Unexpected diversity of bioluminescence in planktonic worms

The vast majority of pelagic bioluminescent organisms emit a blue light with emission maxima (λ_max) ranging from 450 to 490 nm. Among the known outliers, the tomopterids (Annelida: Polychaeta) are usually described as yellow‐emitters (λ_max = 565–570 nm) for which bioluminescence functions as a specific recognition signal. Here, we report the first data regarding the colours emitted by four different tomopterid species, Tomopteris pacifica, T. carpenteri, T. septentrionalis and T. planktonis. Surprisingly, T. planktonis is a blue‐emitter (λ_max = 450 nm). Our pharmacological results on T. planktonis support cholinergic control, as recently demonstrated in the yellow‐emitter, T. helgolandica. Moreover, as revealed by epifluorescence microscopy, the light seems to be produced in both species from the same yellow‐pigmented parapodial glands. Despite these similarities, tomopterids express an unexpected diversity of bioluminescent colour patterns. This leads us to reassess the ecological value of bioluminescence within this group.     

Blunt egg pole holds cues for alien egg discrimination: experimental evidence

Eggshell colour patterns play a crucial role in avian host–parasite coevolution. In contrast to many experiments investigating general host egg discrimination abilities, studies testing where specific recognition cues are located on the eggshells (on blunt, sharp or both egg poles) are lacking. Previous studies suggested that discrimination cues might be located at the blunt egg pole, where the shell patterning is typically concentrated. We tested this hypothesis experimentally in species subject to interspecific (great reed warblers Acrocephalus arundinaceus, reed warblers A. scirpaceus), and also intraspecific parasitism (song thrushes Turdus philomelos, blackbirds T. merula). We examined host responses towards two types of intraspecific eggs painted non‐mimetic immaculate blue either at blunt or sharp poles. All four species rejected eggs manipulated at the blunt pole at significantly higher rates, indicating that they perceive the critical recognition cues in the same egg part. Thus, the presence of egg recognition cues at the blunt egg pole may be a general phenomenon in birds parasitized by both intraspecific and interspecific parasites.     

Tuning of photoreceptor spectral sensitivity in fireflies (Coleoptera: Lampyridae)

Sexual communication between male and female fireflies involves the visual detection of species-specific bioluminescent signals. Firefly species vary spectrally in both their emitted light and in the sensitivity of the eye, depending on the time when each is active. Tuning of spectral sensitivity in three firefly species that occupy different photic niches was investigated using light and electron microscopy, microspectrophotometry, and intracellular recording to characterize the location and spectral absorption of the screening pigments that filter incoming light, the visual pigments that receive this filtered light, and the visual spectral sensitivity. Twilight-active species had similar pink screening pigments, but the visual pigment of Photinus pyralis peaked near 545 nm, while that of P. scintillans had a λ_max near 557 nm. The night-active Photuris versicolor had a yellow screening pigment that was uniquely localized, while its visual pigment was similar to that of P. pyralis. These results show that both screening and visual pigments vary among species. Modeling of spectral tuning indicates that the combination of screening and visual pigments found in the retina of each species provides the best possible match of sensitivity to bioluminescent emission. This combination also produced model sensitivity spectra that closely resemble sensitivities measured either with electroretinographic or intracellular techniques. Vision in both species of Photinus appears to be evolutionarily tuned for maximum discrimination of conspecific signals from spectrally broader backgrounds. Ph. versicolor, on the other hand, appears to have a visual system that offers a compromise between maximum sensitivity to, and maximum discrimination of, their signals. Accepted: 29 September 1999     

ACCLIMATION OF THE PHOTOSYNTHETIC APPARATUS OF PALMARIA PALMATA (RHODOPHYTA) TO LIGHT QUALITIES THAT PREFERENTIALLY EXCITE PHOTOSYSTEM I OR PHOTOSYSTEM II1

