Absolute Brain Size Predicts Dog Breed Differences in Executive Function
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Absolute Brain Size Predicts Dog Breed Differences in Executive Function Daniel J. Horschler1,*, Brian Hare2,3, Josep Call4,5, Juliane Kaminski6, Ádám Miklósi7,8, & Evan L. MacLean1 Supplementary Material 1School of Anthropology, University of Arizona, Tucson, AZ, 85719, United States 2Department of Evolutionary Anthropology, Duke University, Durham, NC, 27708, United States 3Center for Cognitive Neuroscience, Duke University, Durham, NC, 27708, United States 4Department of Developmental and Comparative Psychology, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany 5School of Psychology and Neuroscience, University of St Andrews, St Andrews, United Kingdom 6Department of Psychology, University of Portsmouth, Portsmouth, United Kingdom 7Department of Ethology, Eötvös Loránd University, Budapest, Hungary 8MTA-ELTE Comparative Ethology Research Group, Budapest, Hungary *Correspondence - Email: [email protected]; Phone: 520-621-2646 Method Details Cognitive Data. Cognitive data were compiled from the Dognition.com database. Dognition.com is a citizen science website that provides dog owners with instructions for completing cognitive experiments with pet dogs in their homes. Participants receive written and video instructions for the behavioral protocols, and electronically submit data on their dog’s responses as activities are completed. We included data from ten cognitive tasks in our analyses: Yawning, Eye Contact, Arm Pointing, Foot Pointing, Cunning, Memory vs. Pointing, Memory vs. Smell, Delayed Memory, Inferential Reasoning, and Physical Reasoning. For detailed protocols of each task, see Task Details. We analyzed all subjects from a dataset originally including 18,610 dogs, with the majority of these dogs having completed multiple tasks. To ensure that our sample included only adult dogs, we removed all data from dogs who were 1 year or younger at the time of testing (n = 2,307). Because our analyses focused on breed-average performance, we eliminated data from dogs that were identified as mixed breeds, or for whom the breed identity was not known (n = 5,311; both measures determined by owner report). We also eliminated data from a small subset of poodles with reported body weights intermediate to miniature and standard morphs (n = 22, see Morphological Data). Lastly, to ensure representative samples across breeds we removed data from breeds represented by less than 20 individuals (on a task-wise basis), as well as for breeds which lacked body weight or genetic data (n = 3,573 individuals) from the primary sources these data were compiled from (see below). The final dataset we analyzed included 7,397 dogs. Demographic data for each breed included in our analyses is shown in Supplementary Table 1. Data Collection Process. All text in this section and in the following Task Details section was directly adapted from Stewart et al. (2015). All participants received instructions recommending cognitive tasks be conducted at home in a familiar room reasonably free of distractions. Participants were encouraged to take breaks whenever the dog lost motivation, and were repeatedly reminded not to intentionally influence their dog’s choices, although experimental evidence suggests that the effect of unconscious cuing on dogs during cognitive experiments may not be as ubiquitous as has often been suggested (Hauser, Comins, Pytka, Cahill, & Velez-Calderon, 2011; Pongrácz, Miklósi, Bálint, & Hegedüs, 2013; Schmidjell, Range, Huber, & Virányi, 2012). Regardless, instructions were worded to frequently remind participants that there “are no correct answers” and to let their dog choose freely. For each task, participants watched a how-to video that detailed the experimental setup, procedural protocol, and the set of possible responses from their dog that they would need to live code. Written instructions and a Frequently Asked Questions (FAQ) section were also provided, and a link to the FAQ was available at all times in case questions arose during testing. Participants were not able to advance through tasks or trials out of order, to help ensure that participants correctly followed all steps, and repeated each trial the correct number of times. Once all of the steps were complete for each trial, participants were asked to code their dog’s behavior. The majority of decisions involved either stopping a timer or indicating which of two locations the dog approached first. Participants were instructed to work with a partner unless their dog was able to sit and stay on their own. Video and written instructions emphasized the importance of staring directly ahead after releasing the dog, and of releasing the dog without nudging, pushing, or leading the dog in one direction or the other. In the instructional video, the dog was positioned directly across from the experimenter, approximately 1.8m away. Three post-it notes were placed on a line