Modification of spectral sensitivities by screening pigments in the compound eyes of twilight-active fireflies (Coleoptera: Lampyridae)

1. ERG S(lambda) were determined in dark-adapted intact preparations of 6 North American firefly species (Photinus collustrans, marginellus, pyralis, macdermotti, scintillans and Bicellonycha wickershamorum) which restrict their flashing activity to twilight hours. The curves possess narrow (1/2 bandwidth = 50-60 nm) peaks in the yellow (560-580 nm) and a shoulder in the violet (370-420 nm), with a marked attenuation (1.4-2.2 log units) of sensitivity in the green (480-530 nm) region of the spectrum (Fig. 1). Two additional species (Photuris potomaca and frontalis) which initiate flashing at twilight and continue on late into the night (twi-night) possess broad sensitivity maxima around 560 nm (Fig. 3). 2. Selective adaptation experiments isolated near-UV and yellow in P. scintillans (Fig. 2). In the dorsal frontal region of the compound eyes in P. frontalis, high sensitivity existed only in the short wavelength region (near-UV and blue) with a maximum in the blue (lambda max 435 nm) (Fig. 4). 3. The in situ MSP absorption spectrum of the screening pigments was determined in preparations of firefly retina. a) Two kinds of dark brown granules were found in the clear zone region. These granules absorb all across the spectrum with a gradual increase in optical density in the shorter wavelength region in P. pyralis (Fig. 5). b) Besides dark granules, pink-to-red colored screening pigments were present in the vicinity of the rhabdoms. The absorption spectra of these pigments determined in five species were narrow (1/2 bandwidth = 50-80 nm) with species-specific differences in their peak absorption in the green at 525 nm, 510 nm, 512 nm and 517 nm in P. scintillans, macdermotti, collustrans and pyralis, respectively (Fig. 6). A similar pigment was found in P. marginellus with a lambda max at 512 nm (Fig. 7). In all cases, transmission increased both at long and short wavelengths, but more sharply in the long wavelength region (Figs. 6 and 7). Hence each twilight-restricted species has its own unique colored screening pigment. A yellow pigment whose absorption spectrum differed from those found in genus Photinus was found in twi-night active Photuris potomaca (lambda max 461 nm) and night-active P. versicolor (lambda max 456 nm). The transmission of the Photuris pigment increased sharply only in the long wave-length region (Fig. 8).(ABSTRACT TRUNCATED AT 400 WORDS)     

Discrimination of Variation in a Male Signaling Trait Affects Detection Time in Visual Predators

In many species, expression of elaborate male characteristics likely represents a balance between sexual selection and natural selection via predation, as male traits selected to elicit rapid detection or response on the part of females also increase detection by predators. Predation costs are frequently inferred, but the underlying mechanisms associated with specific traits have rarely been directly explored. Males of the wolf spider Schizocosa ocreata (Araneae: Lycosidae) exhibit a sexually selected signaling trait (dark tufts of bristles) on their forelegs, and are sympatric with a number of visually hunting generalist predators, including cannibalistic conspecifics, that may impact spider survival. Here we use latency of orientation response of the American toad, Bufo americanus (Anura: Bufonidae), to video, 'virtual' courting S. ocreata male stimuli as an index of predator detection, and latency of orientation response of female S. ocreata to the same stimuli as an index of conspecific detection. When compared with stimuli representing the population average, elimination of the signal trait had no significant effect on predator detection but did increase latency to orient in conspecifics. Increasing the size of the signal trait had no effect on conspecific detection but did significantly reduce latency to orient for predatory toads. Results clearly indicate that for a courting male spider of a given size and vigor level, variation in the expression of a secondary sexual characteristic alone can incur differential direct costs and benefits by influencing latency of orientation to visual signals by predators and conspecifics.     

