrees. A systematic over- or underestimation of superficial relative to bone-anchored markers could not be determined.

Inbreeding depression and male fitness in black grouse.

The male lifetime lekking performance was studied, and related to inbreeding-outbreeding in a wild population of black grouse (Tetrao tetrix) in central Finland between 1989 and 1995. Inbreeding was measured as the mean heterozygosity and mean d(2) of 15 microsatellite loci. We found a significantly positive relationship between mean d(2) and lifetime copulation success (LCS), while the relationship between heterozygosity and LCS was close to significant. We also found that males that never obtained a lek territory had significantly lower mean heterozygosity than males that were observed on a territory at least during one mating season in their life. Furthermore, among males that were successful in obtaining a lek territory, LCS and mean d(2) were highest for those males that held central territories. We suggest that inbred males have a disadvantage (or outbred males have an advantage) in the competition for territories that may explain the relationships with LCS and inbreeding. Furthermore, the fact that mean d(2) was positively correlated with LCS whereas heterozygosity was not when we restricted the analysis to territorial males, suggests that mean d(2) provides more information about levels of inbreeding-outbreeding than heterozygosity alone, and potentially highlights the effects of heterosis. To our knowledge, this is the first time that measures of inbreeding and lifetime fitness have been linked in a non-isolated population. This is important in establishing that the relationships found in previous studies are not artefacts of low gene flow created by limited dispersal but a general feature of wild vertebrate populations.

Effects of latitude on the trade-off between reproduction and moult: a long-term study with pied flycatcher.

Long-distance migratory passerine birds are generally time constrained by reproduction and moult, which need to be completed before migration. Breeding and post-nuptial moult may overlap especially under time-constrained conditions (northern latitudes). Here, we analysed the timing of adult moult in relation to latitude, timing of breeding and reproductive effort in pied flycatchers (Ficedula hypoleuca) breeding in four widely separated populations (40-68° N). In males but not females, the proportion of moulting birds while provisioning nestlings increased with increasing latitude. This may suggest that a moult-breeding overlap is a strategy employed by male pied flycatchers to adjust to the short breeding season at northern latitudes. However, the moult-breeding overlap was more pronounced among males in the southernmost study population (Spain). In this population, males may decide not to invest more in reproduction, and start moulting at earlier breeding stage than in northern populations,or, alternatively, birds in the Mediterranean region are time constrained by the hot and dry summer. The trade-off between breeding and post-nuptial moult may be more important in some populations than in others, depending on the latitude of the breeding site. Our results show that a moult-breeding overlap imposes a fitness cost on males in terms of fecundity and breeding success.

Trade-off between reproduction and moult - a comparison of three Fennoscandian pied flycatcher populations.

Organisms that reproduce at high latitudes are assumed to have evolved several adaptations to the short summer. For birds, and especially for long-distance migrants, there is a time constraint because both reproduction and moult must be completed before autumn migration. It has therefore been assumed that birds at northern latitudes must initiate their moult during reproduction more often than birds at low latitudes. To investigate how passerine birds breeding at different latitudes allocate their time between reproduction and moult, we compared timing of these activities during three consecutive breeding seasons in three widely separated populations of the pied flycatcher Ficedula hypoleuca. Our results show that the frequency of individuals with moult-breeding overlap, and moult initiation in relation to breeding stage, varied considerably among populations and years. In all three populations, female moult initiation was restricted to the late nestling period. The males had a more pronounced moult-breeding overlap than the females, but its duration was similar in all three study areas. Thus, there was no evidence for a more pronounced moult-breeding overlap at high compared with low latitudes. These results suggest that pied flycatchers sometimes accept a moult-breeding overlap, but that the time gained by having too extensive an overlap between reproduction and moult does not outweigh the associated costs. Long-distance migrants breeding at northern latitudes apparently experience a trade-off between reproduction and somatic investment during moult. We therefore suggest that a pronounced moult-breeding overlap is not a typical strategy used by long-distance migrants to adjust to the short breeding season at northern latitudes.

HERITABILITY AND SELECTION ON TARSUS LENGTH IN THE PIED FLYCATCHER (FICEDULA HYPOLEUCA).

The heritability estimate (±SE) for tarsus length in the pied flycatcher is 0.53 ± 0.10, based on mother-offspring regressions. The heritability is almost the same (0.50 ± 0.22) for offspring transferred to other nests and reared by foster parents, whereas there is no resemblance (0.04 ± 0.23) between the offspring and their foster mothers. Hence, the nest environment does not affect parent-offspring resemblance. However, a full-sib correlation yields an estimate of the heritability twice as high as the parent-offspring regression did, indicating that shared environment effects, which are not correlated with the tarsus length of parents, must be important. An environmental deviation due to food factors affecting tarsus length is demonstrated. The most important food factors are probably associated with 1) polygyny, which leads to reduced paternal feeding at secondary nests, 2) high breeding density, and 3) progress of the breeding season. All three are associated with reductions in offspring mean tarsus length. We estimate selection on tarsus length for the major components of lifetime reproductive success. Offspring with the shortest tarsi have reduced survival from fledging until breeding, and males with tarsus length close to the mean are most successful in attracting mates. Clutch size increases with female tarsus length, except for individuals with very long tarsi. In general, directional selection is weak, but stabilizing selection is rather strong for survival and male mating success.