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Craniofacial Muscles

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330 L.K. McLoon and F.H. Andrade

17.6 Aging

Aging also affects the craniofacial muscles differently than limb skeletal muscles.

Again, while some alterations from normal have been described, relative to the

signi fi cant atrophy that can be seen in aging skeletal muscle, the craniofacial muscles

remain relatively normal. We will give some examples (Table 17.3 ), but this is

not meant to be a complete summary of the work in this fi eld.

As might be expected from the differential susceptibility to disease, aging affects

each craniofacial muscle group differentially. Extraocular muscles again show the

fewest age-related changes both morphologically and functionally (Yang and

Kapoula 2008 ; Valez et al. 2012 ) , although some changes in connective tissue density

and an increase in mitochondrial defects occur (McMullen et al. 2009 ) . Masseter

also shows constancy in muscle structure and function during aging (Norton et al.

2001 ) . Laryngeal muscles display fewer age-related changes compared to limb

muscle, but still show elevation in connective tissue, some increased variation in

myo fi ber cross-sectional area, changes in neuromuscular junction structure (Connor

et al. 2002 ), plus less force and slower shortening velocity (Kersing and Jennekens

2004 ; McMullen and Andrade 2006, 2009 ) . In facial muscles, while the skin undergoes

signi fi cant loss of elastin, the facial muscles themselves are normal (Lee et al.

2011 ) . Aging changes in tongue muscles are quite interesting; aging retrusive tongue

muscles do not show a decrement in overall contractile speed or force production,

but signi fi cant changes in forces is seen in aging protrusive tongue muscles (Nagai

et al. 2008 ; Connor et al. 2009 ) . The basis for this difference in unclear. However,

all of the changes in craniofacial muscles are relatively minor compared to those

seen in aging limb skeletal muscle where signi fi cant atrophy, fat conversion, fi brosis,

and denervation changes can occur.

17.7 Conclusions and Future Studies

It is clear from the collective chapters in this book that one cannot draw conclusions

about structure and function of craniofacial muscles from the study of limb skeletal

muscles. Their behavior cannot be predicted by studies of limb skeletal muscle;

often they change in diametrically opposed ways in the presence of myogenic signaling

factors (Tzahor et al. 2003 ) or during aging (Monemi et al. 1999 ) . The craniofacial

muscles appear to represent one end of the continuum of skeletal muscle

types in the body, with the relatively more homogeneous soleus at one end and

extraocular and laryngeal muscles at the other. Except for tongue muscles, from

their onset craniofacial muscles are derived from non-segmented, non-somitic

cranial mesoderm, with different genes controlling their formation compared to

those that direct the formation of somitic muscle. The mature muscles each display

a relatively unique set of contractile and metabolic proteins, which are mirrored in

the collective physiological and contractile differences seen in the craniofacial

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