Conspecific odor detection by the male broad-headed skink, Eumeces laticeps: effects of sex and site of odor source and of male reproductive condition

Olfactory stimuli are sufficient for detection and discrimination of sex of conspecific lizards by the male broad-headed skink, Eumeces laticeps, a member of a large group of lizards with pronounced chemosensory abilities, the Autarchoglossa. The capacity of male broad-headed skinks to detect conspecific odors was assessed by measuring tongue extrusion rates in response to odor stimuli presented on moist cotton applicators. Tongue-flick rates of postreproductive males were significantly higher for cloacal odors of postreproductive conspecifics of both sexes than to distilled water and higher to female than male odors over the initial 20- and 60-sec intervals. In a second experiment using testosterone-treated males and estrogen-injected females, testosterone-treated males emitted significantly more tongue flicks to female cloacal odors than to the other stimuli, and two males bit applicators bearing male odors. Testosterone did not affect reaction to male cloacal odors, but markedly increased tongue-flick rates in response to cloacal odors of estrogen-treated females. Postreproductive males also responded to female, but not male, skin odors at a significantly higher rate than to water. Possible sources and presumed adaptive significance of conspecific odors are discussed.     

Action spectra for the photoperiodic control of polymorphism in the aphid Megoura viciae

Energy compensated action spectra are given for the photoperiodic control of polymorphism in Megoura. The production of ‘long day’ parthenogenetic virginoparae and ‘short day’ oviparae mainly depend on the night length. Light has three different effects. ‘Early’ interruptions of the dark phase in a long night cycle reverse the time-measuring dark response. The action spectrum for a 1 hr interruption placed 1.5 hr after the onset of darkness (during dark stage 1) shows a relatively narrow band of activity, mainly in the blue (450–470 nm). The threshold is ca. 0.25 μW cm^?2. ‘Late’ interruptions placed 7.5 or 8 hr after dark hour 0 (during dark stage 3) strongly promote the production of virginoparae without causing a reversal of the response. The action spectrum has the same blue maximum but sensitivity extends into the yellow and red spectral regions. The third photosensitive component, the main photoperiod itself, is required for initiating the dark timing response and has an intermediate action spectrum. Time/intensity curves for a single wavelength (471 nm) show that the responses during stages 1 and 3 depart markedly from reciprocity. Short durations cannot be compensated by high intensities. The shape of the reciprocity curve for an ‘early’ interruption suggests that the stage 1 response is complete after ca. 1.25 hr.The action spectra are believed to be compatible with the view that the photoreceptor is a caroteno-protein. It is suggested that all three pigment forms are related and that time measurement is largely a function of spontaneous ‘dark reaction’ changes in the pigment system. Stage 1 may represent the reversible conjugation phase of the protein/chromophore moieties. In Stage 2 the pigment is presumably photorefractory and is transitional to the highly sensitive broad spectrum form of stage 3.     

Visual scanning by common marmosets (Callithrix jacchus): Functional aspects and the specialrrole of adult males

The rates of visual scanning and vocalizations were studied in a group of captive marmosets (Callinthrix jacchus) after the presentation of five different stimuli (artificial flower, playback of long calls, female/male conspecific, stuffed wild cat) in order to assess the function of visual scanning. Only the presentation of a stuffed cat induced a significant response. The adult males in particular increased their scanning rate and all adult individuals decreased their rate of vocalizations. The results indicate that visual scanning in marmosets is an appropriate measure of vigilance which seems to serve the function of predator detection and avoidance. The strong response of adult males suggests that they play a special role in predator detection.     

Spectral mechanisms of the compound eye in the fireflyPhotinus pyralis (Coleoptera: Lampyridae)

Summary Electroretinograms (ERG) were recorded from dark- and chromatic-adapted compound eyes in the dusk-active firefly,Photinus pyralis , at different wavelengths ranging from 320 to 700 run and over 4.5 log units change in stimulus intensity. ERG waveforms differed in the short (near-UV and violet) and long (yellow) wavelengths (Fig. 1). Waveform differences were quantitated by analysis of rise and fall times as a function of the amplitude of the response. Rise times were found to be relatively constant for all stimulus wavelengths. However, variations in the fall times were detected and followed characteristically different functions for short and long wavelengths (Fig. 2).No significant differences in the slopes of the Vlog-I curves at different stimulus wavelengths were observed (Fig. 3).Spectral sensitivity curves obtained from the ventral sector in dark- and chromatic-adapted conditions revealed peaks in the short ( max 400 nm: Fig. 4; max 430 nm: Fig. 5 A; and max 380 nm; Fig. 5B) and long ( max 570 nm: Figs. 4, 5) wavelengths, suggesting the presence of two spectral mechanisms. The long wavelength (yellow) mechanism was in close tune with the species bioluminescence emission spectrum (Fig. 4B).This investigation was supported in part by NIH Research Grant # EY-00490 (to R.M.C.); Research Grant # 01794N from the Research Foundation of the City University of New York (to A.B.L.); NIGMS Training Grant #1 TO 2 GM 05010-01 MARC (to J.A.H.); and NSF Grant # HES-75-09824 (to C.O.T.). We thank Tom Jensen for technical assistance, Barry Schuttler for his courtesy in allowing us to collect fireflies at his farm, Jean Lall for editorial assistance, and the two anonymous referees whose comments added considerably to the quality of this paper.     

