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Stiftung Tierärztliche Hochschule Hannover Die ontogenetische ...

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Studie II: Shift of the CoM in growing dogs<br />

contrast, the stance time ratio between fore- and hindlimbs increased during the<br />

course of this study despite an increase in relative velocity. Relative fore- and<br />

hindlimb duty factors have been suggested to reflect the antero-posterior mass<br />

distribution of trotting quadrupeds and adding mass to the forelimb led to an increase<br />

in the fore- to hindlimb stance time ratio (Lee et al., 2004). Therefore, the increase in<br />

the stance time ratio observed herein corroborates our conclusion that the CoM shifts<br />

cranially when dogs grow.<br />

Ontogenetic changes in limb loading and the position of the CoM<br />

Between PW11 and PW13 (three dogs) or PW19 (three dogs), the time to peak<br />

vertical force increased significantly in the hind- but not the forelimbs. Therefore,<br />

loading rate was greater in the hindlimbs when the dogs were very young. Various<br />

factors influence the shape of the GRF curve such as the geometric compression of<br />

the leg spring due to the forward motion of the body as well as the limb’s angle of<br />

attack, stiffness and anatomical design (Farley et al., 1993; Witte et al., 2004).<br />

Spring-mass-model simulations, for example, have shown that the trajectory of the<br />

CoM during a stride depends, among other factors, on the angle of attack (Farley et<br />

al., 1993; Seyfarth et al., 2002). That is, steeper angles are associated with a<br />

relatively earlier minimum in the trajectory of the CoM and accordingly an earlier<br />

peak of the vertical force. Compared with when older, puppies appear to protract<br />

their limbs less and hit the ground in a more vertical paw trajectory and a more<br />

flatfooted manner; thus, they seem to have a lower angle of limb retraction before<br />

touch down and therefore a greater rate of limb compression. One consequence of a<br />

steeper angle of touch down is that limb muscle force must be build up more rapidly<br />

(Seyfarth et al., 2002). However, without the according kinematic and electromyographic<br />

data, it remains open whether the increased loading rate in the hindlimbs of<br />

puppies is the consequence of differences in limb behavior.<br />

Numerous studies have shown that the whole-body CoM is situated relatively<br />

cranial in dogs (Bryant et al., 1987; Budsberg et al., 1987; Rumph et al., 1994;<br />

DeCamp, 1997; Lee et al., 1999; ; Bertram et al., 2000; McLaughlin, 2001; Fanchon<br />

et al., 2006; Bockstahler et al., 2007; Walter and Carrier, 2007; Katic et al., 2009;<br />

59

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