Footedness in merlins: Raptors perching in a cold climate.

ABSTRACTPerching or standing on one foot is commonly reported in birds but the level of consistency in using one foot over the other has been less-well documented in most species, particularly birds of prey. For birds experiencing colder temperatures, unipedal perching has been attributed to limiting heat loss through unfeathered legs and feet; individuals should spend longer periods of time perched on one foot as temperatures decrease. Using radio tracking, I collected 486 hours of observations on nine overwintering, free-living merlins (Falco columbarius) in Saskatoon, Canada. Five merlins displayed clear preferences to perch on one foot, however the direction of preference was not consistent and four birds were ambidextrous. There was a curvilinear response in the proportion of time spent in unipedal posture versus temperature, with a peak of ∼22% of the time at moderate temperatures (-10 to -19°C), but lower values at warmer and colder temperatures; the main effect of the squared term for temperature was highly influential while individual foot preference had no impact on the use of unipedal perching. Although preferential use of one foot for perching was displayed by some individuals, thermoregulation may not be the primary driver of this behaviour at colder temperatures.

Behavioral laterality is correlated with problem-solving performance in a songbird.

Cerebral lateralization, which is often reflected in an individual's behavioral laterality (e.g., handedness and footedness), may bring animals certain benefits such as enhanced cognitive performance. Although the lateralization-cognition relationship has been widely studied in humans and other animals, current evidence supporting their relationship is ambiguous and warrants additional insights from more studies. Moreover, the lateralization-cognition relationship in non-human animals has been mostly studied in human-reared populations, and investigations of wild populations are particularly scarce. Here, we test the footedness of wild-caught male yellow-bellied tits (Pardaliparus venustulus) and investigate its association with their performance in learning to solve a toothpick-pulling problem and a drawer-opening problem. The tested birds showed an overall trend to gradually spent less time solving the problems, implying that they learned to solve the problems. Left- and right-footed individuals showed no significant differences in the latency to explore the experimental apparatuses and in the proportions that completed and did not complete the tasks. However, the left-footed individuals learned faster than the right-footed individuals in the drawer-opening experiment, indicating a potential cognitive advantage associated with left-footedness. These results contribute to the understanding of the behavioral differences between differently footed individuals and, in particular, the relationship between lateralization and cognitive ability in wild animals.

Footedness for scratching itchy eyes in rodents.

The neural bases of itchy eye transmission remain unclear compared with those involved in body itch. Here, we show in rodents that the gastrin-releasing peptide receptor (GRPR) of the trigeminal sensory system is involved in the transmission of itchy eyes. Interestingly, we further demonstrate a difference in scratching behaviour between the left and right hindfeet in rodents; histamine instillation into the conjunctival sac of both eyes revealed right-foot biased laterality in the scratching movements. Unilateral histamine instillation specifically induced neural activation in the ipsilateral sensory pathway, with no significant difference between the activations following left- and right-eye instillations. Thus, the behavioural laterality is presumably due to right-foot preference in rodents. Genetically modified rats with specific depletion of Grpr-expressing neurons in the trigeminal sensory nucleus caudalis of the medulla oblongata exhibited fewer and shorter histamine-induced scratching movements than controls and eliminated the footedness. These results taken together indicate that the Grpr-expressing neurons are required for the transmission of itch sensation from the eyes, but that foot preference is generated centrally. These findings could open up a new field of research on the mechanisms of the laterality in vertebrates and also offer new potential therapeutic approaches to refractory pruritic eye disorders.

Relationships between footedness and aging on postural control: Evidence from the Yakumo study.

ABSTRACTThe effects of footedness and aging on postural maintenance function were examined in this study. Using the postural maintenance task, the trajectories of the centre of pressure (COP) were measured with a stabilometer in the lateral and posterior/anterior directions. One hundred and ninety-three right-footers (112 middle-aged: 40-64 years old and 81 old-aged: 65-80 years old) and thirty-two left-footers (14 middle-aged and 18 old-aged) among healthy community members participated in this study. Using the ratio of the COP deviation area sizes for the open- vs. closed-eye conditions as an index, the ratios in the lateral and the posterior/anterior directions were compared in relation to the footedness and the age groups. The results indicated that the left-footers of the older adults group showed a larger COP deviation ratio than that of the middle-aged group, and the left-footers showed a larger deviation ratio in the lateral direction than in the posterior-anterior direction. These findings suggest a weak postural maintenance function of the older adults left footers. As most left-footers are left-handed, health promotion staff should notice that older adults who are left-handers are at a higher risk of falling.