perpendicular to the dog and the experimenter. The center post-it was placed 0.6m directly in front of the experimenter, while the other two were placed 0.9m on either side of the center post-it. Once participants completed this set up, they advanced to step-by-step instructions for each trial. Task Details. The vast majority of dogs were tested using the following task order: Yawning, Eye Contact, Arm Pointing, Foot Pointing, Cunning, Memory vs. Pointing, Memory vs. Smell, Delayed Memory, Inferential Reasoning, and Physical Reasoning. Non-differential rewarding (in which any choice was rewarded) was used in Arm Pointing, Foot Pointing, and Cunning, while differential rewarding was used in Memory vs. Pointing, Memory vs. Smell, Delayed Memory, Inferential Reasoning, and Physical Reasoning. Task Details: Yawning. The Yawning task procedure was based on Joly-Mascheroni et al. (2008). Participants watched a video of instructions that included examples of dogs yawning. Participants were instructed to sit on the floor with the dog, but not to touch or interact with the dog at any point during the yawning conditions. In the control condition, the participant was prompted to say the word “yellow” every 5 seconds for 30 seconds. In the experimental condition, the participant was instructed to yawn clearly and audibly every 5 seconds for 30 seconds. Once this 30-second period ended, a 2-minute timer started. During the 30-second manipulation and the 2 minutes that followed, the participants were instructed to observe if the dog yawned. They were prompted to code whether the dog yawned after each trial (they were not asked for a frequency). There was only one trial of each condition, and all subjects received the control condition first. Task Details: Eye Contact. Procedurally, the Eye Contact task was based on Nagasawa et al. (2009). Participants first conducted a warm-up condition, in which the participant was instructed to stand facing their dog with the dog standing or sitting ~1.8 meters away. The participant was then prompted to call the dog’s name, show the dog a small treat, and place that treat directly beneath their eye and to start the 10-second timer. Once time elapsed, the participant was prompted to give the dog the treat. No coding was required and the warm-up was repeated three times. The experimental condition was the same as the warm-up except that the participant was instructed to stop the timer once the dog broke eye contact for more than two seconds. If the dog never broke eye contact, the trial ended after 90 seconds. Three trials of this condition were conducted. Task Details: Arm Pointing and Foot Pointing. The Arm Pointing task was based on Gácsi et al. (2009) while the Foot Pointing task was based on Lakatos et al. (2009). In the pointing warm- up, the dog was introduced to the two potential locations to find a treat. The participant was instructed to stand ~1.8m away from the dog, call the dog’s name, show the dog a treat, and place the treat down on the ground at arm’s length to either the left or right (~1.2m apart). The instructions indicated which side to place the treat on in each trial. The treat location was counterbalanced across trials and participants never placed a treat on the same side on two consecutive trials. Once the participant had placed the treat, they were instructed to release the dog while giving a release command. After each trial, the participant was asked to record if the dog retrieved the treat. There were six trials in this warm-up. The Arm Pointing and Foot Pointing conditions were identical to the warm-up except that two identical food treats were placed at arm’s length on either side. Participants were instructed which location they were to gesture toward and to allow the dog to retrieve both treats while recording which location the dog approached first. A first approach was illustrated in the video and was defined as crossing the plane created by the post-it notes on the left or right. In the Arm Pointing condition, participants were to point by extending their arm, hand, and index finger toward one location while standing equidistant between the two locations. In the Foot Pointing condition, participants were instructed to take one step toward one location while extending their leg and food in the direction of the other location. In both experimental conditions, gaze and gesture was to be directed at the location being indicated until the dog made their choice. Six trials were conducted in each condition. Task Details: Cunning. The Cunning (Other’s Visual Cues) task was procedurally based on Call et al. (2003). In the baseline watching condition, the dog was positioned ~1.8m from the participant. The participant was then instructed to say the dog’s name and “No” or “Leave it” twice, clearly and loudly, while placing a treat in front of them on the ground. Once the participant stood back up, they activated the countdown timer, which they only stopped when the dog approached and ate the forbidden food.