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Bilateral asymmetry in the lower limbs in medieval humans and La Ferrassie Neandertals LF1 and LF2

Asymétrie bilatérale des membres inférieurs chez des humains médiévaux et les Néandertaliens de La Ferrassie LF1 et LF2
Ameline Bardo et Anna Maria Kubicka

Résumés

L’asymétrie bilatérale des membres inférieurs peut fournir des informations sur les contraintes biomécaniques habituelles et les schémas de mobilité des populations anciennes. Les études précédentes ont généralement analysé les différents éléments du membre inférieur de manière isolée, ce qui a potentiellement négligé les signaux fonctionnels intégrés à l’échelle du membre. Nous présentons ici de nouvelles données sur l’asymétrie bilatérale chez des humains médiévaux et chez les Néandertaliens de La Ferrassie (LF1 et LF2), en combinant des mesures externes à des analyses biomécaniques transversales. Nous avons quantifié l’asymétrie directionnelle (DA) et l’asymétrie absolue (AA) du fémur, du tibia, ainsi que du premier (MT1) et du cinquième métatarsien (MT5), afin d’évaluer les chargements mécaniques habituels. Les humains médiévaux présentaient une asymétrie modérée du MT1 et du tibia, sans dominance latérale constante ni corrélation entre les différents os, ce qui suggère une forte variabilité individuelle. LF1 et LF2 présentaient une DA plus élevée, avec des profils distincts entre individus, probablement liés à des charges mécaniques plus intenses ou plus répétitives au niveau des membres inférieurs. Dans l’ensemble, nos résultats mettent en évidence la variabilité individuelle, un possible dimorphisme sexuel et des adaptations spécifiques selon les segments, tout en soulignant que l’asymétrie des métatarsiens reste encore peu documentée dans les populations fossiles. De plus, les données issues de CT scans à haute résolution ont permis de recalculer plus précisément les propriétés biomécaniques de LF1 et LF2.

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Introduction

1The biomechanical properties of long bones reflect physical activity and habitual behaviour patterns (e.g., Niinimäki, 2012). When a sufficient threshold of mechanical loading is applied, bones respond through changes in their internal structure, including modifications in the orientation, size and distribution of osteons (Frost, 1987; Judex and Carlson, 2009). Consequently, differences in biomechanical properties between the left and right sides of an individual provide a sensitive indicator of limb plasticity and the degree of unilateral or bilateral loading (Kubicka et al., 2018). In the upper limb, such asymmetry is commonly right-biased and largely reflects behavioural lateralization and handedness (e.g., Kubicka et al., 2018; Brzezinski et al., 2023; Temple et al., 2023). In contrast, asymmetry of the lower limb has received far less attention, even though it may provide critical information about habitual locomotor loading. This is partly because lower limb asymmetry tends to be less pronounced than in the upper limb (Auerbach and Ruff, 2006), or because paired skeletal elements in osteological material are often poorly preserved. Nevertheless, understanding lower limb asymmetry is essential for reconstructing habitual loading patterns and skeletal plasticity in past populations.

2In bipedal hominins, the lower limb forms an integrated weight-bearing system that plays a central role in postural support and locomotion (Dowdeswell et al., 2017). Consequently, lower limb bones are generally exposed to greater and more sustained mechanical loads than the upper limb (Auerbach and Ruff, 2006; Shaw and Stock, 2013). However, these loads are not distributed uniformly across the limb. The femur is subjected to substantial bending moments during walking and running, which generate characteristic patterns of cortical distribution and cross-sectional shape (Ruff, 1995; Duda et al., 1997). In contrast, the tibia primarily experiences axial compressive loads transmitted from body mass (Ruff, 1995). Because these two bones are subjected to different loading regimes while functioning together during locomotion, their patterns of structural adaptation and bilateral asymmetry are not expected to be identical. Examining the femur and tibia within the same individuals therefore provides a highly informative framework for assessing how habitual mechanical loading is expressed across the lower limb (Stock and Pfeiffer, 2001; Sugiyama et al., 2010; Macintosh and Stock, 2019).

3Research on the lower limb has focused primarily on reconstructing locomotor behaviour, mobility and adaptations to bipedalism in both fossil and modern populations (Wescott and Cunningham, 2006; Cejudo et al., 2020; Hill et al., 2020; Beato et al., 2021). These studies have consistently shown that femoral and tibial morphology and internal structure vary systematically with locomotor demands, terrain use and the intensity of activity. In particular, variation in femoral diaphyseal curvature or robusticity reflects differences in bending regimes, gait mechanics and overall locomotor loading (Shackelford and Trinkaus, 2002; De Groote, 2011; Shaw and Stock, 2013; Chapman et al., 2018). Likewise, analyses of the trabecular and diaphyseal structural properties of the femur and tibia have applied the bone functional adaptation framework to archaeological and fossil samples, showing that joint loading patterns, mobility strategies and body-mass-related stresses leave a measurable signature in internal bone architecture (Ruff et al., 2006; Georgiou et al., 2020; Cazenave et al., 2022; Daver et al., 2022; Lukova et al., 2024). Together, these studies highlight the value of lower-limb bone structure for investigating patterns of habitual locomotor behaviour. However, despite this extensive literature, these elements are almost always analysed in isolation. As a result, relatively few studies have specifically examined how different bones of the lower limb co-vary within the same individuals in response to shared mechanical constraints. Most population-level studies therefore focus on the femur and tibia alone (e.g. Eckhoff et al., 2016; Auerbach et al., 2017; Mopin et al., 2018), limiting our ability to assess how loading is distributed across the limb as a functional system.

4The lower limb does not end at the tibia: mechanical loads generated during locomotion are ultimately transmitted to the foot, where they are redistributed through the tarsals and metatarsals during stance and propulsion of bipedal locomotion. In particular, the first (MT1) and fifth metatarsals (MT5) occupy functionally distinct positions within the medial and lateral regions of the foot, and experience contrasting loading regimes associated with the longitudinal arch during walking and running (Vereecke et al., 2003; Griffin and Richmond, 2005; Marchi, 2005; Pontzer et al., 2010; Dowdeswell et al., 2017). As a result, MT1 and MT5 provide complementary information to that obtained from the femur and tibia, thus helping to clarify how loads are distributed across the foot during habitual locomotion. Studies of metatarsal asymmetry have documented that these bones respond to both biological and mechanical factors. Developmental differences in metatarsal size and proportional growth are already detectable during foetal development in modern humans (Gawlikowska et al., 2007), and metatarsal proportions vary with age and sex in recent human populations, reflecting normal biological variability in skeletal growth and dimorphism (e.g., Rewekant, 1996; Mountrakis et al., 2010). In adults, asymmetry of the MT1 has been linked to pathological shortening in arthrosis (Davitt et al., 2005) and hallux valgus elongation (Munuera et al., 2008), whereas asymmetry of the MT5 is associated with stress-related fracture risks in athletes (Sun et al., 2024). Together, these studies demonstrate that metatarsal asymmetry reflects differences in mechanical loading and functional use of the foot. Despite the central role of the foot in bipedal locomotion, metatarsal asymmetry has not yet been used to reconstruct habitual biomechanical stress or mobility patterns in past populations. MT1 and MT5 are subjected to markedly different loads during gait, with MT1 experiencing greater pressures during push-off and forward propulsion (Day and Napier, 1964; Wearing et al., 2001; Marchi, 2005) and being structurally more robust than the lateral rays (Danesi et al., 2012). Understanding metatarsal asymmetry therefore offers a unique opportunity to extend biomechanical interpretations beyond the femur and tibia and to capture how foot loading has contributed to locomotor behaviour and skeletal plasticity (Auerbach and Ruff, 2006; Shaw and Stock, 2013; Dowdeswell et al., 2017).

