Predation by echolocating bats has been an important selective pressure on insects, favoring the evolution of ultrasound-sensitive ears (Hoy, 1992;Jones and Rydell, 2003). Aerially hawking bats emit ultrasonic probes and detect flying insect prey by the echoes that return from their bodies. In some hearing species flight is restricted to one gender, and sensitivity to ultrasound is reduced in the non-flying gender, e.g. moths (Cardone and Fullard, 1988) and mantises (Yager, 1990). These observations suggest that sensitivity to ultrasound is closely linked to flight physiologically, as well as evolutionarily.Many cricket species are flight dimorphic, occurring both as long-winged and short-winged forms (Zera, 2004;Roff and Fairbairn, 2007). The long-winged form is initially capable of flight, but individuals may lose this ability when flight muscles undergo histolysis, which becomes increasingly probable with advancing adult age (Shiga et al., 1991). The short-winged form is incapable of flight throughout life. Unlike the cases cited above, in flight-dimorphic crickets the risk of bat predation varies within a single gender and, for long-winged crickets, within a single individual. Moreover, crickets use hearing not only to detect bats but also, in a lower range of frequencies, for intraspecific communication (Moiseff et al., 1978). We compare auditory sensitivity between flightcapable and flight-incapable forms of the cricket Gryllus texensis, both to ultrasonic stimuli and to the sonic frequency of intraspecific signals. We find, using both behavioral and neurophysiological measures, that sensitivity to ultrasound, but not to low sound frequencies, varies according to the ability to fly. Our findings suggest that ultrasound sensitivity is one of a suite of developmental and physiological phenotypes that are linked to flight, and thus to the risk of predation by aerially hawking bats.
Materials and methodsAnimals Gryllus texensis (Cade and Otte) were reared in the laboratory at 25-28°, on a 12·h:12·h photoperiod, with ad libidum access to Purina TM Cat Chow and water. Under these conditions, approximately half the males develop as long-winged, and half as short-winged individuals. The frequency of short-winged females is much lower, approximately 10%. We studied males aged 7-21 days after the final molt. Both long-and short-winged males were selected from the same cohorts, the adult ages of which were known to within 1 week. Thus, both wing morphs were represented approximately equally over the entire range of ages. Experiments were performed during the animals' scotophase.
Sound stimuliStimuli were produced by National Instruments (Austin, TX, USA) digital-to-analog boards (AT-MIO 16E4, 12-bit resolution or PCI-6251, 16-bit) with sampling rate of 100·kHz. Sound level (r.m.s. of a constant tone with the same peak amplitude as actual stimuli) was adjusted by a programmable attenuator (PA4, Tucker-Davis, Alachua, FL, USA or 50P-076, JFW Industries, Indianapolis, IN, USA) and calibrated with Bruël and Kjaer instrum...