Sharp intense line in the bioluminescence emission of the firefly

Numerous investigations have been carried out on the spectral distribution of the light of different species of fireflies. Here we record the emission spectrum of the Indian species of the firefly Luciola praeusta Kiesenwetter 1874 (Coleoptera : Lampyridae : Luciolinae) on a color film. Green and red color-sectors, with an intense yellow one in between, appear in this spectrum. Intensity profile of this spectrum reveals a hitherto undetected strong narrow yellow line, which lies within the full-width-at-half maximum (FWHM) of the intensity profile. The spectrum recorded in a high-resolution spectrometer confirms the presence of this sharp intense line. This finding lends support to an earlier drawn analogy between the in vivo emission of the firefly and laser light.     

Spectral efficiency of the glasshouse whitefly Trialeurodes vaporariorum and Encarsia formosa its hymenopteran parasitoid

Using the electroretinogram (ERG) technique the spectral efficiency of the compound eye of the glasshouse whitefly Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) and its main parasitoid Encarsia formosa (Gahan) (Hymenoptera: Aphelinidae) was measured at selected wavelengths between 340 nm and 670 nm. The form of the ERG for both T. vaporariorum and E. formosa was found to be monophasic in nature. For both male and female T. vaporariorum and female E. formosa a primary peak of efficiency occurred in the blue-green-yellow region, peak 520 nm and a secondary peak in the ultraviolet (UV) region. The compound eye of female E. formosa gave a significantly greater response in the UV region than either the dorsal or ventral regions of the compound eye of T. vaporariorum relative to the responses in the blue-green-yellow region. T. vaporariorum has divided compound eyes with distinct dorsal and ventral regions. In this study it was found that the percentage response in the UV, of the dorsal region of the eye, is significantly greater than that of the ventral region of the eye relative to the percentage response in the blue-green-yellow region and there is a significant shift in the blue-green yellow peak towards the right of the spectrum.     

Discovery of signaling effect of UV-B/C light in the extended UV-A/blue-type action spectra for step-down and step-up photophobic responses in the unicellular flagellate algaEuglena gracilis

Summary Cultures of unicellular algal flagellateEuglena gracilis grown in different conditions were subjected to action spectroscopy for step-down and step-up photophobic responses, respectively. The spectral region was extended into the UV-B/C as well as in the UV-A and visible regions with the Okazaki Large Spectrograph as the monochromatic light source. The photophobic responses of the cells were measured with an individual-cell assay method with the aid of a computerized video motion analyzer. In the UV-A and visible regions, the shapes of the action spectra were the so-called UV-A/blue type. In the newly studied UV-B/C region, new action peaks were found at 270 nm for the step-down response and at 280 nm for the step-up one. The absorption spectrum of flavin adenine dinucleotide (FAD) appeared to fit the action spectrum for the step-up response, whereas the shape of the step-down action spectrum, which has a UV-A peak (at 370 nm) higher than the blue peak (at 450 nm), appeared to be mimicked by the absorption spectrum of a mixed solution of 6-biopterin and FAD. These observations might also account for the fact that the UV-B/C peak wavelength at 270 nm of the action spectrum for the step-down response is shorter by 10 nm than the action spectrum for the step-up response at 280 nm.Abbreviations FAD flavin adenine dinucleotide - FWHM spectral full width at half maximum - NIBB National Institute for Basic Biology - OLS Okazaki Large Spectrograph - PFB paraflagellar body - UV-A ultraviolet light of spectral region between 320 and 400 nm - UV-B/C ultraviolet light of spectral region between 190 and 320 nm     

