Central Mechanisms of Frog Galling

The central mechanisms of release calling and mating callingin the Northern leopard frog (Rana pipiens pipiens) and thegreen tree frog (Hyla cinerea) are discussed. The region ofthe ventral magnoceHifl-ar preoptic nucleus is essential formiating calling. This area probably contains androgen receptorsthat project to, and activate, more posterior calling mechanisms.The region of the main sensory nucleus V is involved, in a mannernot yet clearly understood, in calling. Perhaps this area servesas a sensory correlation center that receives and analyzes avariety of sensory inputs, determines what type of call constitutesan appropriate response, and then excites the calling circuitsat more posterior levels. The region of the hypoglossal andvagus motor nuclei is essential for release calling. This areaprobably serves as a motor coordination center that organizesthe motor patterns of calling.     

Lampenbürstenchromosomen in den Oocytenkernen von Sepia officinalis

The nuclei of the growing oocytes of Sepia officinalis contain lamp-brush chromosomes with clearly demonstrable chromomeres bearing pairs of lateral loops (Fig. 4). Similar to the findings of Callan and Lloyd (1960) in amphibian oocytes the chromosomes of Sepia exhibit some specially organized loops which can be used as landmarkers for recognizing certain chromosomes (Fig. 5). Maximal loop despiralization occurs only in nuclei of follicles during their initial growth phase. The following developmental period is characterized by a successive contraction of loops and chromosome axes (Fig. 7), despite the fact that the diameter of the oocyte nucleus enlarges still three times. In the oldest nuclei hundreds of nucleolus-like granules arise near the chromosomes which later spread over the entire nucleus (Fig. 9).

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Herrn Professor Dr. K. Bier () in dankbarer Verehrung gewidmet.     

Sexual selection at varying population densities in male field crickets,Gryllus veletis andG. pennsylvanicus

Male field crickets call and attract females or they silently search for females in the vicinity of calling males. At high population densities, fewer calling sites are available, defense of calling sites is costly, and an increased proportion of matings should result from searching behavior. To test these predictions, individually marked field crickets, Gryllus veletisand G. pennsylvanicus,were observed for 10 h nightly in large outdoor arenas at relatively high and low densities (2020 and 55, males and females). Data were gathered on body weight, calling duration, movement, and mating frequency for individual crickets. These observations showed that variance in male mating success was significantly greater at a low density in G. pennsylvanicus,and calling duration correlated with mating success at this density. Direct selection on a trait was estimated as the partial regression coefficient (selection gradient, ) and the total selection was estimated (direct and indirect selection on correlated traits) as the covariance (standardized intensity of selection, s) of the trait on the relative mating success. Direct selection favored increased movement at a high density in G. veletis,and direct and total selection favored increased calling duration at a low density in G. pennsylvanicus.Most other comparisons were not statistically significant. The data are discussed in terms of density-dependent fluctuations in sexual selection on correlated male traits.     

Territoriality and Male Mating Success in the Dart-poison Frog,Epipedobates femoralis (Dendrobatidae,Anura)

The social and reproductive behaviour of the dart-poison frog, Epipedobates femoralis, was studied in Amazonian Peru for 14 months. Males defended territories with advertisement calls and, ultimately, fighting. Territory size ranged from 0.25 to 26 m² and was positively correlated with duration of residence and calling activity of the owner. Females were not territorial and were never attacked when approaching calling males. Males and females only mated once and females sampled calling males before mating. Male mating success was closely correlated with territory size and calling activity. No correlation was found between male body size and mating success. Territories provide residents with sufficient space for mate attraction and reproduction without interference from rivals. Since territory size is dependent on calling activity which involves high energetic costs, it is suggested that territory size reveals male quality.     