Acclimation of the photosynthetic apparatus to light absorbed primarily by phycobilisomes (which transfer energy predominantly to photosystem II) or absorbed by chlorophyll a (mainly present in the antenna of photosystem I) was studied in the macroalga Palmaria palmata L. In addition, the influence of blue and yellow light, exciting chlorophyll a and phycobilisomes, respectively, ivas investigated. All results were compared to a white light control. Complementary chromatic adaptation in terms of an enhanced ratio of phycoerythrin to phycocyanin under green light conditions was observed. Red light (mainly absorbed by chlorophyll a) and green light (mainly absorbed by phycobilisomes) caused an increase of the antenna system, which was not preferentially excited. Yellow and blue light led to intermediate states comparable to each other and white light. Growth was reduced under all light qualities in comparison to white light, especially under conditions preferably exciting phycobilisomes (green light-adapted algae had a 58% lower growth rate compared to white light-adapted algae). Red and blue light-adapted algae showed maximal photosynthetic capacity with white light excitation and significantly lower values with green light excitation. In contrast, green and yellow light-adapted algae exhibited comparable photosynthetic capacities at all excitation wavelengths. Low-temperature fluorescence emission analysis showed an increase of photosystem II emission in red light-adapted algae and a decrease in green light-adapted algae. A small increase of photosystem I emission teas also found in green light-adapted algae, but this was much less than the photosystem II emission increase observed in red light-adapted algae (both compared to phycobilisome emission). Efficiency of energy transfer from phycobilisomes to photosystem II was higher in red than in green light-adapted algae. The opposite was found for the energy transfer efficiency from phycobilisomes to photosystem I. Zeaxanthin content increased in green and blue light-adapted algae compared to red, white, and yellow light-adapted algae. Results are discussed in comparison to published data on unicellular red algae and cyanobacteria.     

Intraspecific variation in retinal cone distribution in the bluefin killifish,<Emphasis Type="Italic"> Lucania goodei</Emphasis>

Studies of visual ecology have typically focused on differences among species while paying less attention to variation among populations and/or individuals. Here, we show that the relative abundance of UV, violet, yellow, and red cones varies between two populations of bluefin killifish, Lucania goodei. Animals from a spring population (high-transmission UV/blue light) have a higher frequency of UV and violet cones and a lower frequency of yellow and red cones than animals from a swamp population (low-transmission UV/blue light). Visual sensitivity does not vary significantly between the populations, but spring animals tend to be more sensitive in the UV/blue wavelengths (360–440 nm) and less sensitive in longer wavelengths (560–600 nm) than swamp animals. The results have two important implications. First, the tight conservation of functional regions of opsin genes across taxa does not imply that visual systems are constrained in their evolution; differential sensitivity can arise through differential expression of cone classes within the retina. Second, intraspecific visual signals in this species may evolve to maximize contrast between the signaler and the background (as opposed to brightness); males with blue anal fins are most abundant in swamp habitats where animals express fewer UV and violet cones.Electronic Supplementary Material Supplementary material is available in the online version of this article at .     

Blue–green opponency and trichromatic vision in the greenhouse whitefly (Trialeurodes vaporariorum) explored using light emitting diodes

Visual orientation in the greenhouse whitefly (Trialeurodes vaporariorum Westwood, Hemiptera: Aleyrodidae) is the result of “wavelength‐specific behaviours.” Green–yellow elicits “settling behaviour” while ultraviolet (UV) radiation initiates “migratory behaviour.” The only available physiological study of the photoreceptors' spectral efficiency showed peaks in the green and the UV range and whitefly vision was said to be dichromatic so far. In order to study the visual behaviour of T. vaporariorum, 19 narrow‐bandwidth light emitting diodes (LEDs) covering the UV‐A and visible range were used in combination with light scattering acrylic glass screens in a small‐scale choice arena under greenhouse conditions. Multiple‐choice and dual‐choice assays were performed, resulting in LED‐based behavioural action spectra of settling (green) and migratory behaviour (UV). A potential inhibitory blue–green chromatic mechanism was studied by combining yellow with different bluish LEDs. Intensity dependencies were illustrated by changing LED intensities. Regarding the “settling response,” highest attraction was achieved by a green LED with a centroid wavelength of 550 nm, while a blue LED with 469 nm proved to be most inhibitory. Besides this inhibitory interaction, an intensity dependence was observed within the action spectrum in the green–yellow range. “Migratory behaviour” was elicited the most by the UV LED with the shortest available wavelength of 373 nm. The results provide compelling behavioural evidence for the presence of a green and a yet undescribed blue sensitive photoreceptor and a blue–green opponent mechanism. Furthermore, empirical colour choice models were built and receptor peaks were estimated around 510–520 nm (green), 480–490 nm (blue) and 340–370 nm (UV). Consequently, a trichromatic receptor setup is suggested for T. vaporariorum.     