Cultural and environmental aspects influence lateral preferences.

Transcultural approaches comparing the distribution of lateral preference between countries focused primarily on handedness. However, other laterality dimensions may also be susceptible to cultural variations. The present study compared lateral preference in five dimensions (hand, foot, trunk, hearing, and visual) of young adults from Brazil and Mozambique. Two hundred and two undergraduate students participated in the study, of which 101 were Brazilian (21.7 ± 1.66 years old) and 101 Mozambican (25.6 ± 6.2 years old). The participants' lateral preference direction and degree were assessed using the global lateral preference inventory. Most Brazilians were classified in the moderate preference category, while Mozambicans had a higher frequency of individuals with strong preferences. Hence, Mozambicans presented a higher lateralization degree for all dimensions. A subsequent analysis considering only preference direction (right, left or indifferent), and not degree, did not show the same outcomes, revealing similarities in preference distribution between the two groups. This finding highlights the necessity to incorporate preference degrees in future analysis to not overlook potential differences. We also conclude lateral preference investigations with transcultural approaches should analyse beyond handedness, as other dimensions can also be modulated by cultural characteristics.

Cross-cultural adaptation, validity and reliability of Turkish version of the Waterloo Handedness and Footedness Questionnaire-Revised.

This study investigated the validity and reliability of Turkish versions of the Waterloo Handedness and Footedness Questionnaire-Revised (WHQ-R and WFQ-R). Turkish versions of the WHQ-R and the WFQ-R, and other assessments (the Handedness Questionnaire and the Footedness Preference Test) were applied to 444 healthy participants aged between 18 and 65 years. The translation and cultural adaptation process of the WHQ-R and the WFQ-R was provided by the following proposed guideline. For test-retest reliability, translated versions were re-applied to 88 participants in a 7-day interval. Cronbach's alpha of the WHQ-R was 0.984 and that of the WFQ was 0.905. The test-retest intra-class correlation coefficients (ICC) of the WHQ-R varied from 0.741 to 0.933 and those of the WFQ-R from 0.649 to 0.814. There were moderate to strong correlations between the WHQ-R and the Handedness Questionnaire (r = -853, p < 0.001) and between the WFQ-R and the Footedness Preference Test (r = -0.687, p < 0.001). Confirmatory factor analysis revealed a two-factor structure of the WHQ-R and the WFQ-R. The results of this study demonstrated that Turkish versions of the WHQ-R and the WFQ-R are reliable and valid inventories for assessing handedness and footedness in this population.

The dominant leg is more likely to get injured in soccer players: systematic review and meta-analysis.

In soccer (football), dominant limb kicking produces higher ball velocity and is used with greater frequency than the non-dominant limb. It is unclear whether limb dominance has an effect on injury incidence. The purpose of this systematic review with meta-analysis is to examine the relationship between limb dominance and soccer injuries. Studies were identified from four online databases according to PRISMA guidelines to identify studies of soccer players that reported lower extremity injuries by limb dominance. Relevant studies were assessed for inclusion and retained. Data from retained studies underwent meta-analyses to determine relative risk of dominant versus non-dominant limb injuries using random-effects models. Seventy-four studies were included, with 36 of them eligible for meta-analysis. For prospective lower extremity injury studies, soccer players demonstrated a 1.6 times greater risk of injury to the dominant limb (95% CI [1.3-1.8]). Grouped by injury location, hamstring (RR 1.3 [95% CI 1.1-1.4]) and hip/groin (RR 1.9 [95% CI 1.3-2.7]) injuries were more likely to occur to the dominant limb. Greater risk of injury was present in the dominant limb across playing levels (amateurs RR 2.6 [95% CI 2.1-3.2]; youths RR 1.5 [95% CI 1.26-1.67]; professionals RR 1.3 [95% CI 1.14-1.46]). Both males (RR 1.5 [95% CI 1.33-1.68)] and females (RR 1.5 [95% CI 1.14-1.89]) were more likely to sustain injuries to the dominant limb. Future studies investigating soccer injury should adjust for this confounding factor by using consistent methods for assigning limb dominance and tracking use of the dominant versus non-dominant limb.