条背萤的闪光求偶行为

水栖萤火虫条背萤Luciola substriata (Gorh.)发出单脉冲周期性特异闪光信号进行求偶。室外观测发现,在23℃、86% RH时,雄萤飞行求偶闪光信号脉冲闪光持续时间为0.52 s,间隔时间为0.28 s,闪光信号的最大亮度为0.6 lx; 雌萤求偶信号周期为 0.67 s,最大亮度为0.4 lx。雄萤发出求偶信号0.22 s后,雌萤发出两个连续的回应信号。第一个回应信号为0.49 s,第二个为0.41 s, 两个回应信号的间隔时间为0.11 s。雄萤发现雌萤回应信号后,降落至离雌萤5～10 cm处,继续发出闪光信号,但闪光脉冲频率减小,闪光脉冲时间为1.23 s,间隔时间为0.50 s。条背萤交配时呈“V”或“一”字形交配姿势。     

Neurophysiological response selectivity for conspecific songs over synthetic sounds in the auditory forebrain of non-singing female songbirds

Female choice plays a critical role in the evolution of male acoustic displays. Yet there is limited information on the neurophysiological basis of female songbirds’ auditory recognition systems. To understand the neural mechanisms of how non-singing female songbirds perceive behaviorally relevant vocalizations, we recorded responses of single neurons to acoustic stimuli in two auditory forebrain regions, the caudal lateral mesopallium (CLM) and Field L, in anesthetized adult female zebra finches (Taeniopygia guttata). Using various metrics of response selectivity, we found consistently higher response strengths for unfamiliar conspecific songs compared to tone pips and white noise in Field L but not in CLM. We also found that neurons in the left auditory forebrain had lower response strengths to synthetics sounds, leading to overall higher neural selectivity for song in neurons of the left hemisphere. This laterality effect is consistent with previously published behavioral data in zebra finches. Overall, our results from Field L are in parallel and from CLM are in contrast with the patterns of response selectivity reported for conspecific songs over synthetic sounds in male zebra finches, suggesting some degree of sexual dimorphism of auditory perception mechanisms in songbirds.     

The importance of olfactory signals in the gasterosteid mating system: sticklebacks go multimodal

Stickleback fishes are renowned for the complexity of their nuptial colour signal. In this paper I show that the nuptial signal is in fact multimodal: male-based olfactory cues also transmit information to receptive females. Both female three-spined and brook sticklebacks recognized the scent from each other's males, but discriminated in favour of their own males when asked to choose between conspecific and heterospecific odours. Although females were not attracted to scent from the more distantly related guppy, Poecilia reticulata , changes in their baseline behaviours indicated that they perceived its presence. Olfactory cues act as long distance messages, allowing a female to detect the 'I am here' message from the male before she can actually see him. Studies of interactions between temporally displaced signals indicate that the first cue (in this case chemical) functions to alert the receiver to the presence of the second cue (visual), increasing the probability of its detection and recognition. So, although olfactory cues do not appear to be as species-specific as visual cues, their function as alerting stimuli may not require such fine-tuning. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 555–572.     

The pigment system of the photoreceptor 7 yellow in the fly,a complex photoreceptor

Summary The 7y photoreceptor in the fly (Musca, Calliphora) retina harbours an unusually complex pigment system consisting of a bistable visual pigment (xanthopsin, X and metaxanthopsin, M), a blue-absorbing C₄₀-carotenoid (zeaxanthin and/or lutein) and a uv sensitizing pigment (3-OH retinol).The difference spectrum and photoequilibrium spectrum in single 7y rhabdomeres were determined microspectrophotometrically (Fig. 2).The extinction spectrum of the C₄₀-carotenoid has a pronounced vibrational structure, with peaks at 430, 450 and 480 nm (Fig. 3). The off-axis spectral sensitivity, determined electrophysiologically with 1 nm resolution shows no trace of this fine structure thus excluding the possibility that the C₄₀-carotenoid is a second sensitizing pigment (Fig. 4).The absorption spectra of X and M are derived by fitting nomogram spectra (based on fly R1–6 xanthopsin) to the difference spectrum. max for X is 425 nm, and for M 510 nm (Fig. 5). It is shown that the photoequilibrium spectrum and the difference spectrum can be used to derive the relative photosensitivity spectra of X and M using the analytical method developed by Stavenga (1975). The result (Fig. 6) shows a pronounced uv sensitivity for both, X and M, indicating that the uv sensitizing pigment transfers energy to both X and M. A value of 0.7 for, the relative efficiency of photoconversion for X and M, is obtained by fitting the analytically derived relative photosensitivity spectra to the absorption spectra at wavelengths beyond 420 nm.     

