Biology Letters
Published:https://doi.org/10.1098/rsbl.2017.0446

    Abstract

    According to the fighting hypothesis, frequency-dependent selection gives relatively rarer left-handers a competitive edge in duel-like contests and is suggested as one mechanism that ensured the stable maintenance of handedness polymorphism in humans. Overrepresentation of left-handers exclusively in interactive sports seems to support the hypothesis. Here, by referring to data on interactive ball sports, I propose that a left-hander's advantage is linked to the sports’ underlying time pressure. The prevalence of left-handers listed in elite rankings increased from low (8.7%) to high (30.39%) time pressure sports and a distinct left-hander overrepresentation was only found in the latter (i.e. baseball, cricket and table tennis). This indicates that relative rarity and the interactive nature of a contest are not sufficient per se to evoke a left-hander advantage. Refining the fighting hypothesis is suggested to facilitate prediction and experimental verification of when and why negative frequency-dependent selection may benefit left-handedness.

    1. Background

    Frequency-dependent selection can ensure the maintenance of polymorphisms in animal and non-animal species [1,2]. In humans, the fighting hypothesis proposes that negative frequency-dependent selection may account for the stable maintenance of an unbalanced handedness polymorphism over thousands of years [3,4]. Assuming that handedness is a partly heritable trait, health-related fitness costs potentially associated with left-handedness might be outweighed by fitness benefits associated with its relative rarity, thereby ensuring reproductive success [5]. Fitness benefits are predicted for duel-like situations where unfamiliarity with left-handers may directly affect performance. Sport data support this notion in that left-handed athletes are found overrepresented in the high echelons of interactive (e.g. cricket, baseball) but not non-interactive (e.g. darts, snooker) professional sports [4,6]. The factors strengthening or attenuating the benefits of performing left-handed in interactive contests, however, have yet to be specified.

    Athletes develop task-specific perceptual–cognitive and motor skills to counter the spatio-temporal constraints induced by small distances between competitors, fast movements and/or high ball speeds in interactive sports [7]. Similar to processes of adaptation in the wild [1], predominant exposure to right-handers can result in the acquisition of skills optimal for competition against right- but not left-handers [8]. This is expressed, for example, by better anticipation of right- than left-handers' action intentions [8,9], decision-making preferences [10] or deficiency in responding to unfamiliar left-handed deliveries like in cricket batting [11].

    These issues can promote an advantage of relatively rarer left-handers [6], presumably particularly in competition under severe time pressure. If time pressure is relatively low, inaccuracy in anticipation and related motor control processes, for instance, might still be corrected successfully in time based on an online integration of sensory feedback (e.g. vision) and action [12,13]. In highly time-constrained situations, however, corrections are harder to make timely, despite the athletes’ perceptual–motor system being attuned to uncertainty under time pressure [14]. Therefore, here it was hypothesized that the occurrence and extent of a left-hander's advantage in interactive sports are linked to the contests’ underlying time pressure. Specifically, the advantage should be higher (lower) in more (less) time-constrained sports.

    The recently reported higher proportion of left-handers among the top-100 players in table tennis (a fast sport) compared with tennis, badminton and squash might be interpreted as providing some tentative initial indication for such a relationship [15]. However, that finding was based on a single retrieval date, it did not include measures of time pressure and it was restricted to four sports from men's competition. Here, to test the above hypothesis more comprehensively, the time constraints acting upon athletes in elite competition of baseball, cricket and the above four racket sports were determined and then correlated with the frequencies of left-handed players observed in the respective sport's elite rankings. The latter measure served as proxy for a left-hander’s advantage [3,11,16,17]. The sports were selected owing to similarity in composition, performance demands and game strategy. Specifically, in all the sports, a ball is hit or thrown, a bat or racket is used for interception and players try to force their opponent into not reaching the ball or making a return error. This ensured the determination of a comparable measure of time pressure across sports and precluded consideration of interactive sports that do not share the former features (e.g. boxing). Analyses included data on males and females (racket sports only; table 1 for details), but focus was on male competition as left-handedness was found particularly relevant in male–male contests [3,17].