No evidence that footedness in pheasants influences cognitive performance in tasks assessing colour discrimination and spatial ability.

The differential specialization of each side of the brain facilitates the parallel processing of information and has been documented in a wide range of animals. Animals that are more lateralized as indicated by consistent preferential limb use are commonly reported to exhibit superior cognitive ability as well as other behavioural advantages. We assayed the lateralization of 135 young pheasants (Phasianus colchicus), indicated by their footedness in a spontaneous stepping task, and related this measure to individual performance in either 3 assays of visual or spatial learning and memory. We found no evidence that pronounced footedness enhances cognitive ability in any of the tasks. We also found no evidence that an intermediate footedness relates to better cognitive performance. This lack of relationship is surprising because previous work revealed that pheasants have a slight population bias towards right footedness, and when released into the wild, individuals with higher degrees of footedness were more likely to die. One explanation for why extreme lateralization is constrained was that it led to poorer cognitive performance, or that optimal cognitive performance was associated with some intermediate level of lateralization. This stabilizing selection could explain the pattern of moderate lateralization that is seen in most non-human species that have been studied. However, we found no evidence in this study to support this explanation.

Footedness related differences in femoral bone mineral density in elderly women with osteoporosis.

Purpose: The presumed link between lateralized habitual motor activity and bilateral skeletal asymmetry, as result of bone functional adaptation, is the basis for inferences about bone mineral distribution; consequently, bone loss and osteoporosis. As there is no research connecting directly footedness with osteoporosis, the present study was designed to investigate their relation, in elderly women with osteoporosis.Methods: A sample of 420 white, Caucasian race women, older than 65 years, with previous diagnosis of osteoporosis, was recruited for the study. Finally, 164 women met the inclusion criteria and participated. The femoral neck and total hip bone mineral density (BMD) differences between right and left hips of 105 right-footed and 59 non-right-footed women, and the between footedness groups differences of the above right-left BMD differences, were measured with Dual-energy X-ray Absorptiometry (DXA).Results: In right-footers, total hip right-left BMD differences were statistically significant, favoring left femur [t(104) = -2.79, p < .01, two tailed]. In non-right-footers, femoral neck right-left BMD differences were statistically significant, favoring right femur [t(58) =2.025, p < .05, two tailed]. Femoral neck and total hip right-left BMD differences of right- versus non-right-footed women were statistically insignificant.Conclusion: The dominance of the right lower limb had an asymmetric effect on left total hip BMD, while non-right-footedness favored right femoral neck BMD. The findings highlighted the influence of load bearing effects of the body weight and muscle contractions on femoral BMD. We noted a possible association between footedness and BMD, as a result of bone functional adaptation.

Associations of Bisexuality and Homosexuality with Handedness and Footedness: A Latent Variable Analysis Approach.

Non-right-handedness appears to be more common among bisexuals and homosexuals than among heterosexuals, which might be indirect evidence of effects of prenatal androgen exposure. Current data suggest higher prenatal testosterone levels among bisexual and homosexual women, but are inconclusive for men. This study examined the association between sexual orientation and non-right-handedness for sex differences and whether higher rates of mixed-handedness, rather than left-handedness, might be the driving factor. This allowed for more specific tests regarding the predictions of two competing theories of prenatal androgen exposure, the Geschwind-Galaburda theory and the callosal hypothesis, than in previous research. Being a potentially better indicator of cerebral lateralization than handedness, associations with footedness were also explored. To counter inconsistencies and shortcomings of previous research, we utilized two large discovery and replication datasets (ns = 2368 and 1565) and applied latent variable analysis to reliably classify lateral preferences (i.e., handedness, footedness). This maximized the statistical conclusion validity and allowed for direct tests of replicability. Sexual orientation was differentially associated with lateral preferences among men and women. Associations among women were consistent with predictions of the Geschwind-Galaburda theory, whereas among men they were consistent with predictions of the callosal hypothesis. The results were further consistent with models of homosexuality that suggest a role of parental epigenetic marks on sexually dimorphic fetal development. Research efforts should be increased with regard to footedness and epigenetic theories of homosexuality.