5Although the biomechanical properties of lower-limb bones and metatarsals can change throughout life in response to mechanical loading (Ruff et al., 2006), ontogenetic studies in primates show that aspects of diaphyseal structure are also partly inherited (Hansen et al., 2009; Morimoto et al., 2011; 2014; Cosnefroy et al., 2022). However, genetic canalization is unlikely to generate consistent directional asymmetry between the left and right sides. Bilateral differences in biomechanical properties are therefore informative indicators of habitual mechanical loading and activity patterns in individuals.

6Lower-limb loading does not occur in isolation from the rest of the body. During walking and running, the upper limbs contribute to balance and angular momentum control by moving in opposition to the legs (Pontzer et al., 2009; Shernice et al., 2021; Koo et al., 2025). As a consequence, habitual dominance of one arm can influence loading patterns in the contralateral lower limb. This functional coupling has been demonstrated in modern humans, where individuals with right-biased upper-limb dimensions tend to exhibit slight left-biased biomechanical properties in the leg bones, including diaphyseal breadths and second moments of area (Płochocki, 2004). This pattern, known as crossed symmetry, reflects opposing directional asymmetries between the upper and lower limbs and has been documented in bone lengths, articular surfaces and cross-sectional geometry in modern humans (Schaeffer, 1928; Płochocki, 2002; 2004; Auerbach and Ruff, 2006; Eriksen, 2020). However, this phenomenon has not yet been investigated in Neandertals.

7Neandertal limb bones are characterized by high levels of robusticity relative to recent human populations, generally interpreted as reflecting intense and region-specific mechanical loading rather than greater biological sensitivity to mechanical stress (e.g., for the lower limbs: Lovejoy and Trinkaus, 1980; Ruff et al., 1997; Trinkaus et al., 1991; Trinkaus and Ruff, 1999; Ruff et al., 2006; Churchill and Rhodes, 2009; Rodríguez et al., 2018; Kubicka et al., 2022; for the upper limbs: Trinkaus, 1980; Stock and Pfeiffer, 2001; Kubicka et al., 2018). However, when biomechanical properties are scaled to body size, Neandertals do not consistently exceed Middle Palaeolithic Homo sapiens in femoral and tibial robusticity (Trinkaus and Ruff, 2012), indicating that their skeletal strength reflects specific loading regimes rather than simple hypertrophy. Patterns of bilateral asymmetry also differ between limb segments: the femur and tibia generally show low asymmetry, consistent with balanced axial loading during habitual bipedal locomotion (Lovejoy and Trinkaus, 1980; Trinkaus and Ruff, 1999; Ruff et al., 1997; 2006), whereas the upper limb exhibits more pronounced and more often right-dominant asymmetry reflecting unilateral manipulative behaviours (Trinkaus, 1980; Stock and Pfeiffer, 2001; Pearson et al., 2006; Kubicka et al., 2018). In contrast, metatarsal asymmetry in Neandertals and fossil H. sapiens is still poorly documented, despite the crucial role of these bones in locomotion (Rhoads and Trinkaus, 1977; Auerbach and Ruff, 2006; Shaw and Stock, 2013; Venkadesan et al., 2017; Pablos et al., 2019).

8Despite decades of research on limb bone biomechanics and asymmetry, most studies have examined single skeletal elements in isolation, typically the femur or tibia, rather than evaluating how different segments of the same lower limb respond to shared mechanical constraints. As a result, it remains unclear whether asymmetry patterns are expressed consistently across the femur, tibia and foot of individuals, or whether different bones record distinct aspects of habitual loading. To date, no study has combined the cross-sectional geometry of long bones with metatarsal asymmetry to investigate bilateral asymmetry across the entire lower limb in either modern or fossil humans.

9In this study, we investigate patterns of bilateral asymmetry across the femur, tibia, and foot in the same individuals. Using a medieval agricultural population as a comparative framework, we examine how asymmetry is distributed across the lower limb and whether different bones co-vary in their response to habitual loading. These results are then used to interpret asymmetry patterns in two hunter-gatherer Neandertals from La Ferrassie (LF1 and LF2), whose lower-limb biomechanics have previously been described but never analysed within a multi-element asymmetry framework. We quantified both directional asymmetry (DA) and absolute asymmetry (AA) in the femur, tibia, and the MT1 and MT5. Directional asymmetry refers to the signed difference between the left and right sides for a given parameter, indicating the direction of the side difference. Absolute asymmetry reflects the magnitude of this difference, independently of its direction. We further tested whether asymmetry levels are correlated among these elements within individuals, which allowed us to evaluate whether different segments of the lower limb reflect shared or independent mechanical histories. Finally, the biomechanical properties of two Neandertals (i.e. LF1 and LF2) that were previously analysed by Trinkaus and Ruff (2012) were revisited using high-resolution micro-CT scans and virtual anthropological methods. We hypothesized that:
- MT1 would show greater asymmetry than MT5, reflecting its dominant role in load transmission and forward propulsion during gait (e.g., Day and Napier, 1964; Wearing et al., 2001; Marchi, 2005; Danesi et al., 2012);
- the tibia would exhibit greater asymmetry than the femur, as it is more responsive to mechanical loading during life (Macintosh and Stock, 2019);
- the medieval population would show higher levels of asymmetry than LF1 and LF2, consistent with more repetitive and unilateral lower-limb loading associated with agricultural activities such as digging, carrying and trampling, compared with the more varied mobility of hunter-gatherers (e.g., walking, hunting and gathering);
- the direction of asymmetry would be associated across the lower limb, so that metatarsal asymmetry and long-bone biomechanical properties would tend to show the same side dominance, reflecting shared mechanical loading within the limb.

Material and methods

Material

10In this study, we selected 21 adults (13 males, 8 females) from a large medieval group (of about 200 skeletons), all of whom had well-preserved pairs of the first (MT1) and fifth (MT5) metatarsals, as well as femurs and tibias. To ensure accurate measurements, we only included individuals with well-preserved epiphyses and external shaft surfaces on both sides of the lower limb bones (i.e. femur, tibia and metatarsals). This material comes mostly from a farming community and originates from a cemetery dating from the 10th to the end of the 13th century located in Ostrów Lednicki, Poland (Kubicka et al., 2022). We also examined the bones of two Neandertal adults, La Ferrassie 1 (LF1) and La Ferrassie 2 (LF2), analysing pairs of MT1, MT5, femurs and tibias. Both Neandertal individuals date to 42-47 ka (Guérin et al., 2023).