    Table 1.Handedness in interactive ball games. Players’ task-specific handedness (i.e. throwing, or holding a racket) was obtained from the websites below or from additional searches on the web (e.g. pictures or videos of players in action found on www.youtube.com). Analysis of female competition was restricted to racket sports because no comprehensive ranking data were found for baseball and cricket for the time period investigated. N, total number of different players listed in the year-end rankings considered between 2009 and 2014; n.a., handedness data not available.

    competition sport N left right n.a. % left ranking source
    male badminton 183 23 159 1 12.64 www.bwfbadminton.org
    squash 161 14 147 8.70 www.squashinfo.com
    table tennis 182 47 135 25.82 www.ittf.com
    tennis 194 27 167 13.92 www.atpworldtour.com
    cricket (bowlers) 202 44 158 21.78 www.relianceiccrankings.com
    baseball (pitchers) 204 62 142 30.39 www.espn.go.com/mlb/stats
    female badminton 216 17 199 7.87 www.bwfbadminton.org
    squash 178 14 153 11 8.38 www.squashinfo.com
    table tennis 188 36 151 1 19.25 www.ittf.com
    tennis 196 15 181 7.65 www.wtatennis.com

    2. Material and methods

    (a) Elite rankings and left-hander frequency

    Rankings and players' task-specific handedness (i.e. hand used for throwing or for holding a racket) were retrieved from publicly available online databases (table 1). For each sport, data were collected for six seasons (2009–2014) to account for assumed temporal variation in left-hander frequencies [16,17]. For each racket sport, the top-100 players from the official year-end world rankings were retrieved. For cricket, data on the top-100 Test bowlers listed in the official ICC player rankings were considered. As no world rankings on individual players were found for baseball, ranking data for the top-78 to 94 (depending on the year) pitchers in Major League Baseball (MLB) were taken. Batsmen and hitters were not included because they do not directly impose time pressure upon their opponents. To avoid that identical players were counted multiple times, the proportion of left-handers among all different players listed in the above sports’ rankings over the 6 year period was used for analyses (table 1).

    (b) Time pressure

    Time pressure was defined by the mean time interval (in milliseconds) between the actions of two interacting players in male competition. In racket sports, the time between moments of racket-ball-contact was recorded. In baseball and cricket, the time elapsed from ball release to (estimated) bat–ball-contact was considered. Generally, shorter intervals characterize heavier time constraints. Time intervals were obtained from video analyses of professional matches in badminton (n = 1569 intervals), squash (n = 1976), table tennis (n = 778) and tennis (n = 1941), summary statistics on pitch speeds recorded in the MLB for the full season of 2008 (n = 706 374 pitches) and bowl speeds obtained from automated ball trajectory recordings run by Hawk-Eye Innovations at the Twenty20 Cricket World Cup in South Africa 2007 (n = 5652 deliveries). Accordingly, time pressure was highest in baseball and lowest in squash (electronic supplementary material S1 for details).

    3. Results

    In male competition, binomial tests (one-tailed) revealed a left-hander overrepresentation in baseball (p < 0.001), cricket (p < 0.001) and table tennis (p < 0.001), but not in tennis (p = 0.066), badminton (p = 0.185) and squash (p = 0.784) relative to a normal population estimate of 10.3% [3]. Across sports, left-hander frequency and the mean time interval between interacting players' actions were related negatively (Spearman's ρ = −0.886, p = 0.019; figure 1a), suggesting that left-handedness decreased from high (i.e. small time interval) to low time-constrained sports. The robustness of the correlation is supported by additional analysis, where a cases bootstrap (100 000 replications; see [18] for the same analytic approach) revealed a 95% confidence interval on estimates of Spearman's ρ ranging from −1 to −0.2 (figure 1b; electronic supplementary material S2 for details). When grouped, left-handedness was 2.6 times more likely among ‘high’ (baseball, cricket, table tennis) compared with ‘low’ (badminton, squash, tennis) time pressure contests (table 2). Restriction to racket sports did not alter the effect: left-handedness was more likely in table tennis than in the other racket sports combined and when compared with each sport separately. No notable differences were found between badminton, tennis and squash. Left-handedness was slightly more likely in baseball pitchers than cricket bowlers (table 2).

    Figure 1.

    Figure 1. (a) Left-handedness and time pressure in male competition. Small (large) time values indicate high (low) time pressure. White dots represent originally observed values. Small coloured symbols indicate possible locations of points as obtained from resampling across 100 000 replications. (b) Distribution of bootstrap estimates of Spearman's ρ based on 100 000 replications. The red solid and dashed lines represent the mean correlation and the limits of the 95% confidence interval (percentile method), respectively.