Beyond handedness: assessing younger adults and older people lateral preference in six laterality dimensions.

Lateral preference is influenced by many different factors. Although studies that assessed handedness contributed greatly to our understanding of the phenomenon, looking to other dimensions of motor laterality, such as lower limbs, trunk, visual and hearing preference, may be an interesting strategy to further advance in the field. Comparing different age groups with a complete inventory also contributes to understanding the ageing impact on these variables. Our aim, therefore, was to assess six motor laterality dimensions in younger adults and older people. Two hundred subjects participated in this study (102 adults, 30.6 ± 11.2 years old, and 98 elders, 70.4 ± 7.22 years old) and lateral preference was assessed by the Global Lateral Preference Inventory. We verified significant differences between groups in all dimensions, except the visual one, with proportionally stronger right preference in the older group. Compared to other dimensions, higher frequency of ambidexterity was observed in trunk, hearing and visual preference for both groups. These results might indicate that lateral preference is, indeed, multifactorial, being affected by ageing, task complexity and other possible related aspects.

Paw preferences in the Asian small-clawed otter - using an inexpensive, video-based protocol to study laterality of rare species in the zoo.

It is still debated whether limb-use preferences represent a common trait in vertebrates, which is based on a shared phylogenetic history. Unravelling the evolutionary origin and pattern of paw preferences in vertebrates requires the analysis of a larger number of species within an ecologically relevant setting. We therefore investigated whether observations in a zoo enable the collection of reliable data sets by quantifying paw use in two independent groups of Asian small-clawed otters (Amblonyx cinerea). Employing a continuous focal animal sampling method, each day one of the ten individuals was video recorded from outside of the enclosure during usual activity. We selected four types of unimanual behaviour (reaching for food, reaching for non-food, reaching into a hole, carrying an object) and quantified paw use for each animal. Our study provides first evidences for individual paw preferences in otters, which were in line with previously reported forelimb use pattern in carnivoran species. Preferences differed between motor acts but for "reaching into a hole" a population-level right paw bias was detected. These data support that observations in a zoological setting are useful to explore task-dependent paw preferences and may facilitate future studies investigating paw preferences under experimentally controlled conditions.

Foot-preference underlies bite-scar asymmetry in the gecko Ptyodactylus guttatus.

Scar-asymmetry may reflect brain laterality because scar location may reflect behaviour when being attacked. This has been studied in a few organisms, but never in lizards. Wild geckos (Ptyodactylus guttatus) from Israel were examined for bite-scar numbers and their lateral asymmetry. Social status was documented in the field before capture. Foot-preference, for either the right (R-footed) or left (L-footed) hind leg was determined in six trials for each gecko on adults captured in Jerusalem. I studied 48 geckos: 15 R-footed, 6 ambidextrous and 9 L-footed females; 6 R-footed, 3 ambidextrous and 9 L-footed males. Adults showed significantly more bite-scars than juveniles. The proportion of L-footed males mirrored that of R-footed females. Ambidextrous and L-footed geckos had a higher social status. In males, R-footed individuals had more bite-scars on the right side of the body, while L-footed individuals had more on the left side. R-footed females had more bite-scars on the right side, while L-footed females had bite-scars equally on both sides. Bite-scar asymmetry correlated with hind-leg preference, clearly reflecting brain laterality. Since all ambidextrous males had high social status, that may be the driving factor behind the females' different bite-scar pattern.

Translation, cultural adaption, and test-retest reliability of Chinese versions of the Edinburgh Handedness Inventory and Waterloo Footedness Questionnaire.

