title: Forelimb Kinematics of Rats Using XROMM, with Implications for Small Eutherians and Their Fossil Relatives.
abstract: The earliest eutherian mammals were small-bodied locomotor generalists with a forelimb morphology that strongly resembles that of extant rats. Understanding the kinematics of the humerus, radius, and ulna of extant rats can inform and constrain hypotheses concerning typical posture and mobility in early eutherian forelimbs. The locomotion of Rattus norvegicus has been extensively studied, but the three-dimensional kinematics of the bones themselves remains under-explored. Here, for the first time, we use markerless XROMM (Scientific Rotoscoping) to explore the three-dimensional long bone movements in Rattus norvegicus during a normal, symmetrical gait (walking). Our data show a basic kinematic profile that agrees with previous studies on rats and other small therians: rats maintain a crouched forelimb posture throughout the step cycle, and the ulna is confined to flexion/extension in a parasagittal plane. However, our three-dimensional data illuminate long-axis rotation (LAR) movements for both the humerus and the radius for the first time. Medial LAR of the humerus throughout stance maintains an adducted elbow with a caudally-facing olecranon process, which in turn maintains a cranially-directed manus orientation (pronation). The radius also shows significant LAR correlated with manus pronation and supination. Moreover, we report that elbow flexion and manus orientation are correlated in R. norvegicus: as the elbow angle becomes more acute, manus supination increases. Our data also suggest that manus pronation and orientation in R. norvegicus rely on a divided system of labor between the ulna and radius. Given that the radius follows the flexion and extension trajectory of the ulna, it must rotate at the elbow (on the capitulum) so that during the stance phase its distal end lies medial to ulna, ensuring that the manus remains pronated while the forelimb is supporting the body. We suggest that forelimb posture and kinematics in Juramaia, Eomaia, and other basal eutherians were grossly similar to those of rats, and that humerus and radius LAR may have always played a significant role in forelimb and manus posture in small eutherian mammals.
authors.affiliation: Biology Program, Stockton University, Galloway, New Jersey, 08205, United States of America.
authors.forename: Matthew F
authors.initials: MF
authors.lastname: Bonnan
authors.affiliation: Physics Program, Stockton University, Galloway, New Jersey, 08205, United States of America.
authors.forename: Jason
authors.initials: J
authors.lastname: Shulman
authors.affiliation: Biology Program, Stockton University, Galloway, New Jersey, 08205, United States of America.
authors.forename: Radha
authors.initials: R
authors.lastname: Varadharajan
authors.affiliation: Physics Program, Stockton University, Galloway, New Jersey, 08205, United States of America.
authors.forename: Corey
authors.initials: C
authors.lastname: Gilbert
authors.affiliation: Biology Program, Stockton University, Galloway, New Jersey, 08205, United States of America.
authors.forename: Mary
authors.initials: M
authors.lastname: Wilkes
authors.affiliation: Department of Biology, California State University, San Bernardino, California, 92407, United States of America.
authors.forename: Angela
authors.initials: A
authors.lastname: Horner
authors.affiliation: Department of Ecology and Evolutionary Biology, Brown University, Providence, Rhode Island, 02912, United States of America.
authors.forename: Elizabeth
authors.initials: E
authors.lastname: Brainerd
mesh_terms: D000818:Animals; D001696:Biomechanical Phenomena; D004551:Elbow Joint / Q000033:anatomy & histology / Q000502:physiology; D005552:Forelimb / Q000033:anatomy & histology* / Q000502:physiology*; D005580:Fossils / Q000033:anatomy & histology*; D006811:Humerus / Q000033:anatomy & histology / Q000502:physiology; D008297:Male; D011187:Posture; D011884:Radius / Q000033:anatomy & histology / Q000502:physiology; D016059:Range of Motion, Articular; D051381:Rats / Q000033:anatomy & histology* / Q000502:physiology*; D017207:Rats, Sprague-Dawley; D012785:Shoulder Joint / Q000033:anatomy & histology / Q000502:physiology; D014457:Ulna / Q000033:anatomy & histology / Q000502:physiology; D016138:Walking