Abstract:Bombardier beetles (Brachinini) utilize a rapid series of discrete explosions inside their pygidial gland reaction chambers to produce a hot, pulsed, quinone-based defensive spray. The mechanism of brachinines' spray pulsation was explored using anatomical studies and direct observation of explosions inside living beetles using synchrotron X-ray imaging. Quantification of the dynamics of vapor inside the reaction chamber indicates that spray pulsation is controlled by specialized, contiguous cuticular structures located at the junction between the reservoir (reactant) and reaction chambers. Kinematics models suggest passive mediation of spray pulsation by mechanical feedback from the explosion causing displacement of these structures.One Sentence Summary: Spray pulsation in bombardier beetles was determined to be controlled by the displacement of specialized cuticular structures between the reservoir (reactant) and reaction chambers.
Main Text:When threatened, bombardier beetles (Fig. 1A) expel a hot spray from their pygidial glands (1, 2). The spray contains p-benzoquinones (3), chemical irritants commonly employed by arthropods (4). However, bombardier beetles are unique in utilizing an internal explosive chemical reaction to simultaneously synthesize, heat, and propel their sprays (2, 3). The spray dynamics have been investigated by high-speed photography of the spray, spray impact force measurements, recordings of explosion sounds, and simulations (5-7). Species in the tribe Brachinini (brachinines) achieve spray temperatures of ~100 °C (2) with ranges of several centimeters (1) and velocities of ~10 m/s via a "biological pulse jet" (5), where the spray consists of a rapid succession of pulses formed in discrete explosions. Pulse repetition rates of 368-735 Hz were measured from audio recordings for Stenaptinus insignis (5).It is well known that brachinines' ability to produce internal explosions is facilitated by the two-chambered construction of their pygidial glands (3) (Fig. 1B-E). Each of the beetle's two pygidial glands comprises a reservoir chamber (RSC), reaction chamber (RXC), and exit channel (EC) which vents near the abdomen tip (Fig. 1B). The distal ends of the exit channels curve dorsally to form reflector plates (Fig. 1B, RP) used for spray aiming (8). An inter-chamber valve (Fig. 1D,E, ICV) is contiguous with the walls of the reaction and reservoir chambers and separates the chambers' contents when closed (2). The pygidial glands are constructed of cuticle, a composite of chitin, proteins, and waxes (9), which protects the beetle from the toxic chemicals, high temperatures, and high pressures during explosions. The muscle-enveloped, flexible reservoir chamber (5) stores an aqueous reactant solution of ~25% hydrogen peroxide and ~10% p-hydroquinones (3), along with ~10% alkanes as a nonreactive second liquid phase (10). Valve muscles (Fig. 1D, VM) span between the valve and the reservoir chamber to facilitate valve opening. During spray emission, reactant solution flows from the reserv...