Differential Response to Colors by the Lizard Uta palmeri (Reptilia: Iguanidae)

Many lizards have brightly colored body parts thought to be signals, but color perception in lizards has not been well studied. We present data documenting differential response to colors in an iguanid lizard. Free-living Uta palmeri differentially approached and bit at a red-colored object when simultaneously presented objects of four different colors (red, blue, yellow, black). This response to red was statistically significant for all three age-sex classes of lizards tested (adult males, adult females, juveniles). In addition, juveniles and adult females exhibited intermediate levels of responses (approaching and sniffing, and licking) to the yellow object but adult males did not. Seasonally important food items of this species are red (fish regurgitated by nesting seabirds) and yellow (yolk from broken seabird eggs), and response behavior (biting versus licking) was similar to that exhibited during feeding. Thus, as presented in our experimental paradigm, red and yellow elicited feeding responses. The spectral reflectance curves of the red and yellow stimuli were found to exhibit peaks at wavelengths of 630 nm (range 570–670), and at 600 nm (range 550–700), respectively. These values overlap with the spectral range of the gravid coloration of female U. palmeri (peak 615 nm, range 550–690) that is thought to be a signal to conspecifics. Furthermore, the reflectance curves are similar in shape and height (intensity). Thus, the similarities of the reflectance curves of the red paint and of the natural coloring suggest that palmeri can detect the female gravid coloration. Whether the response to red objects is due to discriminating differences in hue or in relative brightness remains to be examined.     

The function of anal fin egg-spots in the cichlid fish Astatotilapia burtoni

Color and pigmentation patterns of animals are often targets of sexual selection because of their role in communication. Although conspicuous male traits are typically implicated with intersexual selection, there are examples where sex-specific displays play a role in an intrasexual context, e.g. when they serve as signals for aggression level and/or status. Here, we focus on the function of a conspicuous male ornament in the most species-rich tribe of cichlid fishes, the haplochromines. A characteristic feature of these ca. 1500 species are so-called egg-spots in form of ovoid markings on the anal fins of males, which are made up of carotenoid based pigment cells. It has long been assumed that these yellow, orange or reddish egg-spots play an important role in the courtship and spawning behavior of these maternal mouth-brooding fishes by mimicking the eggs of a conspecific female. The exact function of egg-spots remains unknown, however, and there are several hypotheses about their mode of action. To uncover the function of this cichlid-specific male ornament, we used female mate choice experiments and a male aggression test in the haplochromine species Astatotilapia burtoni. We manipulated the number and arrangement of egg-spots on the anal fins of males, or removed them entirely, and tested (1) female preference with visual contact only using egg-traps, (2) female preference with free contact using paternity testing with microsatellites and (3) male aggression. We found that females did not prefer males with many egg-spots over males with fewer egg-spots and that females tended to prefer males without egg-spots over males with egg-spots. Importantly, males without egg-spots sired clutches with the same fertilization rate as males with egg-spots. In male aggression trials, however, males with fewer egg-spots received significantly more attacks, suggesting that egg-spots are an important signal in intrasexual communication.     

Photomovement in Dunaliella salina: Fluence rate-response curves and action spectra

We determined the action spectra of the photophobic responses as well as the phototactic response in Dunaliella salina (Volvocales) using both single cells and populations. The action spectra of the photophobic responses have maxima at 510 nm, the spectrum for phototaxis has a maximum at 450–460 nm. These action spectra are not compatible with the hypothesis that flavoproteins are the photoreceptor pigments, and we suggest that carotenoproteins or rhodopsins act as the photoreceptor pigments. We also conclude that the phototactic response in Dunaliella is an elementary response, quite independent of the step-up and step-down photophobic responses. We also determined the action spectra of the photoaccumulation response in populations of cells adapted to two different salt conditions. Both action spectra have a peak a 490 nm. The photoaccumulation response may be a complex response composed of the phototactic and photophobic responses. Blue or blue-green light does not elicit a photokinetic response in Dunaliella.Diagrams of the optical set-ups used for measuring the responses at the single-cell level and of the plans for building the phototaxometer described in this paper are available to the interested readerWe thank Mr. M. Kubota for a tremendous amount of technical assistance and Mr. R. Nagy for building the phototaxometer. We thank T. Kondo, Professor H. Imaseki and the members of the Laboratory of Biological Regulation, NIBB, for their help and support in various aspects of this research. This research was supported, in part, from grants from the Okazaki Large Spectrograph (Project Nos. 86-535, 87-518, 88-523), the Japanese Society for the Promotion of Science, and the College of Agriculture and Life Sciences at Cornell University to R. W.     

