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The Distal Hindlimb Musculature of the Cat: Multiaxis Moment Arms at the Ankle Joint

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The Distal Hindlimb Musculature of the Cat: Multiaxis Moment Arms at the Ankle Joint Exp Brain Res (1993) 96:141-151 Experimental Brain Research Springer-Verlag 1993 The distal hindlimb musculature of the cat: multiaxis moment arms at the ankle joint R.P. Young, S.H. Scott, G.E. Loeb MRC Group in Sensory-Motor Physiology and Bio-Medical Engineering Unit, Queen's University, Kingston, Ontario, Canada K7L 3N6 Received: 21 August 1992 / Accepted: 6 May 1993 Abstract. The cat hindlimb muscles have been classified, that are spanned by the muscle. The torque produced by traditionally, as flexors and extensors, based on their ac- a muscle is the product of its tension (active and passive) tions in the parasagittal plane and their patterns of re- and its moment arm at the joint in question (see Fig. 1). cruitment during locomotion and reflex responses. This The moment arm is given by the distance from the joint study provides a detailed examination of the relative center to the line of action of the muscle. This vector can magnitudes of the various moment arms of the cat ankle be decomposed into components that correspond to the muscles and the interdependent effects of position in the anatomical degrees of freedom. It is also the case that the various axes of motion. We used a method based on ob- amount of tension that can be generated by a muscle serving small sliding movements of tendon in response to depends on the length and velocity of its sarcomeres, small angular displacements of the joint. Surprisingly, we which in turn depends on the position and motion of the found that the ankle joint of the cat permits substantial joints that it crosses, acting through the moment arms at motion in three axes (eversion/inversion and abduction/ those joints. Thus the nature and amount of work that a adduction as well as extension/flexion) and many muscles given muscle can do on the skeleton depends at lea.st as crossing the ankle joint have their largest moment arms much on the exact course of its tendons around each joint about axes other than extension/flexion. These moment as it does on the cross-sectional area of the muscle. arms often depended on the joint position in the axis of The feline ankle is a typical joint, in that it has numer- the moment arm and, to a lesser degree, on the extension/ ous muscles which act upon multiple degrees of freedom. flexion angle as well. For some muscles (notably peroneus The muscles crossing this joint traditionally have been longus) there was sufficient variability that the predomi- considered to act in the main axis of motion during loco- nant action in neutral posture (axis with the largest mo- motion, plantar-flexion/dorsi-flexion (here designated ment arm) could change from animal to animal, which physiologically as extension or flexion, respectively; may be related to heterogeneities of locomotor and reflex Abraham and Loeb 1985; Crouch 1969). Secondary con- recruitment reported in the companion paper. sideration is usually given to eversion/inversion (rotation along the long axis of the foot). In a preliminary report Key words: Muscles - Ankle - Moment arm - Cat (Young et al. 1992), we have described a surprisingly large range of motion and large muscle moment arms in ab- duction/adduction (rotation of the foot with respect to the long axis of the shank). By comparison, abduction/ Introduction adduction of the human foot does not occur in the ankle joint itself, but in the talonavicular joint of the foot When the nervous system commands a movement, either (where it is coupled to eversion/inversion) and in rotation through reflex or voluntary control, the results depend of the tibia (about its long axis) at the knee (Levens et al. on the intrinsic mechanical properties of the muscu- 1948). In this report, we describe in detail the relative loskeletal system. Therefore, a full understanding of the moment arms in each of these three axes for all of the neural mechanisms underlying the control of any motor muscles that cross the ankle joint. In particular, we have behavior depends upon a thorough understanding of examined the changes in moment arms that occur as the musculoskeletal structure and function. joint is moved over the range of motion in the axis of the The contractile force generated by a muscle produces moment arm and simultaneously, in an orthogonal axis. skeletal motion by producing torques about the joints We have also described surprisingly large variability be- Correspondence to: