Mechanisms of fluid production in smooth adhesive pads of insects.

Insect adhesion is mediated by thin fluid films secreted into the contact zone. As the amount of fluid affects adhesive forces, a control of secretion appears probable. Here, we quantify for the first time the rate of fluid secretion in adhesive pads of cockroaches and stick insects. The volume of footprints deposited during consecutive press-downs decreased exponentially and approached a non-zero steady state, demonstrating the presence of a storage volume. We estimated its size and the influx rate into it from a simple compartmental model. Influx was independent of step frequency. Fluid-depleted pads recovered maximal footprint volumes within 15 min. Pads in stationary contact accumulated fluid along the perimeter of the contact zone. The initial fluid build-up slowed down, suggesting that flow is driven by negative Laplace pressure. Freely climbing stick insects left hardly any traceable footprints, suggesting that they save secretion by minimizing contact area or by recovering fluid during detachment. However, even the highest fluid production rates observed incur only small biosynthesis costs, representing less than 1 per cent of the resting metabolic rate. Our results show that fluid secretion in insect wet adhesive systems relies on simple physical principles, allowing for passive control of fluid volume within the contact zone.