Edith A. Widder

Bioluminescence in the Ocean: Origins of Biological, Chemical, and Ecological Diversity

In Living Color Light produced using the chemical process of bioluminescence spans the entire range of the visible spectrum. Bioluminescence has evolved independently several times in the tree of life. However, the majority of bioluminescent organisms reside in the open ocean, where their bioluminescence helps species in over 700 genera evade predators, attract mates, and find food. Widder (p. 704) reviews recent advances in understanding the evolution and distribution of bioluminescence in marine systems. From bacteria to fish, a remarkable variety of marine life depends on bioluminescence (the chemical generation of light) for finding food, attracting mates, and evading predators. Disparate biochemical systems and diverse phylogenetic distribution patterns of light-emitting organisms highlight the ecological benefits of bioluminescence, with biochemical and genetic analyses providing new insights into the mechanisms of its evolution. The origins and functions of some bioluminescent systems, however, remain obscure. Here, I review recent advances in understanding bioluminescence in the ocean and highlight future research efforts that will unite molecular details with ecological and evolutionary relationships.

Crepuscular and nocturnal illumination and its effects on color perception by the nocturnal hawkmoth Deilephila elpenor.

SUMMARY Recent studies have shown that certain nocturnal insect and vertebrate species have true color vision under nocturnal illumination. Thus, their vision is potentially affected by changes in the spectral quality of twilight and nocturnal illumination, due to the presence or absence of the moon, artificial light pollution and other factors. We investigated this in the following manner. First we measured the spectral irradiance (from 300 to 700 nm) during the day, sunset, twilight, full moon, new moon, and in the presence of high levels of light pollution. The spectra were then converted to both human-based chromaticities and to relative quantum catches for the nocturnal hawkmoth Deilephila elpenor, which has color vision. The reflectance spectra of various flowers and leaves and the red hindwings of D. elpenor were also converted to chromaticities and relative quantum catches. Finally, the achromatic and chromatic contrasts (with and without von Kries color constancy) of the flowers and hindwings against a leaf background were determined under the various lighting environments. The twilight and nocturnal illuminants were substantially different from each other, resulting in significantly different contrasts. The addition of von Kries color constancy significantly reduced the effect of changing illuminants on chromatic contrast, suggesting that, even in this light-limited environment, the ability of color vision to provide reliable signals under changing illuminants may offset the concurrent threefold decrease in sensitivity and spatial resolution. Given this, color vision may be more common in crepuscular and nocturnal species than previously considered.

MARINE BIOLUMINESCENCE SPECTRA MEASURED WITH AN OPTICAL MULTICHANNEL DETECTION SYSTEM

The emission spectra of 70 bioluminescent marine species were measured with a computer controlled optical multichannel analyzer (OMA). A 350 nm spectral window is simultaneously measured using a linear array of 700 silicon photodiodes, coupled by fiber optics to a microchannel plate image intensifier on which a polychromator generated spectrum is focused. Collection optics include a quartz fiber optic bundle which allows spectra to be measured from single photophores. Since corrections are not required for temporal variations in emissions, it was possible to acquire spectra of transient luminescent events that would be difficult or impossible to record with conventional techniques. Use of this system at sea on freshly trawled material and in the laboratory has permitted acquisition of a large collection of bioluminescence spectra of precision rarely obtained previously with such material. Among unusual spectral features revealed were organisms capable of emitting more than one color, including: Umbellula magniflora and Stachyptilum superbum (Pennatulacea), Parazoanthus lucificum (Zoantharia), and Cleidopus gloria-maris (Pisces). Evidence is presented that the narrow bandwidth of the emission spectrum for Argyropelecus affinis (Pisces) is due to filters in the photophores.

Far Red Bioluminescence from Two Deep-Sea Fishes

Spectral measurements of red bioluminescence were obtained from the deep-sea stomiatoid fishes Aristostomias scintillans (Gilbert) and Malacosteus niger (Ayres). Red luminescence from suborbital light organs extends to the near infrared, with peak emission at approximately 705 nanometers in the far red. These fishes also have postorbital light organs that emit blue luminescence with maxima between 470 and 480 nanometers. The red bioluminescence may be due to an energy transfer system and wavelength-selective filtering.

Bioluminescence in the Monterey Submarine Canyon: image analysis of video recordings from a midwater submersible

Video images of bioluminescence were recorded in situ during a 1985 study of the midwater environment of the Monterey Canyon, using a single-person, untethered submersible. Gelatinous organisms were responsible for the most brilliant bioluminescent displays, often exhibiting elaborate kinetics in response to mechanical stimulation. Images of bioluminescent displays recorded from identified organisms are shown and display patterns are described. All bioluminescence emission spectra from captured specimens were blue, with peak emissions between 460 and 494 nm. Image-analysis of recordings of mechanically stimulated bioluminescence revealed source densities between 43 and 175 m-3 and intensities between 2.5 and 37.3 μW sr-1 m-3. The predominant display type at all depths studied (between 100 and 560 m) was luminous secretions. Despite high intensities of mechanically stimulated bioluminescence, no spontaneous light production was recorded in the absence of mechanical stimulation.

