SUMMARYIn the western Caribbean Sea, about an hour after the sun sets, a complex and ritualized light show of precise, vertically placed luminescent pulses erupts over shallow grassbeds. These are among the most complex displays known in marine systems. Displays consist of repeated trains of secreted bioluminescent pulses in a specific pattern ejected into the water column as courtship signals by male Vargula annecohenae, which are small (<2 mm) myodocopid ostracod crustaceans. Although these animals display in near darkness, we have used image intensification and infrared videography and three-dimensional analysis in the lab to demonstrate that each luminescent display train, which can be up to 60 cm long, consists of two distinct luminescent and swimming phases. The first, or 'stationary,' phase consists of three (usually) bright, longer pulses placed close together, with the male swimming in a looping pattern. We hypothesize that this pattern acts as an attention-grabbing signal for receptive females. The stationary phase is followed by the 'helical phase,' which consists of about a dozen evenly placed dimmer, shorter pulses secreted by an individual male rapidly spiraling upward in a helical pattern. We hypothesize that this phase, which has very uniform interpulse intervals and distances, helps an approaching female target and intercept the rapidly moving male. Here we provide details of these two phases, and produce a three-dimensional model of a multiply-displaying male.

In the calcium-activated photoprotein aequorin, light is produced by the oxidation of coelenterazine, the luciferin used by at least seven marine phyla. However, despite extensive research on photoproteins, there has been no evidence to indicate the origin of coelenterazine within the phylum Cnidaria. Here we report that the hydromedusa Aequorea victoria is unable to produce its own coelenterazine and is dependent on a dietary supply of this luciferin for bioluminescence. Although they contain functional apophotoproteins, medusae reared on a luciferin-free diet are unable to produce light unless provided with coelenterazine from an external source. This evidence regarding the origins of luciferin in Cnidaria has implications for the evolution of bioluminescence and for the extensive use of coelenterazine among marine organisms.

Vision is important for locomotion in complex environments. How it is used to guide stepping is not well understood. We used an eye search coil technique combined with an active marker-based head recording system to characterize the gaze patterns of cats walking over terrains of different complexity: (1) on a flat surface in the dark when no visual information was available, (2) on the flat surface in light when visual information was available but not required, (3) along the highly structured but regular and familiar surface of a horizontal ladder, a task for which visual guidance of stepping was required, and (4) along a pathway cluttered with many small stones, an irregularly structured surface that was new each day. Three cats walked in a 2.5 m corridor, and 958 passages were analyzed. Gaze activity during the time when the gaze was directed at the walking surface was subdivided into four behaviors based on speed of gaze movement along the surface: gaze shift (fast movement), gaze fixation (no movement), constant gaze (movement at the body’s speed), and slow gaze (the remainder). We found that gaze shifts and fixations dominated the cats’ gaze behavior during all locomotor tasks, jointly occupying 62–84% of the time when the gaze was directed at the surface. As visual complexity of the surface and demand on visual guidance of stepping increased, cats spent more time looking at the surface, looked closer to them, and switched between gaze behaviors more often. During both visually guided locomotor tasks, gaze behaviors predominantly followed a repeated cycle of forward gaze shift followed by fixation. We call this behavior “gaze stepping”. Each gaze shift took gaze to a site approximately 75–80 cm in front of the cat, which the cat reached in 0.7–1.2 s and 1.1–1.6 strides. Constant gaze occupied only 5–21% of the time cats spent looking at the walking surface.

Nocturnal behaviors that vary as a function of light intensity, either from the setting sun or the moon, are typically labeled as circadian or circalunar. Both of these terms refer to endogenous time-dependent behaviors. In contrast, the nightly reproductive and feeding behaviors of Vargula annecohenae, a bioluminescent ostracod (Arthropoda: Crustacea) fluctuate in response to light intensity, an exogenous factor that is not strictly time-dependent. We measured adult and juvenile activity of V. annecohenae throughout lunar cycles in January/February and June 2003. Overnight and nightly measurements of foraging and reproductive behavior of adult V. annecohenae indicated that activity was greatest when a critical "dark threshold" was reached and that the dark threshold for adult V. annecohenae is met when less than a third of the moon is visible or at the intensity of light 2-3 min before the start of nautical twilight when no moon is illuminated. Juvenile V. annecohenae were also nocturnally active but demonstrated little or no response to lunar illumination, remaining active even during brightly moonlit periods. In addition to light level, water velocity also influenced the behaviors of V. annecohenae, with fewer juveniles and adults actively foraging on nights when water velocity was high (>25 cm/s). Our data demonstrate that the strongest environmental factor influencing adult feeding and reproductive behaviors of V. annecohenae is the availability of time when illumination is below the critical dark threshold. This dependence on darkness for successful growth and reproduction allows us to classify darkness as a resource, in the same way that the term has been applied to time, space and temperature.