Methods

11To analyse the level of asymmetry in MT1 and MT5, we took seven linear measurements as described by Barrio and collaborators (2006) (figure 1; table 1). Measurements of epicondylar diameters were not included in this study as this feature was not well preserved in LF1 and LF2. Each of the seven measurements was performed twice by one observer using a digital calliper, with an accuracy of 0.01 mm, on both sides of the body and for each metatarsal and sample. The complete right foot of LF2 is still attached to a large block of sediment. The 3D models of the MT1 and MT5 of this specimen were then measured virtually twice by two observers using Gom Inspect software (v 2.0.1). The MT measurements and the right foot measurements of LF2 were consistent between the two observers, with Interclass Correlation Coefficients (ICC) ranging from 0.92 to 0.99, p<0.05.

Figure 1

Figure 1

Measurements taken on the left MT1 and MT5. The abbreviations used are described in table 1 |
Mesures prises sur les MT1 et MT5 gauches. Les abréviations utilisées sont décrites dans le tableau 1

Table 1

Table 1

Description of linear measurements of the MT1 and MT5 (after Barrio et al., 2006) |
Description des mesures linéaires réalisées sur les MT1 et MT5 (d’après Barrio et al., 2006)

12Pairs of femurs and tibias from the medieval group were scanned in the craniocaudal direction using cone-beam computed tomography (CBCT) with the standard protocol (0.625 mm3, 120kV). The long bones of LF1 and LF2, and the complete right foot of LF2, were digitised using a medical CT scan (Philips iCT128 platform, 0.312 mm3, 125kV). Three-dimensional (3D) reconstructions were created for each scanned femur and tibia using 3D Slicer software (Fedorov et al., 2012). Manual segmentation and virtual reconstruction were performed for the bones of LF1 and LF2 because some specimens contained imaging artefacts that could have affected the biomechanical analysis. The biomechanical length of the fully preserved femurs and tibiae of the Medieval population was measured using Gom Inspect software (v2.0.1). The 3D models were then oriented in 3D using MeshLab software (Cignoni et al., 2008), with the x-y plane representing the transverse plane and the x-z and y-z planes defining the coronal and sagittal planes respectively (see Profico et al., 2021 for details). The poorly preserved bones of LF1 and LF2 were oriented using medieval long bones with preserved epiphyses to help determine the distal ends of the femurs and tibiae in the 3D space. The fully preserved bones of both Neandertals were located based on their biomechanical length using MeshLab software (Cignoni et al., 2008). For this study, we only analysed the biomechanical properties of the femur and tibia because the distal ends of these long bones in LF1 and LF2 are not preserved, so that linear measurements could not be obtained.

13Biomechanical properties were extracted using the morphomap R package (Profico et al., 2021) at five standardized cross-sections (CS) locations at 20%, 35%, 50%, 65% and 80% of the biomechanical length (figure 2). The following biomechanical properties were calculated for each femur and tibia: total area (TA), cortical area (CA), the polar moment of area (J), the maximum second moment of area (Imax), the minimum second moment of area (Imin). The relative amount of cortical area (%CA), which characterizes the resistance of the shaft to axial loadings (Kubicka et al., 2022), was calculated using the CA/TA ratio. We included %CA because it also correlates with J and may reflect adaptive responses to increased mechanical loads during life (Kubicka et al., 2022). The J provides information on the resistance to torsional and bending rigidity loads, and is therefore used as an indicator of robustness (Kubicka et al., 2022). Imax and Imin are then used to calculate the ratio of the maximum to minimum second moments of area (i.e. Imax/Imin). This ratio describes the torsional strength and, because it is based on the main axes, is less sensitive to section orientation than second moments calculated relative to fixed anatomical axes (Ix/Iy) (Stock and Pfeiffer, 2001).

Figure 2

Figure 2

Illustration of the five cross-sections (CS) analysed at different percentages of the length of a right femur (left) and a right tibia (right) of a medieval individual from Ostrów Lednicki |
Illustration des cinq coupes transversales (CS) analysées à différents pourcentages de la longueur d’un fémur droit (à gauche) et d’un tibia droit (à droite) d’un individu médiéval provenant d’Ostrów Lednicki

14Trinkaus and Ruff (2012) estimated the biomechanical properties of LF1 and LF2 using parallax-corrected radiographs, most likely derived from figures published by Heim (1982). Because fossil long bones are often partially reconstructed and filled with sediment, such indirect approaches may limit the accurate identification of cortical boundaries. In this study, biomechanical properties were recalculated using high-resolution CT scans of the femurs and tibias (figure 3), which allow precise discrimination between preserved and reconstructed cortical regions and minimize distortions related to taphonomic processes, as documented in previous methodological studies (Ruff et al., 1989; Ryan and Shaw, 2015; Kivell, 2016). This approach enabled us to both reassess previously published values for LF1 and LF2 and calculate new biomechanical properties for four cross-sections of LF2 that had not been analysed previously. The newly computed values correspond to CS20% of the left and right femora, CS35% of the left femur and CS80% of the right femur of LF2 (figure 3).

Figure 3

Figure 3

Reconstructions of the LF1 (A) and LF2 (B) lower bones, femurs at the top and tibias below, with illustration of the CS analysed |
Reconstitutions des os long des jambes de LF1 (A) et LF2 (B), les fémurs en haut et les tibias en bas, avec une illustration des sections transversales analysées

Statistical analysis

15Shapiro-Wilk Normality tests revealed that some of the population data deviated significantly from a normal distribution (p<0.05), so non-parametric tests were used. Wilcoxon signed-rank tests were used to assess side differences in metatarsal measurements and in femoral and tibial biomechanical properties, including both raw variables (J) and derived indices (%CA and Imax/Imin), at each of the five CS. The dimensions of the MTs and the biomechanical properties of the tibias and femurs were then converted into percentages of directional asymmetry (%DA) and absolute asymmetry (%AA) according to Auerbach and Ruff (2006):

16%DA indicates biased asymmetry, with positive values expressing a right bias and negative values expressing a left bias. %AA shows the total amount of bilateral asymmetry, calculated by removing the signs from %DA. Both of these asymmetry indicators were used to illustrate our results. %DA and %AA were tested for outliers in all MT and biomechanical tibia and femur measurements, in accordance with Hoaglin and Iglewicz (1987). Wilcoxon signed-rank tests were performed with and without outliers, depending on their presence. We also tested for correlation of DA and AA across nine measurements showing significant differences between body sides in medieval humans, using Spearman rank correlation tests: ML PE and AP midshaft of the MT1; %CA (CS 20% and CS 80%), Imax/Imin CS 50% and CS 80%) and J (CS 65% and CS 80%) for the tibias; and J (CS 35%) for the femurs. Due to the small sample size, the statistical tests were performed without dividing by sex, but the descriptive statistics were presented separately for each sex in tables 2 and 3. The statistical analyses were performed using RStudio software (version 4.3.3), with a significance level of 0.05.