Giant Deep-Sea Protist Produces Bilaterian-like Traces

One of the strongest paleontological arguments in favor of the origin of bilaterally symmetrical animals (Bilateria) prior to their obvious and explosive appearance in the fossil record in the early Cambrian, 542 million years ago, is the occurrence of trace fossils shaped like elongated sinuous grooves or furrows in the Precambrian. Being restricted to the seafloor surface, these traces are relatively rare and of limited diversity, and they do not show any evidence of the use of hard appendages. They are commonly attributed to the activity of the early nonskeletonized bilaterians or, alternatively, large cnidarians such as sea anemones or sea pens. Here we describe macroscopic groove-like traces produced by a living giant protist and show that these traces bear a remarkable resemblance to the Precambrian trace fossils, including those as old as 1.8 billion years. This is the first evidence that organisms other than multicellular animals can produce such traces, and it prompts re-evaluation of the significance of Precambrian trace fossils as evidence of the early diversification of Bilateria. Our observations also render indirect support to the highly controversial interpretation of the enigmatic Ediacaran biota of the late Precambrian as giant protists.

Bioluminescence and the Pelagic Visual Environment

Bioluminescence is a relatively rare phenomenon in the terrestrial world. Fireflies are the best-known examples, although there are also bioluminescent earthworms, beetles, millipedes, centipedes, flies, snails and fungi. None of these creatures are found in high abundance and do not appear to play a significant role in the balance of nature on land. In contrast, bioluminescent organisms in the ocean are so abundant that their significance in marine ecosystems is beyond question. However, bioluminescence is frequently overlooked or underestimated in assessments of the visual ecology of the marine environment. Although it is a generally accepted fact that light is a critical determinant of organism distribution patterns and behavior in the oceans, the impact of bioluminescence in this regard is little known. Also, while bioluminescence dominates the submarine light field at depths below the penetration of sunlight and at night in surface waters, it is poorly characterized compared to the solar light field. This article provides a general description of the pelagic visual environment with an emphasis on the current state of knowledge regarding the light field produced by bioluminescent plankton. The argument is made that, given the degree to which light has shaped life in the pelagic realm, we should expect to find comparable adaptations to the bioluminescent light field.

Comparative study of the spectral sensitivities of mesopelagic crustaceans

Abstract The spectral sensitivities of 12 species of mesopelagic crustaceans were studied by means of electrophysiological recordings. Nine of the species are vertical migrators, while 3 are not, and 9 species possess bioluminescent organs, while 3 are not bioluminescent. All species had a single peak of spectral sensitivity with maxima between 470 nm and 500 nm. There was no apparent correlation between sensitivity maxima and daytime depth distribution, migratory behavior, or the presence or absence of bioluminescent organs. With the exception of the hyperiid amphipod Phronima sedentaria, the spectral sensitivities of these mesopelagic crustaceans demonstrate a better match for maximum sensitivity to bioluminescence than to downwelling light.In a previous companion paper, we presented the details of our algorithms for performing nonlocal density functional theory (DFT) calculations in complex 2D and 3D geometries. We discussed scaling and parallelization, but did not discuss other issues of performance. In this paper, we detail the precision of our methods with respect to changes in the mesh spacing. This is a complex issue because given a Cartesian mesh, changes in mesh spacing will result in changes in surface geometry. We discuss these issue using a series of rigid solvated polymer models including square rod polymers, cylindrical polymers, and bead-chain polymers. By comparing the results of the various models, it becomes clear that surface curvature or roughness plays an important role in determining the strength of structural solvation forces between interacting solvated polymers. The results in this paper serve as benchmarks for future application of these algorithms to complex fluid systems.

UV Light in the Deep‐Sea: In Situ Measurements of Downwelling Irradiance in Relation to the Visual Threshold Sensitivity of UV‐Sensitive Crustaceans

Several species of deep‐sea crustaceans possess unusually high spectral sensitivity to UV light, measured elec‐trophysiologically. In addition, behavioral experiments indicate that these species are equally sensitive to near‐UV and blue‐green light. These results raised questions about the functional significance of this short wavelength sensitivity, since it is commonly believed that UV light is virtually absent in the deep‐sea. Using submersible based technology, we conducted measurements of downwelling irradiance at two wavelengths ‐ 380 and 480 nm. These measurements indicate that the irradiance of 380 nm light at 500–600 m, the daytime depth range of the crustaceans with a U V sensitivity peak, is high enough to be detected by these species. In addition, measurements of the spectrum of the downwelling light field conducted at dusk demonstrate that spectral changes visible at the surface are not visible at 150 m. These results are discussed with respect to hypotheses on the functional significance of ...

Propagation and Perception of Bioluminescence: Factors Affecting Counterillumination as a Cryptic Strategy

Many deep-sea species, particularly crustaceans, cephalopods, and fish, use photophores to illuminate their ventral surfaces and thus disguise their silhouettes from predators viewing them from below. This strategy has several potential limitations, two of which are examined here. First, a predator with acute vision may be able to detect the individual photophores on the ventral surface. Second, a predator may be able to detect any mismatch between the spectrum of the bioluminescence and that of the background light. The first limitation was examined by modeling the perceived images of the counterillumination of the squid Abralia veranyi and the myctophid fish Ceratoscopelus maderensis as a function of the distance and visual acuity of the viewer. The second limitation was addressed by measuring downwelling irradiance under moonlight and starlight and then modeling underwater spectra. Four water types were examined: coastal water at a depth of 5 m and oceanic water at 5, 210, and 800 m. The appearance of the counterillumination was more affected by the visual acuity of the viewer than by the clarity of the water, even at relatively large distances. Species with high visual acuity (0.11° resolution) were able to distinguish the individual photophores of some counterilluminating signals at distances of several meters, thus breaking the camouflage. Depth and the presence or absence of moonlight strongly affected the spectrum of the background light, particularly near the surface. The increased variability near the surface was partially offset by the higher contrast attenuation at shallow depths, which reduced the sighting distance of mismatches. This research has implications for the study of spatial resolution, contrast sensitivity, and color discrimination in deep-sea visual systems.