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Our formula shows that respiratory gain increases with body size and with increased rate of fluttering. Therefore, insects can regulate their rate of oxygen uptake by varying the rate of fluttering while keeping the spiracles closed during a large fraction of the time during the flutter phase. We also use a mathematical model to show that water loss is approximately proportional to the percentage of time the spiracles are open. Thus, insects can achieve both high oxygen intake and low water loss by keeping the spiracles closed most of th
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