G.E. Loeb, Abrarnsky Hall, Queen's University, tween specimens in the relative actions of some of these Kingston, Ontario, Canada K7L 3N6 muscles, which may relate to variability in their physio- 142 Fig. l. The relationship between moment arm (MA), change in joint angle (A~p), and change in muscle length (Ax). The position of a point on the muscle relative to the tibia is recorded at a joint angle ~ and at an an- .... t-t ....L-t gle q~ + A~p. The moment arm of the muscle equals the change in muscle length divided by the change in joint angle (in radians). The amount of torque ('~) generated by a muscle spanning the ankle joint is the product of the muscle's force (F~) and its moment arm about the ankle joint. The force transmitted to the end of the foot (Fp) equals the muscle force (Fro) multiplied by its moment arm about the ankle joint and divided by the length of the foot (L) Fig. 2. Anatomy of the feline ankle mus- culature. Left, lateral view of the leg and ankle; middle, cross-section of the ankle at the level of the malleoli; right, medial view of the leg and ankle. Muscle abbre- viations: PL, peroneus longus, PB, per- oneus brevis, PT, peroneus tertius, TP, tibialis posterior, FDL, flexor digitorum longus, FHL, flexor hallucis longus, TA, tibialis anterior, El)L, extensor digitorum LATERAL CROSS-SECTION MEDIAL longus, and SO, soleus logical recruitment described in the companion paper fore, moment arms were computed from the tendon dis- (Loeb 1993). placements measured during small increments of motion Figure 2 shows gross anatomical views of the course of applied to each anatomical axis (An et al. 1983, 1984; the muscles studied and their tendons about the ankle. A Spoor and Van Leeuwen 1992; Spoor et al. 1990; Young cross section was taken through the medial and lateral et al. 1992). This technique makes no assumptions re- malleoli, the anatomical landmarks that denote the ap- garding the actual location, unicentricity, or even con- proximate center of rotation in flexion/extension. It centricity of the centers of rotation for each degree of shows that many of these muscles lie very close to this freedom. presumed center of rotation, so that any attempt to esti- mate moment arm from direct anatomical measurements would be very sensitive to small errors in locating the Materials and methods exact center of rotation, which may not even be fixed over the anatomical range of motion. Moment arms in the The moment-arm data reported here are from 19 cat preparations (adult animals of either sex). An additional six animals had been other axes depend on the often oblique course of these used in preliminary studies to refine the experimental technique and tendons through the retinacula and over the boney pul- one cadaver of similar mass (described below) was used to survey leys of the distal tibia, fibula, and tarsal bones, which the musculotendinous architecture of these muscles. The animals cannot be appreciated in two-dimensional views. There- were deeply anesthetized with intraperitoneal pentobarbital sodium 143 Table 1. Muscles and their path length changes Muscle Abbreviation Mass Fasc a AEx/F1 AEv/Iv AAb/Ad ALPsum FS (g) (mm) (mm) (mm) (mm) (mm) Peroneus longus PL 1.81 12.8 4.8 1.6 3.9 10.3 0.80 Peroneus brevis PB 1.05 8.8 0.3 1.8 4.1 6.2 0.70 Peroneus tertius PT 0.47 12.0 0.7 2.0 3.0 5.7 0.48 Tibialis posterior TP 1.16 5.2 0.6 0.3 4.3 5.2 1.00 Flexor digitorum FDL 1.56 17.0 3.0 1.4 3.6 8.0 0.47 longus Flexor hallucis longus FHL 5.10 15.6 8.3 0.6 1.4 10.3 0.66 Tibialis anterior TA 7.41 49.1 15.8 2.6 1.3 19.7 0.40 Extensor digitorum EDL 3.65 25.3 19.0 1.7 2.0 22.7 0.91 longus Soleus SO 2.93 a 33.5 22.0 b 22.0 b 0.66 Medial gastrocnemius MG 10.58 a 19.7 22.0 b 22.0 b 1.12 Lateral gastrocnemius LG 11.65 a 18.4 22.0 b 22.0 b 1.19 FS, fascicle stroke (ALPsum/Fasc) where ALPsum is the sum of b Moment arm was measured only for soleus and only about the length changes possible in each of the three anatomical axes: Ex/F1, Ex/F1 axis; we assumed that there is no significant moment arm extension/flexion; Ev/Iv, eversion/inversion; Ab/Ad, abduction/ad- about the other axes and that the other muscles contributing to the duction; Fasc, fascicle length; common Achilles tendon have a similar range of motion a Obtained from 2.8-kg cat; sarcomere length normalized to 2.3 gm and, as required, further doses of pentobarbital sodium were admin- mur fixing the knee at an angle of 90-100 ~ The transmitting istered intravenously. At completion of the experiment, the animals piezoelectric crystal was positioned along the line of pull of the were killed with an overdose of anesthetic.
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