Table 2

Table 2

Descriptive statistics (averages±standard errors) for each measurement on the MT1 and MT5, presented for the medieval sample as a whole, and for males (M) and females (F) separately, with percentage of directional asymmetry (%DA) and percentage of absolute asymmetry (%AA) |
Statistiques descriptives (moyenne±erreurs-standards) pour chaque mesure sur les MT1 et les MT5 présentés pour l’échantillon médiéval dans son ensemble, et séparément pour les hommes (M) et les femmes (F), avec le pourcentage d’asymétrie directionnelle (%DA) et le pourcentage d’asymétrie absolue (%AA)

Table 3

Table 3

Mean percentage of directional asymmetry (%DA) and absolute asymmetry (%AA), ±standard errors, for the three biomechanical properties (%CA, Imax/Imin, J) for the five cross-sections (CS) of the femur and the tibia for the medieval humans (M: males; F: females), and values for LF1 and LF2 from this study and a previous one by Trinkaus and Ruff (2012). The bold values are new LF2 measurements not presented in Trinkaus and Ruff (2012) |
Pourcentage moyen d’asymétrie directionnelle (%DA) et d’asymétrie absolue (%AA), ±erreurs standards, pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du fémur et du tibia pour les humains médiévaux (M : mâles ; F : femelles), et les valeurs pour LF1 et LF2 qui proviennent de cette étude et d’une précédente étude par Trinkaus et Ruff (2012). Les valeurs en gras sont de nouvelles mesures pour LF2 qui ne sont pas présentées dans Trinkaus et Ruff (2012)

Results

Directional asymmetry of metatarsals

17The MT1s of the medieval sample showed significant differences between sides for ML PE (W=108, p<0.05), indicating left-side dominance, and for AP midshaft (W=25, p<0.05), indicating right-side dominance (figure 3A). When we tested without the two outliers present for ML PE, we found no significant difference between sides (W=77, p>0.05) (figure 3A).

18There were no significant differences between sides for any of the MT5 measurements of the medieval sample (p>0.05) (figure 3B). When we tested the data after excluding the outliers, we found significant differences between the left and right sides for the ML PE and ML midshaft measurements (both W=67, p<0.05), indicating left-side dominance. Significant differences were found between sides for all measurements for both MT1 and MT5 when tested using absolute values (p<0.01) (see Supplementary table 1).

19Within both LF1 and LF2, DA levels were higher for the MT1 than for the MT5. LF1 exhibited left-side dominance at the AP midshaft of the MT1 (%DA=-4.72), whereas LF2 exhibited right-side dominance (%DA=1.92), similar to the medieval humans (%DA=2.02). LF1 exhibited right-side dominance for ML midshaft of the MT1 (%DA=0.87), comparable to that observed in the medieval humans (%DA=1.59). Conversely, LF2 exhibited stronger right-side dominance (%DA=5.86; figure 4; table 2). For the MT5, LF1 and LF2 generally showed %DA values close to those for the medieval humans, except at the AP midshaft and ML PE. Right-side dominance was observed at the AP midshaft in the medieval humans (%DA=4.35) and LF1 (%DA=5.06), whereas LF2 showed left-side dominance (%DA=-5.46). Left-side dominance was observed for ML PE in medieval humans (%DA=-4.10), while LF2 showed a right-side dominance (%DA=2.72).

Figure 4

Figure 4

Percentage of directional asymmetry (%DA) for the seven linear measurements (see table 1) of the MT1 (A) and MT5 (B) for the medieval sample and LF1 and LF2. A significant level of DA was found for the mediolateral diameter of the proximal epiphysis (ML PE) and the antero-posterior diameter at the midshaft (AP midshaft) of the MT1 (* p<0.05). Each box plot shows the median value (bold line) and the interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-bias, while negative values indicate a left-bias |
Pourcentage d’asymétrie directionnelle (%DA) pour les sept mesures linéaires (voir tableau 1) du MT1 (A) et du MT5 (B) pour l’échantillon médiéval et LF1 et LF2. Un niveau significatif de DA a été trouvé pour le diamètre médiolatéral de l’épiphyse proximale (ML PE) et le diamètre antéro-postérieur au milieu de la diaphyse (AP midshaft) du MT1 (* p<0,05). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "valeurs aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche

Directional asymmetry of femurs

20Medieval humans showed a significant side difference in the femur only for J at 35% of biomechanical length (W=191, p<0.01), indicating left-side dominance (figure 5). When outliers were excluded, left-side dominance was also observed at 50% and 65% of biomechanical length (p<0.05). Analyses based on absolute values revealed significant side differences for all biomechanical properties and cross-sections (p<0.001; Supplementary table 2).

Figure 5

Figure 5

Percentage of directional asymmetry (%DA) for the three biomechanical properties (%CA, Imax/Imin, J) across five cross-sections (CS) of the femur in the medieval humans and in LF1 and LF2. A significant %DA was found for J at CS 35% (** p<0.01). Each box plot shows the median value (bold line) and the interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-side bias, while negative values indicate a left-side bias |
Pourcentage d’asymétrie directionnelle (%DA) pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du fémur, chez les humains du médiéval et LF1 et LF2. Un pourcentage significatif de DA a été observé pour J à CS 35 % (** p<0.01). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "données aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche

21For %CA, LF2 exhibited left-side dominance, whereas LF1 exhibited right-side dominance, similarly to the medieval humans (figure 5). For Imax/Imin, LF2 displayed weak right-side dominance at 35% of the biomechanical length, close to the range for medieval humans, while LF1 showed stronger left-side dominance (figure 5). At the remaining cross-sections (20%, 50%, 65%, and 80% of biomechanical length), both LF1 and LF2 exhibited higher levels of DA than medieval humans. LF2 generally showed higher DA for J than LF1, with right-side dominance relative to both medieval humans and LF1, except at 35% of biomechanical length, where LF1 exhibited right-side dominance (figure 5; table 3).

Directional asymmetry of tibias

22Medieval humans exhibited significant side differences for all three biomechanical properties across the tibia (figure 6; Supplementary table 2). %CA showed right-side dominance at 20% and left-side dominance at 80% of biomechanical length (W=49, p<0.02 and W=180, p<0.05 respectively; figure 6). No significant difference was found when testing after excluding outliers at 80% of biomechanical length. Imax/Imin exhibited left-side dominance at 50% (W=181, p<0.05) and 80 % of biomechanical length (W=194, p<0.01), whereas J showed right-side dominance at 65% (W=25, p<0.001) and 80% of biomechanical length (W=50, p<0.05). When outliers were excluded, J also showed a right-side difference at 50% of biomechanical length (W=54, p<0.05). Analyses based on absolute values indicated significant side differences for all biomechanical properties and cross-sections (p<0.001; Supplementary table 2).