Optical releasers of the innate proboscis extension in the hoverfly Eristalis tenax L. (Syrphidae,Diptera)

Freshly emerged, inexperienced imagos of the hoverfly Eristalis tenax L. extend their proboscis towards small, yellow colour stimuli, such as anther parts and artificial floral guides. The releasing of this behaviour, which is adapted to pollen feeding, was investigated in behavioural tests using white, UV-reflecting artificial flowers with four small screens illuminated with test stimuli serving as artificial floral guides. The releasing of the innate proboscis extension was tested using monochromatic test lights. Within an intensity range from approx. 5·10¹¹ to approx. 10¹⁴ quanta · cm^-2· s^-1, the flies extended their proboscis only towards green and yellow test lights (approx. 520–600 nm). The inhibition of the innate proboscis extension was tested using mixed light stimuli composed of a yellow monochromatic reference light (560 nm, 10¹³ quanta·cm^{-2 -1}) and of a monochromatic test light. When the reference light was mixed with ultraviolet or blue test lights, the releasing of the innate proboscis extension was strongly inhibited, whereas admixing green/yellow light slightly promoted it; admixing red light had no effect. The results indicate that the releasing of the innate proboscis extension is mediated by the photoreceptor type R8y. Other receptor types which could cause the inhibition of the proboscis reaction are discussed.     

Action spectra of the female's response in the firefly Photinus pyralis (Coleoptera: Lampyridae): evidence for an achromatic detection of the bioluminescent optical signal

Behavioural action spectra of the threshold of the Photinus pyralis female response to light stimuli simulating the bioluminescent optical signal of the conspecific male firefly were determined in the laboratory. The action spectra (Fig. 1) were narrow and peaked in the yellow region of the spectrum. The females responded only to stimuli of wavelengths longer than 480 nm and not to stimuli in the blue (420-460 nm) part of the spectrum. The shape of the function corresponds with (a) the electroretinographic spectral sensitivity function in the long wavelength (520-660 nm) region of the spectrum, (b) the action spectrum of the female response (Fig. 1), (c) the species yellow bioluminescence emission spectrum and (d) the action spectrum of the intracellular response from single retinular cells (Fig. 2) of the compound eyes in the firefly. Such a correspondence suggests that the narrow yellow receptors of the female mediate the detection and processing of the optical signal of the conspecific male. Since the bioluminescent optical signal is processed exclusively by a single receptor class, signal detection is achromatic.     

Colour choice behaviour in the pollen beetle Meligethes aeneus (Coleoptera: Nitidulidae)

The pollen beetle Meligethes aeneus Fabricius (Coleoptera, Nitidulidae), a pest of oilseed rape (Brassica napus), is known to respond to coloured stimuli; however, current understanding of the underlying mechanisms of colour choice in this species is limited. In the present study, physiological and behavioural experiments are conducted to determine the response of the pollen beetle to colours in the field. Spectral sensitivity is measured in 10 animals using the electroretinogram technique. Light flashes (100 ms) at varied wavelengths (340–650 nm, 10‐nm steps) and at different light intensities are applied to the eye after dark adaptation. In behavioural experiments in the field, 100 water traps of varying colours (from yellow to green to blue with varying amounts of white and black added, and with known spectral reflectance) are set out on a bare soil field in May 2008. The mean spectral sensitivity curve of M. aeneus peaks at 520 nm; however, a model template fitted to the long wavelength tail of the observed curve reveals a peak at approximately 540 nm (green). A secondary sensitivity peak is observed in the ultraviolet (UV) range (370 nm). A total of 2482 pollen beetles are captured in the coloured traps. The results show that the pollen beetles' preference for yellow over other colours can be modelled as a colour opponent mechanism (green versus blue); however, further experiments are needed to specify responses to colours with higher UV reflectance. These findings may be used to optimize trap colours for monitoring to help develop integrated pest management strategies for pollen beetle control.     