Quantitative assessments of handedness and footedness are often required in studies of human cognition and behaviour, yet no reliable Chinese versions of commonly used handedness and footedness questionnaires are available. Accordingly, the objective of the present study was to translate the Edinburgh Handedness Inventory (EHI) and the Waterloo Footedness Questionnaire-Revised (WFQ-R) into Mandarin Chinese and to evaluate the reliability and validity of these translated versions in healthy Chinese people. In the first stage of the study, Chinese versions of the EHI and WFQ-R were produced from a process of translation, back translation and examination, with necessary cultural adaptations. The second stage involved determining the reliability and validity of the translated EHI and WFQ-R for the Chinese population. One hundred and ten Chinese participants were tested online, and the results showed that the Cronbach's alpha coefficient of internal consistency was 0.877 for the translated EHI and 0.855 for the translated WFQ-R. Another 170 Chinese participants were tested and re-tested after a 30-day interval. The intra-class correlation coefficients showed high reliability, 0.898 for the translated EHI and 0.869 for the translated WFQ-R. This preliminary validation study found the translated versions to be reliable and valid tools for assessing handedness and footedness in this population.

Age-related differences in Fukuda stepping and Babinski-Weil tests, within-day variability and test-retest reliability.

BACKGROUND: The Fukuda stepping and Babinski-Weil tests are associated with unperceived body rotation and linear displacements in young adults, but performance in older adults on these two tests has yet to be determined. AIM: The main purpose was to compare the performance and reliability of the Fukuda stepping and Babinski-Weil tests in young and older adults. METHODS: Fifty older and 50 young adults executed three trials of each test on day 1 (test) and day 2 (retest). Lateral and longitudinal displacements and body rotation relative to the starting position were measured. Means and standard deviations (SD) were compared between the two groups with Mann-Whitney tests. Test-retest reliability was assessed with intra-class correlation coefficients (ICC). Foot preference was determined from the score on the Waterloo Footedness Questionnaire and correlated with test scores. RESULTS: Lateral and longitudinal displacements were significantly larger in older than young participants on the Fukuda stepping test (p < .01) and significantly smaller on the Babinski-Weil test (p < .001). Older participants displayed a significantly smaller SD on the three Babinski-Weil test trials (p < .001). Displacement and rotation measures ICC ranged between 0.25 and 0.77 in older and between 0.58 and 0.80 in young participants. Foot preference correlated with rotation on the Fukuda stepping test in young (p < .05), but not in older participants (p > .05). DISCUSSION AND CONCLUSION: Linear displacements, but not body rotation, were different between older and young adults. There was no clear age-related differences in test-retest reliability, but the moderate reliability indicates that performance can vary from day to day in both age groups.

Effectiveness of pedalling retraining in reducing bilateral pedal force asymmetries.

Previous studies have been limited to describe asymmetries during pedalling and suggest possible repercussion on performance and/or injury risks. However, few studies have presented strategies to mitigate asymmetries. The purpose of this study was to assess the effectiveness of a pedalling retraining intervention to reduce bilateral pedal force asymmetries. Twenty cyclists were assessed and 10 enrolled in a pedalling retraining method receiving visual and verbal feedback of pedal forces. The asymmetry index was computed for comparison of bilateral peak pedal forces and used during retraining (12 trials at 70% of peak power). Significantly larger asymmetry was observed for asymmetrical cyclists at the first three trials (P < 0.01 and ES = 1.39), which was reduced when post-retraining was compared to measures from symmetrical cyclists (P = 0.69 and ES = 0.18). Cyclists with larger asymmetry (>20%) in bilateral pedal forces reduce their asymmetries using sessions of pedalling retraining and achieve asymmetry indices similar to symmetrical cyclists.

Asymmetry of Achilles tendon mechanical and morphological properties between both legs.