Figure 6

Figure 6

Percentage of directional asymmetry (%DA) c (%CA, Imax/Imin, J) for the five cross-sections (CS) of the tibia for the medieval humans and LF1 and LF2. A significant %DA was found for Imax/Imin (CS 50% and CS 80%) and J (CS 65% and CS 80%) (*** p<0.001; ** p<0.01; * p<0.05). Each box plot shows the median value (bold line) and interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-side bias, while negative values indicate a left-side bias |
Pourcentage d’asymétrie directionnelle (%DA) pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du tibia, chez les humains du médiéval et LF1 et LF2. Un pourcentage significatif de DA a été observé pour Imax/Imin (CS 50 % et CS 80 %) et J (CS 65 % et CS 80 %) (*** p<0,001; ** p<0,01; * p<0,05). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "données aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche

23LF1 and LF2 showed distinct patterns of DA and AA (table 3). For %CA, LF2 displayed stronger left-side dominance, whereas LF1 showed right-side dominance closer to the values for medieval humans at 20%, 35% and 50% of biomechanical length (figure 6). Both LF1 and LF2 exhibited higher AA for %CA at 65% of biomechanical length than medieval humans (table 3). For Imax/Imin, both LF1 and LF2 showed left-side dominance with low DA values at 20% of the biomechanical length, similarly to medieval humans, but right-side dominance at 65%, with particularly high DA for LF2 (%DA=21.28; figure 6). At 35% of biomechanical length, LF2 showed left-side dominance with low DA values, close to the medieval humans, whereas LF1 showed right-side dominance. At 50% of the biomechanical length, LF1 showed left-side dominance, similarly to the medieval humans, while LF2 showed right-side dominance (figure 6; table 3). For J, LF1 exhibited stronger left-side dominance at 20% of biomechanical length than LF2 and the medieval humans. At 50% of biomechanical length, LF1 showed left-side dominance, whereas LF2 and the medieval humans showed right-side dominance. At 65% of biomechanical length, LF1 displayed right-side dominance similarly to the medieval humans, while LF2 showed left-side dominance (figure 6, table 3).

Correlations between directional asymmetry of the lower limb bones

24A significant positive correlation of DA was found in the tibia between 65% and 80% of biomechanical length for J (R=0.73, p<0.01).

Discussion

25This study aimed to evaluate bilateral asymmetry in the lower-limb bones of pre-industrial humans and two La Ferrassie Neandertals (LF1 and LF2) to assess whether asymmetry patterns are consistent with habitual locomotor loading. In the medieval humans, MT1 showed greater DA than MT5, and the tibia exhibited greater DA than the femur; however, directional biases were not consistent at the group level, suggesting substantial inter-individual variation. LF1 and LF2 generally showed higher DA and AA in MT1, the tibia and the femur than the medieval humans, which may reflect more intense and/or repetitive lower limb loading. Overall, bilateral asymmetry likely reflects multiple interacting factors, including activity patterns, locomotor behaviour, and potential sexual dimorphism, which are discussed below.

Asymmetry in the foot bones

26In our medieval sample, we found that MT1 showed a significant level of DA relative to MT5, but there was no significant pattern of side dominance. Indeed, we found two different biases in two regions of MT1: the antero-posterior diameter at the midshaft showed a right-side dominance, indicating a more robust midshaft in the right MT1, whereas the mediolateral diameter of the proximal epiphysis showed left-side dominance, indicating a more robust base in the left MT1. Furthermore, significant asymmetry was observed in all MT1 and MT5 measurements when testing for AA. These results indicate asymmetry in these two bones, but no common direction of dominance at the group level.

27We cannot exclude the possibility that farmers exhibit different patterns to hunter-gatherers, who may represent a more appropriate comparative sample than medieval individuals for inferring mechanical stress due to locomotor adaptation in past populations. Data on the humerus, femur and tibia suggest that agricultural populations tend to develop more pronounced asymmetry in bones associated with increasing sedentism and repetitive or unilateral tasks related to farming activities (e.g., Bridges et al., 1989; Auerbach and Ruff, 2006; Ruff et al., 2006; Wescott and Cunningham, 2006; Shaw and Stock, 2011; Sládek et al., 2016; Kubicka et al., 2018; Sikdar and Mushrif-Tripathy, 2023). Conversely, hunter-gatherers often display greater symmetry due to more variable or ambidextrous activities (Kubicka et al., 2018). These studies demonstrate that the mode of subsistence alters the distribution of loads and laterality in weight-bearing bones (humerus, femur, tibia). Although there are no direct studies on MT1 or MT5 asymmetry comparing hunter-gatherers and agricultural populations, the effects of unilateral activity could also be visible in the metatarsals given the agricultural activities in our comparative sample (Kubicka et al., 2018).

28It was found that approximately 76% of present-day humans prefer to use their right foot for activities involving a single foot, such as kicking a ball or stamping on an imagined insect (Gentry and Gabbard, 1995). This indicates that the left foot is more frequently weight-bearing and therefore subjected to greater stress in this context. There are two main limitations of our results that should be discussed. First, when testing after removing individuals showing extreme values, we did not find any significant side dominance, indicating an intra-group effect in our sample. DA appears to be more prevalent at the individual level, potentially reflecting differences in intrinsic foot musculature rather than a common pattern of pedal pressure, as observed in non-human catarrhines (Patel, 2024). Furthermore, recent research has indicated a strong phylogenetic but weak functional signal in MT1 among different species of primates, suggesting that locomotor behaviours are only partially reflected in the form-function relationships of key skeletal elements (Tomizawa et al., 2024). Secondly, no division by sex was made in the analyses due to our small sample size, which could have influenced the results, since it has been shown that the emergence and intensification of agricultural practices are accompanied by a more pronounced sexual and functional division of labour (Murdock and Provost, 1973; Masclans et al., 2021). Taken together, these findings suggest that metatarsal morphology is influenced by factors beyond mechanical stress. This has also been observed in other skeletal elements, such as femurs, where genetic (e.g., Hansen et al., 2009), phylogenetic (e.g., Morimoto et al., 2012; Cosnefroy et al., 2024a), sex-specific activity patterns (e.g., Ruff, 1987; Shaw and Stock, 2011; Laffranchi et al., 2020) and ontogenetic factors (e.g., Morimoto et al., 2018) have been identified as influences. Further research into metatarsal asymmetry in various primate species and diverse human groups is required to determine the factors influencing their bone structure.