Acoustic stimuli from predators trigger behavioural responses in aggregate caterpillars

Prey detect their predators through predator signals and cues and, consequently, respond with anti‐predatory behaviours to inhibit the action of their aggressors. Lepidopterans can intercept signals emitted by predators and may defend themselves through chemical, morphological or behavioural responses. In this study, we investigated the effect of acoustic stimuli of different predators on defensive behaviour of gregarious caterpillars. Our results demonstrated that Hylesia nigricans (Lepidoptera, Saturniidae) caterpillars alter their behaviour (i.e. abruptly raising the head) in response to the acoustic stimulus of the predators (i.e. predation risk signals from birds and wasps). The magnitude of this response depended on predator identity and caterpillar body size. Larger caterpillars responded more strongly to predatory stimuli than smaller caterpillars. However, regardless of the size of the caterpillars, they responded more strongly to the stimuli of wasps. In addition, we identified that H. nigricans caterpillars emit ultrasonic noise after detecting the stimuli of the predators – this noise seems to function as an alert about the risk of predation during the early stages of development (second and fifth instars). The duration of ultrasonic emission (i.e. milliseconds) increases with the number of repetitions of the stimuli (i.e. wing‐beat sounds of the wasps and insectivorous birds). These results provide novel information about predation risk in interactions among caterpillars and their predators, and indicate possible communication among invertebrates mediated by the risk of predation.     

The disruption of dominance hierarchies by a non-native species: an individual-based analysis

We studied the effects of the exotic rainbow trout (Oncorhynchus mykiss) on the performance and the dominance hierarchy of native Atlantic salmon (Salmo salar) at the group and individual level using laboratory and semi-natural experiments. At the group level, we compared the effects of interspecific and intraspecific competition (substitutive and additive design) on behavioural responses and growth of young-of-the-year Atlantic salmon. At the individual level, the same design was used to evaluate: (1) the temporal consistency of behavioural responses, dominance hierarchy and growth rate of Atlantic salmon; (2) the pattern of correlations between behaviours; and (3) the relationship between individual growth rate and behaviour. In the laboratory, group-level analyses revealed a weak but similar effect of rainbow trout and intraspecific competition on the behaviour and growth of Atlantic salmon. In contrast, individual-based analyses demonstrated that rainbow trout (but not intraspecific competition) strongly affected behavioural strategy, dominance hierarchy and growth trajectory of individual Atlantic salmon. Specifically, behaviours, dominance status and growth rate of salmon were temporally consistent in the intraspecific environment, while these patterns were disrupted when rainbow trout were present. Similarly, we found that rainbow trout strongly affected behavioural correlations and the relationships between individual growth rate and behaviour. The semi-natural experiments confirmed these results as interspecific competition affected relationships between individual growth rate of salmon, initial weight and activity index. Overall, individual-based analyses highlighted important mechanisms that were concealed at the group level, and that may be crucial to understand ecological and evolutionary consequences of exotic species. Moreover, these results demonstrated that competition with an exotic species disrupts the hierarchical relationship among native individuals and may therefore represent a potential for a shift in selective pressure.     

Acoustic communication in the pine engraver bark beetle: do signals vary between behavioural contexts?

Acoustic communication is taxonomically widespread in bark beetles and is proposed to play an important role in a variety of social and defensive behavioural contexts. Yet our understanding of how signals vary between contexts is currently limited. The present study tests the hypothesis that acoustic signals vary between behavioural contexts in the female pine engraver beetle Ips pini (Say) (Coleoptera: Curculionidae: Scolytinae). Female Ips pini produce acoustic chirps using a vertex‐pronotal stridulatory organ. Randomly sampled chirps generated under three contexts (i.e. distress, predation and premating) are compared for their duration, number of pulses, interpulse intervals, pulse rate and amplitude envelope shapes. The results obtained show that, during premating events, chirps are significantly longer in duration and tend to have a higher proportion of descending amplitude envelopes than chirps occurring during distress and predation events. Chirps produced during distress and predation conditions are indistinguishable from one another. By contrast to the results from previous bark beetle studies, no support is found for categorizing chirps as ‘interrupted’ or ‘uninterrupted’ types based on temporal patterns. The functional significance of context‐dependent variation in chirp characteristics is discussed. Previous studies on acoustic communication in bark beetles are limited as a result of a general lack of objective sampling and measurement criteria for characterizing signals. Recommendations are outlined for future studies on the functions and evolution of acoustic communication in bark beetles.     

