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Several studies have assessed different components of lateralization. To date, the relationships between the 3 measures of laterality, structural, motor, and sensory, have not widely been assessed. Specifically, the relationships between structural (hair whorl characteristics) and motor or sensory lateralization have largely been overlooked. This study investigated the associations between these measures of lateralization in dogs (n = 114), using hair whorl characteristics (structural), the Kong and First-stepping Tests (motor), and the Sensory Jump Test (sensory). Several associations emerged, revealing the first evidence of a relationship between structural asymmetry (both the presence and direction of a hair whorl in various regions of the body) and sensory lateralization. Specifically, the presence of a whorl on the dog's left side of the head (cephalic) and thorax was associated with a right visual bias. In addition, right visual bias was probable if the ventral mandibular whorl was present in a counter-clockwise direction ( P = 0.008). Our data also demonstrated an association between structural and motor laterality (paw preference). Most notably, dogs with clockwise chest whorls were significantly more right-preferent in the First-stepping Test than those with counter-clockwise whorls ( P = 0.010). In addition, an association between measures of motor and sensory lateralization also emerged, representing some of the first evidence of such a relationship in nonhuman animals.

Motor laterality in potential guide dogs (n = 113) was determined using two methods: the Kong™ Test (the current benchmark test) and our innovation, a First-stepping Test. Kong™ studies record paw use during foraging from a rubber cylinder. However, motivation to feed varies markedly among dogs and breeds, and the test comprises 50 observations which for a single dog can take up to 4 hours to collect. The First-stepping Test bypasses hunger, and simply records the first foot advanced after standing with both forelegs level. In contrast to the Kong™ Test, it allows 50 observations to be gathered in 20 minutes. The findings of the current study indicate that lateralized behavior in the domestic dog is strongly task-dependent. The First-stepping Test revealed more significant paw preferences (PPs) than the Kong™ Test (76.1% vs. 52.2%). A significant right population bias ( P = 0.036) was revealed using this novel test, whereas there was no bias in the direction of laterality observed using the Kong™ Test ( P = 0.30). Instead, there was a significant population bias for the category of ambidextrous PP using the Kong™ ( P = 0.005). Strength of laterality (regardless of direction) was significantly high in the First-stepping Test ( P 0.001), but only weak PPs were observed for the Kong™ Test ( P = 0.65). Measures of laterality from the First-stepping Test were not influenced by dog factors, whereas age, breed, and sex of the dog were significantly associated with several laterality measures during the Kong™ Test. The occurrence of using both paws simultaneously to either hold the Kong™, or to step-off by means of jumping, was associated with a decrease in lateral strength for both tests. The First-stepping Test was repeatable, and the intraobserver (κ = 0.988) and interobserver (κ = 0.975) reliabilities were high. It offers a simple, quick and, when compared with the Kong™ Test, a more discriminative method of determining PP that is less influenced by dog factors.

Hair whorl characteristics were assessed in the domestic dog (Canis familiaris) in the regions of cephalic, cervical (dorsal, ventral, and lateral), thoracic and brachial axillary regions, the chest, shoulders, elbows, ventral abdominal region, and on the caudal thighs (ischiatic). They were classified as simple or tufted, and their position was recorded as the distance between their centers and bony landmarks within each region. The distribution of whorls was explored in a cohort of domestic dogs (N = 120) comprising a variety of breeds and cross-breeds, sourced from shelters (N = 60) and the general public (N = 60). Whorls observed in the majority of dogs in this cohort typically occurred on the chest, brachial axillary region, elbows, and ischiatic region. Atypical whorls were present in fewer than 20% of the population, and included those on the head (cephalic), cervical regions (dorsal, ventral, and lateral), shoulders, thoracic axillary region, and on the ventral abdominal region. The majority of whorls on dogs were classified as simple. In contrast, those located on the elbows and the majority of chest whorls were tufted. The presence and position of whorls were often associated with several variables including coat length and thickness, and the sex and source of the dog. The palpation and hair-cluster method of whorl assessment described in this article is best suited to dogs with short-to-medium coat lengths. The current methodology developed to assess hair whorl characteristics provides a framework for future investigations into any associations between hair whorl characteristics and other canine traits such as temperament.

In horses and cattle, hair whorls have been shown to act as a structural marker of reactivity and behavioral lateralization. Few studies on canine whorls have been reported and none have assessed whorl position or direction of flow. This study describes the distribution and characteristics of whorl in each of 10 regions in which whorls are typically located in dogs. Hair whorls were assessed in dogs (n = 120) and were recorded as clockwise or counterclockwise in the cephalic, cervical (dorsal, lateral, ventral), thoracic and brachial axillary, chest, shoulders, elbows, abdominal, and ischiatic regions. Bilateral whorls, including brachial axillary, elbow, abdominal and ischiatic whorls, rotated in opposing directions, allowing the dog's overall hair coat to be symmetrical. Cephalic, brachial axillary, and ischiatic whorls were consistent in their direction; cephalic and ischiatic whorls were clockwise on the right side of the body, and counterclockwise on the left, whereas right brachial axillary whorls were counterclockwise and left were clockwise. The central chest whorl was predominantly counterclockwise (91.21%). Direction of whorls was associated with several factors, including coat length, coat thickness, sex and source of the dog.