29As in the medieval sample, LF1 and LF2 exhibited higher levels of DA and AA in MT1 than in MT5. However, the values for LF1 and LF2 were higher than those for the medieval sample. Neandertals differ from extant H. sapiens in having more robust bones, including robust lower limb bones, which has been interpreted as an adaptation to increased locomotor loading (Trinkaus, 1983; DeSilva et al., 2019; Pablos et al., 2019). In this context, the higher levels of asymmetry in MT1 in LF1 and LF2 may indicate more intense and/or repetitive foot loading than in the medieval sample. As in the medieval sample, however, we did not find a clear pattern of side dominance for LF1 and LF2. Rather, we found a right- or left-side bias depending on the measurement considered. Nevertheless, when we examined midshaft robusticity (i.e., the mediolateral and antero-posterior diameters at the midshaft), we found a right-side bias for LF2 and the medieval humans, indicating a more robust right MT1. LF1 showed a less clear pattern, with a more robust left MT1 in the antero-posterior direction and a more robust right MT1 in the mediolateral direction. It should be noted that the MT1s of LF1 are not well enough preserved to be studied in full, making it more difficult to define asymmetry patterns for this individual. In contrast, LF2 has more complete MT1s and differs from the medieval sample in having a larger head (i.e., distal epiphysis) on the left MT1; the medieval sample showed no such bias. In modern humans, the MT1 head is subject to high compressive forces during push-off (Rodgers, 1995; Donahue and Sharkey, 1999; Vereecke et al., 2003; D’Août et al., 2004) and is thought to enhance joint stability and facilitate close packing of the metatarsophalangeal joint during this stage (Susman and Brain, 1988; Hetherington et al., 1989; Susman and de Ruiter, 2004; Pontzer et al., 2010; Fernandez et al., 2015). In the light of our results, it is possible that the left foot of LF2 experienced greater loading during push-off than the right, and that this asymmetry may have affected gait, balance and overall stability, potentially reflecting habitual lateralized behaviours or adaptation to specific locomotor or environmental demands. Furthermore, as the foot appears to be correlated with handedness (e.g., Singh, 1970; Auerbach and Ruff, 2006), it is reasonable to ask whether LF1 and LF2 exhibit similar right- or left-side dominance in both upper and lower limbs. Bilateral asymmetry in the hand bones of Neandertals and other fossil hominins has not yet been explicitly investigated (Bardo et al., 2023), although such data could inform discussions of crossed symmetry patterns, side dominance and asymmetrical mechanical loading related to pathology. Conversely, analyses of upper-limb asymmetry in Neandertals have identified several markers consistent with right-handedness (e.g., Trinkaus, 1980; Trinkaus et al., 1994; Vandermeersch and Trinkaus 1995; Shaw et al., 2012), although De Groote (2011) highlighted substantial inter-individual variation among humans. Lower limb asymmetry in Neandertals has often been interpreted as relatively limited, consistent with more varied, bilateral or highly mobile activities, such as walking, hunting and gathering (e.g., Auerbach and Ruff, 2006; De Groote, 2011). Furthermore, the foot bones of LF1 and LF2, two relatively well-preserved specimens, warrant further study; to our knowledge, Heim’s monograph (1982) remains the only comprehensive description to date.

Asymmetry in the leg bones

30In the medieval sample, we found significant AA across all measurements for both the femur and the tibia across all biomechanical properties, indicating substantial bilateral differences. However, within this group, our results showed that the tibia exhibited greater DA than the femur across all biomechanical properties. This is consistent with evidence that the tibia is more responsive to habitual mechanical loading (Ruff et al., 1997; 2006; Trinkaus and Ruff, 1999; Macintosh and Stock, 2019). This pattern suggests that the tibia may be more sensitive than the femur to differences in activity or locomotor stress between limbs, reflecting the specific loading demands placed on the lower limb during daily behaviours. However, as Stock (2006) noted, only the polar second moment of area shows a clear correlation with mobility type. This suggests that additional factors, such as genetic influences or activity-specific loading patterns, may also contribute to the shape and asymmetry of the tibial shaft.

31In Neandertals and Pleistocene H. sapiens, this pattern appears to be similar: lower limb-bones, particularly the tibia, exhibit structural adaptations consistent with high and repetitive biomechanical loading (Ruff et al., 1997; 2006; Trinkaus and Ruff, 1999). Cross-sectional geometry analyses of Neandertal tibiae, for example, show greater cortical thickness and bending rigidity relative to the femur, suggesting that the tibia may have been particularly sensitive to habitual locomotor stress. This could reflect frequent ambulation over varied terrain or high-intensity terrestrial mobility (Lovejoy and Trinkaus, 1980; Trinkaus and Ruff, 1989; Ruff et al., 1997; Kubicka et al., 2022), and underscores the fact that differences in plasticity and responsiveness to loading between bones are not unique to modern humans, but probably represent a broader feature of hominin lower limb adaptation.

32Interestingly, when examining DA, tibial cross-sections closer to the epiphyses exhibited greater asymmetry than those near the midshaft. According to Pearson (2000), functional plasticity varies along bones, with the articular ends generally being less responsive to mechanical loading than the shafts. Despite this, we did not detect a consistent DA pattern in tibiae. Examination of cortical area (%CA), which reflects the magnitude of habitual loading during life, revealed a right bias at the proximal CS and a left bias at the distal CS; however, statistical analyses suggest that these biases are largely driven by a few outlier individuals. Further analyses illustrate this variability. The polar moment of area (J), which indicates resistance to bending and torsional rigidity, showed a right bias at the distal shaft of the tibiae. In contrast, Imax/Imin, which reflects cross-sectional shape, showed a left bias at the distal CS and at the midshaft when outliers were included. Medieval humans displayed only a significant left bias for femoral J at 35% of biomechanical length, and at midshaft when outliers were removed. The sample studied here mostly comprises farmers (Kubicka et al., 2018), whose mobility levels may not have been sufficient to exert a strong influence on DA in the lower limb. This same group does, however, exhibit pronounced DA in the upper limb, reflecting habitual unimanual activities (Kubicka et al., 2016; 2018). However, the absence of strong directional patterns in lower limb remodelling or DA is not necessarily specific to farming; it may also reflect an increasing homogeneity of activity patterns in more recent populations compared with older ones (Ruff, 1987). To further explore the influence of mobility on DA in the lower limbs, future studies should include larger and more diverse samples, including Pleistocene H. sapiens, as has been done in analyses of lower limb morphology (e.g., Shackelford and Trinkaus, 2002; Trinkaus and Ruff, 2012; Shaw and Stock, 2013; Cosnefroy, 2025).

33In this study, we present both newly generated data and revised values for the two La Ferrassie Neandertals (table 4). The differences between our results and those reported by Trinkaus and Ruff (2012) most likely reflect differing methodologies rather than biological variation, as the use of high-resolution CT data allows more accurate identification of cortical boundaries in fossil material affected by sediment infilling and post-depositional alterations. LF1 and LF2 display higher DA than the medieval comparative sample, indicating that these individuals most likely experienced more intensive or repetitive loading of the lower limbs. Interestingly, LF1 and LF2 differ in both the magnitude and direction of asymmetry, which could reflect sex-based differences. Specifically, the greater overall robustness observed in LF1 (male) relative to LF2 (female) might signal sexual dimorphism in biomechanical properties, consistent with patterns documented in both modern humans and Neandertals (Trinkaus, 1983; Ruff, 1987; Ruff et al., 1993; Sládek et al., 2016). However, given the extremely limited sample size, these differences should be interpreted with caution as they could also arise from individual variability, habitual activity or pathological conditions. Several Neandertal individuals do indeed demonstrate pronounced asymmetry or unusual robusticity associated with trauma or degenerative conditions. For example, Shanidar 1 exhibits extensive remodelling linked to repeated injury (Ivanhoe and Trinkaus, 1983), and La Chapelle-aux-Saints shows atypical lower-limb robusticity probably due to osteoarthritis and compensatory loading (Trinkaus, 1985; Haeusler et al., 2019). Such cases suggest that LF2’s higher DA may result partly from compensatory responses to injury or a localized pathology rather than from purely behavioural or sexual dimorphism effects. Notably, sexual dimorphism in asymmetry tends to be more evident in the upper limb, while the lower limb in modern humans generally shows only modest differences between sexes (Auerbach and Ruff, 2006). Analysis of cortical area (%CA) indicates that LF2 had a more robust left lower limb, whereas LF1 was more robust on the right, although this pattern is less clear for J and Imax/Imin. Overall, LF2 exhibits higher DA magnitudes than LF1, which may reflect variation in activity patterns or pathological influences. Finally, comparing upper- and lower-limb asymmetry following Auerbach and Ruff (2006) could reveal a crossed-symmetry pattern, in which right-biased asymmetry in the upper limb corresponds to left-biased asymmetry in the lower limb, as commonly observed in modern humans (Plochocki, 2002; 2004; Auerbach and Ruff, 2006; Eriksen, 2020).