Color change as a potential behavioral strategy

Within species, color morphs may enhance camouflage, improve communication and/or confer reproductive advantage. However, in the male cichlid Astatotilapia burtoni, body color may also signal a behavioral strategy. A. burtoni live in a lek-like social system in Lake Tanganyika, Africa where bright blue or yellow territorial (T) males (together ~ 10–30% of the population) are reproductively capable and defend territories containing food with a spawning site. In contrast, non-territorial (NT) males are smaller, cryptically colored, shoal with females and have regressed gonads. Importantly, males switch between these social states depending on their success in aggressive encounters. Yellow and blue morphs were thought to be adaptations to particular habitats, but they co-exist both in nature and in the laboratory. Importantly, individual males can switch colors so we asked whether color influences behavioral and hormonal profiles. When pairing territorial males with opposite colored fish, yellow males became dominant over blue males significantly more frequently. Moreover, yellow T males had significantly higher levels of 11-ketotosterone than blue T males while only blue NT males had higher levels of the stress hormone cortisol compared to the other groups. Thus color differences alone predict dominance status and hormone profiles in T males. Since T males can and do change color, this suggests that A. burtoni may use color as a flexible behavioral strategy.     

Lighting environment predicts the relative abundance of male colour morphs in bluefin killifish (Lucania goodei) populations

Animal communication occurs when an animal emits a signal, the signal is transmitted through the environment, and then detected by the receiver. The environment in which signalling occurs should govern the efficacy of this process. In this study, I examine the relationship of lighting environment (light transmission and tree cover), location and the relative abundances of male colour morphs across seven drainages and 30 populations in the bluefin killifish, Lucania goodei. I found that males with blue anal fins were more common in populations with low transmission of ultraviolet (UV) and blue wavelengths. By contrast, males with red anal fins (and to a lesser extent, males with yellow anal fins) were more common in populations with high transmission of UV and blue wavelengths. High UV-blue light transmission should create a blue visual background and may make blue males less conspicuous and red males more conspicuous to conspecifics. Colour contrast with the visual background may be more important than total brightness of the colour pattern. These results indicate that natural selection for effective intraspecific communication drives the relative abundance of male colour morphs in different lighting habitats.     

Peripheral encoding of behaviorally relevant acoustic signals in a vocal fish: single tones

The midshipman fish, Porichthys notatus, generates acoustic signals for intraspecific communication. Nesting males produce long-duration “hums” which attract gravid females and can be effectively mimicked by pure tones. In this study we examine the encoding of tonal signals by the midshipman peripheral auditory system. Single-unit recordings were made from afferents innervating the sacculus while presenting sounds via an underwater loudspeaker. Units were characterized by iso-intensity spike rate and vector strength of synchronization curves, as well as by peri-stimulus time histograms. Additionally, response-intensity curves and responses to long-duration (up to 10 s) stimuli were obtained. As has been seen in other teleosts, afferents had highly variable activity profiles. Excitatory frequencies ranged from 60 to over 300 Hz with most units responding best around 70 or 140 Hz. Thresholds at 90 Hz ranged from 95 to 145 dB re 1 μPa. Strong synchronization provided a robust temporal code of frequency, comparable to that described for goldfish. Spike rate showed varying degrees of adaptation but high rates were generally maintained even for 10-s stimuli. The midshipman peripheral auditory system is well suited to encoding conspecific communication signals, but nonetheless shares many response patterns with the auditory system of other teleosts. Accepted: 10 February 1999     