Sex-allocation behaviour of a solitary ectoparasitoid: effects of host-patch characteristics and female density

The sex-allocation behaviour of the solitary ectoparasitoid Dinarmus basalis (Hymenoptera: Pteromalidae) was investigated by examining the female's response to two proximal factors: the host-patch characteristics and the conspecific female density in the patch. The offspring sex-ratio of single females presented with unparasitised hosts was female biased (approximately three daughters to one son in the progeny) whatever the host density tested. The sex-ratio in the presence of hosts parasitised by a conspecific 48 h beforehand was male biased. The proportion of male offspring also increased when the number of conspecific females exploiting the patch changed from a single female to a group of females, whatever the group size. No sexual differences in mortality as a result of larval competition on superparasitised hosts were observed. The offspring sex-ratios observed in response to the host-patch characteristics were the result of manipulation of the egg fertilisation by the females during the oviposition phase. Thus, the variability of the sex-ratio in response to an increase in the number of conspecific females in the same patch represented a reduction in egg fertilisation. These results are analysed with regard to the predictions of Local Mate Competition theory and the females' host discrimination ability.     

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     

Female mate choice and the potential for ornament evolution in túngara frogs Physalaemus pustulosus

The potential for ornament evolution in response to sexual selection rests on the interaction between the permissive-ness or selectivity of female preferences and the constraints on male development of signaling related traits. We investigate the former by determining how latent female preferences either exaggerate the magnitude of current traits (I.e. Elaborations) or favor novel traits (I.e. Innovations). In tungara frogs, females prefer complex mating calls (whine-chucks) to simple calls (whine only). The whine is critical for mate recognition while the chuck further enhances the attractiveness of the call. Here we use a combina-tion of synthetic and natural stimuli to examine latent female preferences. Our results show that a diversity of stimuli, including conspecific and heterospecific calls as well as predator-produced and human-made sounds, increase the attractiveness of a call when added to a whine. These stimuli do not make simple calls more attractive than a whine-chuck, however. In rare cases we found stimuli that added to the whine decrease the attractiveness of the call. Overall, females show strong preferences for both elaborations and innovations of the chuck. We argue that the emancipation of these acoustic adornments from mate recognition allows such female permissiveness, and that male constraints on signal evolution are probably more important in explaining why males evolved their specific adornment. Experimentally probing latent female preferences for stimuli out of the species' range is a useful means to gain insights about the potential of female choice to influence signal evolution and thus the astounding diversity in male sexually-selected traits.     

Female sexual arousal in amphibians

Rather than being a static, species specific trait, reproductive behavior in female amphibians is variable within an individual during the breeding season when females are capable of reproductive activity. Changes in receptivity coincide with changes in circulating estrogen. Estrogen is highest at the point when females are ready to choose a male and lay eggs. At this time female receptivity (her probability of responding to a male vocal signal) is highest and her selectivity among conspecific calls (measured by her probability of responding to a degraded or otherwise usually unattractive male signal) is lowest. These changes occur even though females retain the ability to discriminate different acoustic characteristics of various conspecific calls. After releasing her eggs, female amphibians quickly become less receptive and more choosy in terms of their responses to male sexual advertisement signals. Male vocal signals stimulate both behavior and estrogen changes in amphibian females making mating more probable. The changes in female reproductive behavior are the same as those generally accepted as indicative of a change in female sexual arousal leading to copulation. They are situationally triggered, gated by interactions with males, and decline with the consummation of sexual reproduction with a chosen male. The changes can be triggered by either internal physiological state or by the presence of stimuli presented by males, and the same stimuli change both behavior and physiological (endocrine) state in such a way as to make acceptance of a male more likely. Thus amphibian females demonstrate many of the same general characteristics of changing female sexual state that in mammals indicate sexual arousal.