Original researchFirst-stepping Test as a measure of motor laterality in dogs (Canis familiaris)
Introduction
Lateralization refers to the specialized functions of the right and left cerebral hemispheres and may control both motor responses and preferential processing of stimuli. It may also be used to predict more general behavioral attributes. For example, in dogs, associations between motor laterality and noise sensitivity have been reported (Branson and Rogers, 2006). Specifically, lack of paw preference (PP) in a food-retrieval task is associated with noise-related fearfulness. Fearfulness, including noise sensitivity, is one of the reasons for failure of guide dogs during training. So, knowing a given animal's laterality may be predictive of its fearfulness. Furthermore, this information may also explain the performance of a dog in guiding tasks because convention dictates that dogs work to the left of their handlers. This convention may disadvantage some dogs that might be perfectly useful if worked on the right of their handlers. Motor laterality can be task-dependent (Fagot and Vauclair, 1991, Spinozzi and Truppa, 1999, Wells, 2003, Trouillard and Blois-Heulin, 2005, Batt et al., 2008a). Various methodologies for determining motor lateralization in dogs have been reported, including reaching for food (Aydinlioğlu et al., 2000), circling direction (Branson, 2006), paw on bone/Kong™ (Branson and Rogers, 2006, Batt et al., 2007), removal of tape from eyes (Tan, 1987, Tan and Caliskan, 1987) or nose (Quaranta et al., 2004, Quaranta et al., 2006, Poyser et al., 2006, Batt et al., 2007), blanket removal, and paw-shaking (Wells, 2003). Although a variety of tests have been devised to determine PP, the way in which they chiefly reflect individual dogs' physical ability may limit their ability to reveal true brain asymmetry. In addition, some tests may be questionable on welfare grounds (e.g., the removal of adhesive tape from eyes or nose). The Kong™ (a rubber cylindrical dog toy) Test, based on foraging behavior, is the current benchmark test for determining PP in dogs (Branson, 2006, Branson and Rogers, 2006, Batt et al., 2007, Batt et al., 2008a, Batt et al., 2008b, Batt et al., 2009). However, motivation to feed varies markedly among dogs and breeds (Rooney and Bradshaw, 2004), and the test comprises 50 observations which for a single dog can take up to 4 hours to collect. Consequently, there remains a need to develop a method of determining PP without reliance on motivation to feed.
This study examined, as a potential indicator of motor bias, the first foot placed forward (first-stepping) when moving forward from a standing position. van Alphen et al. (2005) previously described a stepping-off technique, but their dogs (n = 36) commenced from a stand or sit position, and stepped off with their owners who may have biased the leg used by the dog to initiate walking. Dogs in the current study stepped-off from only a stand position, and neither the dog handler (assistant) nor researcher stepped-off with the dog when the dog was cued to step-off. This removed any potential biases caused by these operator actions. In addition, PP in the Kong™ Test was determined in the same cohort of dogs. This permitted comparisons with previous studies, as well as a comparison between the results of the two laterality tests in this study to determine whether PP alters with task. The aims of the current study were to (i) develop a methodology to assess motor laterality in dogs without the reliance on food (First-stepping Test); (ii) use this methodology to determine both strength and direction of lateralization in a cohort of potential guide dogs, and (iii) compare motor laterality findings from the First-stepping and Kong™ Tests.
Section snippets
Animals
Motor laterality was determined in potential guide dogs (n = 113) using two tasks based on food-retrieval (the Kong™ Test) and simple locomotion (the First-stepping Test). Dogs were aged between 13 and 17 months and neutered (males, n = 52; females, n = 61). They included Labrador retrievers (LR; n = 96), golden retrievers (GR; n = 9), and Labrador-golden retriever crosses (LR × GR; n = 8). Dogs were assessed for PP in both tasks at the NSW/ACT Guide Dog Training Centre, Glossodia, NSW,
Results
First-stepping Test exposed more significant PPs than the Kong™ Test (First-stepping, 76.1%; Kong™, 52.2%). A significant right population bias (P = 0.036) was observed in the First-stepping Test, whereas the Kong™ Test did not reveal a bias in the direction of laterality (DL; P = 0.30). However, when assessing the categories of PP, a clear ambidextrous bias emerged from the Kong™ data (P = 0.005) and the right-preferent bias was confirmed from the First-stepping data (P = 0.012; Figure 4). The
Discussion
Our findings indicate that lateralized behavior in the domestic dog is strongly task-dependent. There was no association between the First-stepping and Kong™ Tests for direction and strength of lateralization, or for PP category. The First-stepping Test exposed significant PPs in more dogs than the Kong™ Test (76.1% vs. 52.2%). A significant right population bias was identified in the cohort of potential guide dogs using the First-stepping Test, whereas no significant left or right population
Conclusion
This study introduces the First-stepping Test as a repeatable and consistent indicator of motor bias in the dog and has revealed a significant right population bias in a cohort of potential guide dogs undertaking this task. A comparison between measures of laterality using the Kong™ and First-stepping Tests has confirmed that lateralized behavior in the domestic dog is strongly task-dependent. The First-stepping Test was quicker, revealed stronger PPs, and exposed a significant left or right PP
Acknowledgments
The authors thank Kent Williams for assisting in the First-stepping data collection and Kate Marshall for her assistance with the interobserver ratings. The Australian Research Council and NSW/ACT Guide Dogs provided funding for this project. None of the financial contributors had any involvement in the preparation of this article. Guide Dogs NSW/ACT provided access to 113 of their trainee guide dogs for the study, but the experimental design was the product of the author's (L.T.) work.
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2021, Applied Animal Behaviour ScienceCitation Excerpt :Tomkins et al. (2012) also found that strength (although not direction) of paw use as assessed by the first stepping task was a significant predictor of outcome success on a guide dog training programme. From an animal welfare perspective, the non-invasive nature of the first stepping task is a strong positive, and from a more methodological point of view this test offers a number of advantages, being straightforward to carry out and suitable for use with all animals, regardless of their age, sex, size or breed (Tomkins et al., 2010a). The non-food oriented nature of the task also makes it suitable for subjects that are not hungry or food-motivated.