The activity of jaw and hyolingual muscles during the entire feeding sequence is examined in the lizard Agama stellio, with special focus on the intraoral transport and swallowing stages. Correlation of electromyography (EMG) data with kinematics shows that the kinematic phases (slow opening, SO; fast opening, FO; fast closing, FC; slow closing/power stroke, SC/PS) are characterised by distinct activities in the jaw and hyolingual muscles. The SO phase is clearly the result of tongue protraction (upon protraction, the tongue is pulled against the prey and consequently the lower jaw is pushed down), whereas the FO phase is caused by activity in the jaw opener and dorsal cervical muscles. Both the FC and SC/PS phases are characterised by pronounced activity in the jaw adductor muscles. Tongue retraction is produced by activity in the hyoid and tongue retractor muscles. A quantitative analysis of time-related EMG data shows that, in accordance with the kinematic analyses, three different stages can be recognised as components of the feeding cycle: prey capture, intraoral transport and swallowing. However, analysis of intensity-related data allowed a fourth stage, crushing, to be detected. Whereas there are indications that prey capture, intraoral transport and swallowing are controlled by different motor patterns, the differences between crushing and transport are likely to be caused by feedback mechanisms. Our results show the importance of including intensity-related data in quantitative analyses of EMG recordings in order to discriminate between feeding stages. Additionally, it is shown that both the jaw and the hyolingual muscles play crucial roles during feeding. During all stages, movements of the hyolingual apparatus are an essential part of the feeding cycle. Thus, when examining lizard feeding mechanisms, the activity patterns of the hyolingual muscles should not be neglected.
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