Table 4

Table 4

Non-standardized femoral and tibial biomechanical properties of La Ferrassie 1 and La Ferrassie 2 compiled from the sectional measurements (CS20%, CS35%, etc.) published by Trinkaus and Ruff (2012) and from the measurements collected in this study using high-resolution CT scans |
Propriétés biomécaniques non standardisées des fémurs et des tibias de La Ferrassie 1 et La Ferrassie 2 ont été établies à partir des mesures sectionnelles (CS20%, CS35%, etc.) publiées par Trinkaus et Ruff (2012), ainsi que les mesures obtenues dans la présente étude à partir de scans tomographiques à haute résolution

Directional asymmetry across the lower limb bones

34No correlations were found between tibiae, femora and the metatarsals. In LF1 and LF2, the long bones are incompletely preserved, preventing direct measurement of femoral and tibial epiphyseal lengths and widths. Consequently, biomechanical properties provided more informative data than linear measurements for these specimens. According to the bone functional adaptation concept, bone tissue remodels throughout life to resist and accommodate local mechanical loading (Currey, 2002; Ruff et al., 2006), suggesting that both external and internal morphology adapt together. However, external bone shape, particularly articular surfaces, is generally less plastic than internal bone structure (Ruff et al., 1991; Lieberman et al., 2001; but see also Karakostis et al., 2017). The lack of correlation between the bones analysed may reflect differences in mechanical constraints between the leg and the foot, leading to distinct functional signals. Additionally, biomechanical properties and overall morphology may respond to different magnitudes of mechanical stimuli (Kubicka and Myszka, 2020), so that changes in these two skeletal markers may not occur in parallel. Very few studies have examined internal and external bone structure jointly (Chapman et al., 2018; Kubicka and Myszka, 2020; Cosnefroy et al., 2024b; Tanner et al., 2024), which underlines the need for further research to determine which structural features, or combinations thereof, best capture bilateral asymmetry.

Conclusion

35This study integrates long-bone cross-sectional geometry with metatarsal measurements to evaluate bilateral asymmetry across multiple lower-limb elements within the same individuals. Across the medieval sample, asymmetry was present but showed no consistent directional pattern, highlighting substantial inter-individual variability. The two La Ferrassie individuals generally exhibited higher asymmetry values than the comparative sample, but with no uniform side dominance, suggesting that lower-limb asymmetry may reflect a combination of loading history, individual behaviour, and possibly sex- or health-related factors. These results emphasize the value of a multi-element approach for interpreting locomotor loading in past populations and point to the need for broader comparative analyses using larger samples, subsistence contrasts and independent evidence for pathology.

Supplementary information

36Supplementary table 1. Results of the Wilcoxon signed rank tests used to test for significance of both indicators (%DA and %AA) for each of the seven measurements for the MT1 and MT5. Statistically significant results are shown in bold | Résultats des tests de Wilcoxon utilisés pour tester la signification statistique des deux indicateurs (%DA et %AA) pour chacune des sept mesures pour le MT1 et le MT5. Les résultats statistiquement significatifs sont indiqués en gras

Table S1a

Table S1a

Results including all individuals |
Résultats incluant tous les individus

Table S1b

Table S1b

Results after exclusion of outliers (NA indicates measurements for which no outliers were detected) |
Résultats après exclusion des valeurs aberrantes (NA indique les mesures pour lesquelles aucune valeur aberrante n’a été détectée)

37Supplementary table 2. Results of the Wilcoxon signed rank tests used to test for significance of the biomechanical properties (%CA, J, Imax/Imin) in each of five CS for the femur and the tibia. Statistically significant results are shown in bold | Résultats des tests de Wilcoxon utilisés pour tester la signification statistique des propriétés biomécaniques (%CA, J, Imax/Imin) dans chacun des cinq CS pour le fémur et le tibia. Les résultats statistiquement significatifs sont indiqués en gras

Table S2a

Table S2a

Results including all individuals |
Résultats incluant tous les individus

Acknowledgements: We thank Anna Wrzesińska from the Museum of the First Piasts at Lednica for allowing us to use skeletal material from Ostrów Lednicki. Thanks also to Szymon Stelting from Poznań University of Life Sciences for technical support in scanning the osteological material, and to Prof. Dominique Grimaud-Hervé from the MNHN in Paris for giving us access to the CT-images of LF1 and LF2. This project was funded by the National Science Centre in Poland (AK; grant number: NCN 2015/19/N/NZ8/00177), the French Research Agency (AK and AB; project "Bringing the brain of Homo erectus and Neandertals back to life", ANR-20-CE27-0009-01), and the ERC-Adv. LATEUROPE project (AB; Grant agreement ID [101052653]).

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Bibliographie

Results after exclusion of outliers (NA indicates measurements for which no outliers were detected) |
Présente les résultats après exclusion des valeurs aberrantes (NA indique les mesures pour lesquelles aucune valeur aberrante n’a été détectée)