    Table 2.Handedness comparisons between sports.

    category comparison male competition
    female competition
    odds ratio 95% CI pa odds ratio 95% CI pa
    all sports high versus low time pressureb 2.60 (1.89, 3.58) <0.001
    racket sports high versus low time pressurec 2.57 (1.69, 3.93) <0.001 2.76 (1.72, 4.43) <0.001
    table tennis versus badminton 2.41 (1.39, 4.17) 0.001 2.79 (1.51, 5.16) <0.001
    table tennis versus tennis 2.15 (1.27, 3.64) 0.003 2.88 (1.52, 5.46) <0.001
    table tennis versus squash 3.66 (1.93, 6.94) <0.001 2.61 (1.35, 5.03) 0.002
    badminton versus tennis 0.89 (0.49, 1.63) 0.416 1.03 (0.50, 2.12) 0.542
    badminton versus squash 1.52 (0.75, 3.06) 0.159 0.93 (0.45, 1.95) 0.500
    tennis versus squash 1.70 (0.86, 3.36) 0.085 0.91 (0.42, 1.94) 0.474
    baseball and cricket pitchers versus bowlers 1.57 (1.00, 2.45) 0.031d

    aBased on Fisher's exact tests, one-tailed.

    bHigh time pressure sports include baseball, cricket and table tennis. Low time pressure sports comprise badminton, tennis and squash. The comparison in the ‘all sports’ category is calculated for male competition only, because data were not available for females in baseball and cricket.

    cTable tennis (high pressure) versus the group of relatively low-pressure racket sports (badminton, tennis, squash).

    dOutside the range for a classification as ‘statistically significant’ when multiple comparison corrections are applied to male competition data.

    In female competition, left-handers were again overrepresented in table tennis (p < 0.001), but not in badminton (p = 0.499), tennis (p = 0.548) or squash (p = 0.507) relative to a normal population estimate of 7.7% [3]. Left-handedness was 2.76 times more likely in table tennis compared with the other racket sports combined and when compared with each sport separately (table 2). No meaningful differences in left-hander frequency were found between badminton, tennis and squash. Overall, left-handedness was more prevalent in male (15.44%) than female (10.70%) racket sport competition, p = 0.004, odds ratio = 1.52, 95% CI (1.12, 2.07).

    4. Discussion

    A left-hander’s advantage does not emerge to a similar extent across structurally similar interactive sports. Left-handed male and female players were more prevalent at the elite level of highly time-constrained sports (baseball, cricket, table tennis) as opposed to sports where players have more time available between actions (e.g. tennis, squash). Only in the former type of sports was a clear left-hander overrepresentation relative to the normal population found. This suggests that left-handers' relative rarity and the interactive nature of a contest may not be sufficient to provoke a measurable fitness benefit. Rather, its occurrence appears linked to the time pressure underlying interactions, being more likely in high compared to relatively lower time-constrained contests.

    The present findings refine the fighting hypothesis as to the conditions likely required to elicit a left-handers’ performance advantage. An excess of left-handedness in high but not low time pressure sports, combined with no meaningful frequency differences between badminton, tennis and squash, suggest that left-dominance may be reliably identified only below a yet unknown time pressure threshold. In the latter contests, achievement may depend to a larger extent on basic physiological and psychological aspects (e.g. agility, concentration, emotion regulation) irrespective of an opponent's laterality (see also the discussion by Malagoli Lanzoni et al. [15]). Team-strategic considerations might additionally explain part of the excess of left-handedness in baseball and cricket. This, however, neither applies to racket sports [17] nor contradicts the hypothesis of time pressure being one, but possibly not the only, factor moderating left-dominance [6]. The relevance of time constraints and the existence of a hypothesized threshold for eliciting left-dominance need next be tested experimentally.

    Humans in close-distance fights are especially under severe time pressure and these fights' outcomes are assumed to have contributed to the maintenance of left-handedness in humans [3]. Consequently, the data at hand could provide additional tentative support for the fighting hypothesis. However, direct experimental evidence has recently been claimed to put that hypothesis to real test [18]. The present results indicate that researchers then need to ensure that the experimental task is performed under adequate time pressure. Otherwise, proper verification of negative frequency-dependent selection may fail, for example, because an initially suboptimal behavioural strategy against a left-handed opponent, resulting from adaptation to common right-handed opponents in the course of an experiment [810], could still be corrected in time without measurable performance impairment. The current findings illustrate that research should take a closer look at the interplay between specific performance demands and frequency-dependent selection in interactive contests.

    Data accessibility

    The datasets supporting this article have been uploaded as part of the electronic supplementary material.

    Competing interests

    I declare I have no competing interests.

    Funding

    This research was supported by an internal grant of the University of Kassel (Zentrale Forschungsförderung) awarded to F.L. It was also closely connected with the project ‘Laterality in sports’ funded by the German Research Foundation (DFG; HA 4361/5-2 and STR 490/11-2).

    Acknowledgements

    I thank Sophia Köhler, Carlotta Stern and Miriam Krone for helping with video analyses and Chris McManus for discussing bootstrap resampling and sharing Matlab code.

    Footnotes

    Electronic supplementary material is available online at https://dx.doi.org/10.6084/m9.figshare.c.3923299.

    Published by the Royal Society. All rights reserved.