Abstract
We examined masseter recruitment and firing patterns during chewing in four adult ring-tailed lemurs (Lemur catta), using electromyography (EMG). During chewing of tougher foods, the working-side superficial masseter tends to show, on average, 1.7 times more scaled EMG activity than the balancing-side superficial masseter. The working-side deep masseter exhibits, on average, 2.4 times the scaled EMG activity of the balancing-side deep masseter. The relatively larger activity in the working-side muscles suggests that ring-tailed lemurs recruit relatively less force from their balancing-side muscles during chewing. The superficial masseter working-to-balancing-side (W/B) ratio for lemurs overlaps with W/B ratios from anthropoid primates. In contrast, the lemur W/B ratio for the deep masseter is more similar to that of greater galagos, while both are significantly larger than W/B ratios of anthropoids. Because ring-tailed lemurs have unfused and hence presumably weaker symphyses, these data are consistent with the symphyseal fusion-muscle recruitment hypothesis stating that symphyseal fusion in anthropoids provides increased strength for resisting forces created by the balancing-side jaw muscles during chewing. Among the masseter muscles of ring-tailed lemurs, the working-side deep masseter peaks first on average, followed in succession by the balancing-side deep masseter, balancing-side superficial masseter, and finally the working-side superficial masseter. Ring-tailed lemurs are similar to greater galagos in that their balancing-side deep masseter peaks well before their working-side superficial masseter. We see the opposite pattern in anthropoids, where the balancing-side deep masseter peaks, on average, after the working-side superficial masseter. This late activity of the balancing-side deep masseter in anthropoids is linked to lateral-transverse bending, or wishboning, of their mandibular symphyses. Subsequently, the stresses incurred during wishboning are hypothesized to be a proximate reason for strengthening, and hence fusion, of the anthropoid symphysis. Thus, the absence of this muscle-firing pattern in ring-tailed lemurs with their weaker, unfused symphyses provides further correlational support for the symphyseal fusion late-acting balancing-side deep masseter hypothesis linking wishboning and symphyseal strengthening in anthropoids. The early peak activity of the working-side deep masseter in ring-tailed lemurs is unlike galagos and most similar to the pattern seen in macaques and baboons. We hypothesize that this early activity of the working-side deep masseter moves the lower jaw both laterally toward the working side and vertically upward, to position it for the upcoming power stroke. From an evolutionary perspective, the differences in peak firing times for the working-side deep masseter between ring-tailed lemurs and greater galagos indicate that deep masseter firing patterns are not conserved among strepsirrhines.
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