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Table des illustrations

Titre Figure 1
Légende Measurements taken on the left MT1 and MT5. The abbreviations used are described in table 1 |Mesures prises sur les MT1 et MT5 gauches. Les abréviations utilisées sont décrites dans le tableau 1
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-1.png
Fichier image/png, 62k
Titre Table 1
Légende Description of linear measurements of the MT1 and MT5 (after Barrio et al., 2006) |Description des mesures linéaires réalisées sur les MT1 et MT5 (d’après Barrio et al., 2006)
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-2.png
Fichier image/png, 30k
Titre Figure 2
Légende Illustration of the five cross-sections (CS) analysed at different percentages of the length of a right femur (left) and a right tibia (right) of a medieval individual from Ostrów Lednicki |Illustration des cinq coupes transversales (CS) analysées à différents pourcentages de la longueur d’un fémur droit (à gauche) et d’un tibia droit (à droite) d’un individu médiéval provenant d’Ostrów Lednicki
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-3.png
Fichier image/png, 164k
Titre Figure 3
Légende Reconstructions of the LF1 (A) and LF2 (B) lower bones, femurs at the top and tibias below, with illustration of the CS analysed |Reconstitutions des os long des jambes de LF1 (A) et LF2 (B), les fémurs en haut et les tibias en bas, avec une illustration des sections transversales analysées
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-4.png
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URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-5.png
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URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-6.png
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Titre Table 2
Légende Descriptive statistics (averages±standard errors) for each measurement on the MT1 and MT5, presented for the medieval sample as a whole, and for males (M) and females (F) separately, with percentage of directional asymmetry (%DA) and percentage of absolute asymmetry (%AA) |Statistiques descriptives (moyenne±erreurs-standards) pour chaque mesure sur les MT1 et les MT5 présentés pour l’échantillon médiéval dans son ensemble, et séparément pour les hommes (M) et les femmes (F), avec le pourcentage d’asymétrie directionnelle (%DA) et le pourcentage d’asymétrie absolue (%AA)
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-7.png
Fichier image/png, 480k
Titre Table 3
Légende Mean percentage of directional asymmetry (%DA) and absolute asymmetry (%AA), ±standard errors, for the three biomechanical properties (%CA, Imax/Imin, J) for the five cross-sections (CS) of the femur and the tibia for the medieval humans (M: males; F: females), and values for LF1 and LF2 from this study and a previous one by Trinkaus and Ruff (2012). The bold values are new LF2 measurements not presented in Trinkaus and Ruff (2012) |Pourcentage moyen d’asymétrie directionnelle (%DA) et d’asymétrie absolue (%AA), ±erreurs standards, pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du fémur et du tibia pour les humains médiévaux (M : mâles ; F : femelles), et les valeurs pour LF1 et LF2 qui proviennent de cette étude et d’une précédente étude par Trinkaus et Ruff (2012). Les valeurs en gras sont de nouvelles mesures pour LF2 qui ne sont pas présentées dans Trinkaus et Ruff (2012)
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-8.png
Fichier image/png, 1,2M
Titre Figure 4
Légende Percentage of directional asymmetry (%DA) for the seven linear measurements (see table 1) of the MT1 (A) and MT5 (B) for the medieval sample and LF1 and LF2. A significant level of DA was found for the mediolateral diameter of the proximal epiphysis (ML PE) and the antero-posterior diameter at the midshaft (AP midshaft) of the MT1 (* p<0.05). Each box plot shows the median value (bold line) and the interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-bias, while negative values indicate a left-bias |Pourcentage d’asymétrie directionnelle (%DA) pour les sept mesures linéaires (voir tableau 1) du MT1 (A) et du MT5 (B) pour l’échantillon médiéval et LF1 et LF2. Un niveau significatif de DA a été trouvé pour le diamètre médiolatéral de l’épiphyse proximale (ML PE) et le diamètre antéro-postérieur au milieu de la diaphyse (AP midshaft) du MT1 (* p<0,05). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "valeurs aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-9.png
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Titre Figure 5
Légende Percentage of directional asymmetry (%DA) for the three biomechanical properties (%CA, Imax/Imin, J) across five cross-sections (CS) of the femur in the medieval humans and in LF1 and LF2. A significant %DA was found for J at CS 35% (** p<0.01). Each box plot shows the median value (bold line) and the interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-side bias, while negative values indicate a left-side bias |Pourcentage d’asymétrie directionnelle (%DA) pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du fémur, chez les humains du médiéval et LF1 et LF2. Un pourcentage significatif de DA a été observé pour J à CS 35 % (** p<0.01). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "données aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-10.png
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Titre Figure 6
Légende Percentage of directional asymmetry (%DA) c (%CA, Imax/Imin, J) for the five cross-sections (CS) of the tibia for the medieval humans and LF1 and LF2. A significant %DA was found for Imax/Imin (CS 50% and CS 80%) and J (CS 65% and CS 80%) (*** p<0.001; ** p<0.01; * p<0.05). Each box plot shows the median value (bold line) and interquartile range of %DA. Vertical lines indicate variability outside the upper and lower quartiles, except for ‘‘outliers’’ (dots). Positive values indicate a right-side bias, while negative values indicate a left-side bias |Pourcentage d’asymétrie directionnelle (%DA) pour les trois propriétés biomécaniques (%CA, Imax/Imin, J) pour les cinq sections transversales (CS) du tibia, chez les humains du médiéval et LF1 et LF2. Un pourcentage significatif de DA a été observé pour Imax/Imin (CS 50 % et CS 80 %) et J (CS 65 % et CS 80 %) (*** p<0,001; ** p<0,01; * p<0,05). Chaque boîte à moustache indique la valeur médiane (ligne en gras) et les intervalles interquartiles de %DA. Les lignes verticales indiquent la variabilité à l’extérieur des quartiles supérieur et inférieur, sauf pour les "données aberrantes" (points). Les valeurs positives expriment un biais à droite tandis que les valeurs négatives expriment un biais à gauche
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-11.png
Fichier image/png, 115k
Titre Table 4
Légende Non-standardized femoral and tibial biomechanical properties of La Ferrassie 1 and La Ferrassie 2 compiled from the sectional measurements (CS20%, CS35%, etc.) published by Trinkaus and Ruff (2012) and from the measurements collected in this study using high-resolution CT scans |Propriétés biomécaniques non standardisées des fémurs et des tibias de La Ferrassie 1 et La Ferrassie 2 ont été établies à partir des mesures sectionnelles (CS20%, CS35%, etc.) publiées par Trinkaus et Ruff (2012), ainsi que les mesures obtenues dans la présente étude à partir de scans tomographiques à haute résolution
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-12.png
Fichier image/png, 884k
Titre Table S1a
Légende Results including all individuals |Résultats incluant tous les individus
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-13.png
Fichier image/png, 196k
Titre Table S1b
Légende Results after exclusion of outliers (NA indicates measurements for which no outliers were detected) | Résultats après exclusion des valeurs aberrantes (NA indique les mesures pour lesquelles aucune valeur aberrante n’a été détectée)
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-14.png
Fichier image/png, 183k
Titre Table S2a
Légende Results including all individuals |Résultats incluant tous les individus
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-15.png
Fichier image/png, 207k
Titre Table S2b
URL http://journals.openedition.org/bmsap/docannexe/image/17966/img-16.png
Fichier image/png, 200k
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Référence électronique

Ameline Bardo et Anna Maria Kubicka, « Bilateral asymmetry in the lower limbs in medieval humans and La Ferrassie Neandertals LF1 and LF2 »Bulletins et mémoires de la Société d’Anthropologie de Paris [En ligne], 38 (1) | 2026, mis en ligne le 21 avril 2026, consulté le 16 mai 2026. URL : http://journals.openedition.org/bmsap/17966 ; DOI : https://doi.org/10.4000/164bt

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Auteurs

Ameline Bardo

UMR 7194 - HNHP, CNRS-MNHN, Département Homme et Environnement, Musée de l’Homme, Paris, France ; Department of Human Origins, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany ; https://orcid.org/0000-0003-1840-6423 ; ameline.bardo[at]mnhn.fr

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Anna Maria Kubicka

UMR 7194 - HNHP, CNRS-MNHN, Département Homme et Environnement, Musée de l’Homme, Paris, France ; Department of Zoology, Poznań University of Life Sciences, Poznań, Poland ; https://orcid.org/0000-0002-7844-9225 ; amkkubicka[at]gmail.com

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