Although symmetry of Achilles tendon (AT) properties between legs is commonly assumed in research and clinical settings, different loading profiles of both legs in daily life (i.e., foot dominance) may affect the tendon properties in a side-depended manner. Therefore, AT properties were examined with regard to symmetry between legs. Thirty-six male healthy adults (28 ± 4 years), who were physically active but not involved in sports featuring dissimilar leg load participated. Mechanical and morphological AT properties of the non-dominant and dominant leg were measured by means of ultrasound, magnetic resonance imaging and dynamometry. The AT of the dominant leg featured a significant higher Young's modulus and length (P < 0.05) but a tendency toward lower maximum strain (P = 0.068) compared with the non-dominant leg. The tendon cross-sectional area and stiffness were not significantly different between sides. The absolute asymmetry index of the investigated parameters ranged from 3% to 31% indicating poor AT side symmetry. These findings provide evidence of distinct differences of AT properties between both legs in a population without any sport-specific side-depended leg loading. The observed asymmetry may be a result of different loading profiles of both legs during daily activities (i.e., foot dominance) and challenges the general assumption of symmetrical AT properties between legs.

Lower-limb dominance does not explain subject-specific foot kinematic asymmetries observed during walking and running.

Studies of human locomotion have observed asymmetries in lower-limb kinematics, especially at the more distal joints. However, it is unclear whether these asymmetries are related to functional differences between the dominant and non-dominant limb. This study aimed to determine the effect of lower-limb dominance on foot kinematics during human locomotion. Range of motion for the metatarsophalangeal joint (MPJ) and medial longitudinal arch (MLA), as well as time duration of windlass mechanism engagement, were recorded from healthy young adults (N = 12) across a range of treadmill walking and running speeds. On the group level, there were no differences in MPJ or MLA range of motion, or windlass engagement timing, between the dominant and non-dominant limb (p > 0.05). While not explained by limb dominance, between-limb differences in MPJ and MLA ranges of motion were observed for individual participants on the order of ∼2-6°, which could be clinically relevant or impact interpretation of research data.

Inventories of Human Lateral Preference: A Systematic Review.

There are a variety of inventories available to evaluate human lateral preference, but no previous review has systematically analyzed and compared them. We conducted a systematic literature review to identify these inventories and describe their characteristics (e.g., dimensions, scales, tasks, psychometric properties). We included 26 articles, each presenting a different inventory to assess lateral preference, published between 1900 and 2022, selected from the following databases: PubMed, SportDiscus, APA PsycNET, and Web of Science. These inventories analyzed tasks performed in everyday life (domestic, work-related, sports, and leisure/recreation activities), covering hand, foot, hearing, visual and other non-usual lateral preference dimensions, such as trunk and head. Most inventories classified individuals based on the direction (left, indifferent or right preference) and degree (consistent or moderate) of laterality. However, many of the reviewed inventories lack established psychometric verification in their original publication, such as validity, reliability, responsiveness, and practical applicability. Inventories also presented tasks that may be specific to a given cultural setting, limiting their application across different countries. Based on these findings, we provide a comprehensive guide for researchers to select a lateral preference assessment tool, but also advocate for a new inventory with a broader approach, containing more than one dimension of lateral preference (i.e., not only handedness) and tasks that are common to different cultures to enhance practical applicability worldwide.

The development of hand, foot, trunk, hearing, and visual lateral preference throughout the lifespan.

This research investigated the developmental process of five lateral preference dimensions (hand, foot, trunk, hearing, and visual preference). A total of 1236 volunteers participated in this study, divided into five age groups: 07-12 (n = 247); 13-17 (n = 234); 18-30 (n = 227); 31-60 (n = 225); and 61-90 years old (n = 303). Lateral preference was assessed via questionnaire with the Global Lateral Preference Inventory. By assessing the degree and direction of lateral preference in different ages, our results revealed a pattern of lateralization strengthening with aging in all the analyzed dimensions. We also verified significant correlation between hand preference and the other dimensions for all age groups, but correlation was stronger in the 7-12 group for all correlation pairs. Our results lead to the suggestion of an underlying general lateralization process in early ages (7-12 years old) followed by specific developmental trajectories of each preference dimension (13 years forward), likely startled by hemisphere and functional specialization related to innate developmental patterns of neural structures and social/environmental influences.