The Role of Wing Pigmentation, UV and Fluorescence as Signals in a Neotropical Damselfly

Pigmentation patterns, ultraviolet reflection and fluorescent emission are often involved in mate recognition and mate quality functions in many animal taxa. We investigated the role of wing ultra-violet reflection, fluorescence emission, and pigmentation on age and sexual signals in the damselfly Mnesarete pudica. In this species, wings are sexually dimorphic in colour and exhibit age dependency: males and females show a smoky black colouration when young, turning red in mature males while it turns brown in females. First, we investigated wing UV patterns through reflectance and emission spectra. Second, behavioural experiments were undertaken to show male and female responses to manipulated wing pigmentation and experimentally reduced UV (UV-). Reflectance spectra of the wings of juvenile and mature males and females were used to show the differences between controls and individuals with manipulated colouration used in the behavioural experiment. UV-reduced, females with wings painted red, and control males and females were tethered and presented to conspecific males and females, and their behavioral responses were recorded. The male red wing pigmentation and females with red wings elicited an aggressive response in territorial males and a sexual response in females. Both males and females showed neutral responses towards individuals with reduced UV. Wing signals of juvenile individuals also provoked neutral responses. These results suggest that UV, together with pigmentation, plays a role during mate recognition in males and females. Other than butterflies and spiders, it seems that fluorescence signals and UV reflectance can also be part of communication in odonates.     

Search behaviour of two hemipteran species using vibrational communication

The ability of conspecifics to recognize and locate each other in the environment depends on the efficiency of intraspecific communication. We compared the mate searching strategies of southern green stinkbug Nezara viridula (male searches for a continuously calling female) and the leafhopper Aphrodes makarovi (partners form a precisely coordinated duet). Males of both species were tested on plants in playback experiments. One leaf was vibrated with unaltered conspecific female signals or with various conspecific signals using modified temporal parameters. The results showed that the onset of searching was faster in A. makarovi than in N. viridula. Changes in temporal parameters of female replies had negative effect on the searching behaviour of A. makarovi. Males located the source of longer female replies faster than the short female call and they failed to locate the source of a female reply with temporal parameters outside the species-specific values. In contrast, in N. viridula, searching males successfully located also the source of a female song with parameters outside the species-specific values. The results are discussed with regard to male behavioural strategies in species with different vibrational communication systems and different male mating investment.     

Responses of the blister beetle Hycleus apicicornis to visual stimuli

Insect attraction to host plants may be partly mediated by visual stimuli. In the present study, the responses of adult Hycleus apicicornis (Guér.) (Coleoptera: Meloidae) to plant models of different colours, different combinations of two colours, or three hues of blue of different shapes are compared. Single‐colour models comprised the colours sky blue, bright green, yellow, red, white and black. Sky blue (reflecting light in the 440–500 nm region) is the most attractive, followed by white, which reflects light over a broader range (400–700 nm). On landing on sky blue targets, beetles exhibit feeding behaviour immediately. When different hues of blue (of different shapes) are compared, sky blue is preferred over turquoise, followed by dark blue, indicating that H. apicicornis is more attracted to lighter hues of blue than to darker ones. No significant differences are found between the three shapes (circle, square and triangle) tested, suggesting that reflectance associated with colour could be a more important visual cue than shape for host location by H. apicicornis. The preference of H. apicicornis for sky blue can be exploited in designing an attractive trap for its management.     

Spectral sensitivity of the compound eyes in two day-active fireflies (Coleoptera: Lampyridae: Lucidota)

Summary The electroretinographic visual spectral sensitivity functions in day-active fireflies Lucidota luteicollis and Lucidota atra show a broad green sensitivity and a shoulder in the near-ultraviolet region of the spectrum (Figs. 1, 2) as is commonly found among day-active insects. The nomogram for P530 visual pigment matches the spectral sensitivity curves in the green. The adult L. luteicollis retains its larval bioluminescent light organ which has a peak emission at 562 nm. The _max of the ERG spectral sensitivity does not match the bioluminescent peak (Fig. 1B) as it does in twilight- and dark-active fireflies. Some relevant behavioural observations with respect to mating are presented.