Calling behavior of the female german cockroach,Blattella germanica (Dictyoptera: Blattellidae)

Virgin German cockroach females, Blattella germanica(L.), were observed, for the first time, to exhibit a characteristic calling behavior during which females emit a volatile sex pheromone. Under a photoperiod of 12L12D, the percentage of 7-day-old virgin females that exhibited this behavior peaked before the end of the scotophase in a similar pattern to the diel periodicity of mating. A clear relationship was evident between calling and stages of sexual receptivity during successive gonotrophic cycles. Females initiated calling 5–6 days after the imaginal molt, when their basal oocytes were 1.6 mm long. If not mated, females continued to exhibit bouts of calling during the next 3–4 days until 24 h before ovulation. Calling was completely suppressed by mating as well as the presence of an egg case in the genital atrium in both virgin and mated gravid females. We suggest that calling and the emission of a volatile sex pheromone serve to attract males from a distance as well as to potentiate responses to contact sex pheromone in aggregations.     

Multiscale determinants of tree frog (Hyla arborea L.) calling ponds in western Switzerland

A tree frog (Hyla arborea L., 1758) metapopulation in western Switzerland was studied during spring 2001. All potential calling ponds in an area of 350km² were searched for tree frog calling males. Twenty-nine out of 111 ponds sheltered between 1 and 250 callers. Most ponds were occupied by less than 12 males. Pond parameters were measured at three different levels using field analysis and a Geographical Information System (GIS). The first level was water chemistry and pond-associated measures. The second level was the surrounding land use in a 30m buffer around the pond. The third level consisted of landscape indices on a broader scale (up to 2km). Logistic regression was applied to identify parameters that can predict the presence of calling males in a pond. Response variable was the presence or absence of callers. Four significant parameters allowed us to explain about 40% of the total deviance of the observed occupational pattern. Urbanization around the pond had a highly negative impact on the probability of presence of calling males. Hours of direct sunlight on the pond was positively correlated with callers. Higher water conductivity was associated with a lesser probability of species presence. Finally, the further the closest two-lane road, the higher the probability of callers presence. Our results show that presence or absence of callers is influenced by factors acting at various geographical scales.     

Acoustic orientation in adult, female crickets (Gryllus bimaculatus de Geer) after unilateral foreleg amputation in the larva

Summary InGryllus bimaculatus females one foreleg was amputated at the coxa-trochanter joint in the 2nd, 4th or 8th/9th larval instar. A leg of up to normal length is regenerated (Fig. 1) but it lacks a functional ear. In spite of the, usually shorter, regenerated foreleg, the adult one-eared crickets show no impairments in walking when tested on a locomotion compensator. Without sound they walk erratically and most of them weakly circle towards the intact side (Fig. 2).With calling song presentation three response types can be distinguished:tracking (Fig. 3A), hanging on (Fig. 3B) or continuouscircling towards the intact side (Fig. 3C, D). Turning tendencies in monaurals increase with song intensity and exceed those of intact and bilaterally operated animals (Fig. 4). Course deviations towards the intact side also slightly increase with intensity (Fig. 5). Course stability is reduced compared to that of intact animals but exceeds that of bilaterally operated crickets (Figs. 5, 6). It is best at 60 dB and deteriorates at higher sound intensities (Fig. 6). The percentage of monaurals tracking or hanging on decreases with increasing intensity (Fig. 7B). Tracking is established in most animals but it is limited to a narrow intensity range (Fig. 7A, C). Apart from an increased percentage of tracking after early operations (Fig. 7D), there are no prominent changes in orientational parameters with the date of foreleg amputation.Reamputation of the regenerated leg in the adult monaurals does not significantly impair acoustic orientation (Figs. 8, 9), but occlusion of the ipsilateral prothoracic spiracle does (Figs. 10, 11).An attempt is made to correlate the behavioral performance with the activity of auditory interneurons which have undergone morphological and physiological changes (Fig. 12).     

Mating-induced loss of sex pheromone and sexual receptivity in insects with emphasis on Helicoverpa zea and Lymantria dispar

Mating in most species of insects leads to a transient or permanent loss in sexual receptivity of the females. Among moths, this loss of receptivity is often accompanied with a loss of the sex pheromone in the absence of calling, which also could be temporary or permanent. Most of the earlier work on changes in reproductive behavior after mating was done with Diptera in which sperm and/or male accessory gland secretions were shown to be responsible for termination of receptivity. In the corn earworm moth, Helicoverpa zea, mated females become depleted of pheromone and become nonreceptive to further mating attempts, but only for the remainder of the night of mating. A pheromonostatic peptide isolated from the accessory glands of males may be responsible for the depletion of pheromone, while the termination of receptivity is independently controlled. In the gypsy moth, Lymantria dispar, the changes in behavior following mating are permanent. In this species, the switch from virgin to mated behavior involves three steps: a physical stimulation associated with mating, transfer of viable sperm to the spermatheca, and commencement of oviposition. Signals generated by these factors operate through neural pathways and, unlike in H. zea, accessory gland factors seem not to be involved. © 1994 Wiley-Liss, Inc.

1 This article is a US Government work and, as such, is in the public domain in the United States of America.

     

The spectral sensitivity of a littoral annelid:Nereis mediator

Summary The spectral sensitivities of the four eyes ofNereis mediator (Polychaeta, Annelida) were measured from 400 nm to 600 nm by determining the reciprocal of the number of quanta required to elicit a constant electroretinogram of 350 V.N. mediator is maximally sensitive at 480 nm, although it is very sensitive over a broad range from 400 nm to 540 nm (Fig. 4). Selective adaptation experiments (Fig. 5) and changes in the time characteristics of the electroretinogram under conditions of dark and green adaptation (Fig. 2) indicate the presence of a multi-receptor system.We thank Dr. Olga Hartman and Dr. Kristian Fauchald for their help with the natural history and identification of specimens. We also thank Josephine Yingst for her help in collection of specimens. This research was supported by a USC Biomedical Grant 5730 and by AFOSR Contract F 44620-70-C-0113.     

The 23 kDa Polypeptide from Common Frog Lens: Isolation,Properties, and Cellular Localization

The major water-soluble polypeptide with molecular weight of approximately 23 kDa (the 23 kDa polypeptide) was identified in the lens of common frog Rana temporaria L. According to the gel filtration data, the peptide is a part of an oligomeric protein with molecular weight over than 300 kDa (-crystallin fraction). A highly pure fraction of the 23 kDa polypeptide was isolated by two-step ion-exchange chromatography and SDS electrophoresis and the specific antibodies were obtained. Immunohistochemistry showed the presence of the 23 kDa polypeptide in the cytoplasm of lens epithelial cells (including its central region) and in the zones neighboring the plasma membranes in the cortical fibers. The 23 kDa polypeptide was not found in the lens central zone (nucleus). It was also present in the retina (in the cells of inner nuclear layers), but not in the other tissues and organs of adult frog. Immunochemical analysis showed that the 23 kDa polypeptide was different from all known crystallins of frogs and other animals (bull, mouse, rat, and chicken). The nature of the 23 kDa polypeptide and the relation of its expression with the lens cell differentiation are discussed.     

Behavioral experiments on the visual processing of color stimuli inPieris brassicae L. (Lepidoptera)

Summary In spontaneous-choice experiments on the butterflyPieris brassicae L. (Pieridae), spectral-effectiveness and spectral-sensitivity functions were obtained for various behaviors.Pilot experiments with colored PVC films, for which the relative number of reflected quanta with regard to the given illumination had been calculated, showed that the feeding response is distinctly intensity-dependent (Fig. 4). The animals are also capable of color discrimination independent of this intensity discrimination;P. brassicae prefers blue to other colors (e.g., orange, red and purple) with higher relative quantum numbers (Fig. 3) and distinguishes golden yellow and red from gray shades as well as from black and white (Fig. 5a, b).The results of subsequent spontaneous-choice experiments, using as stimuli monochromatic lights with known quantum flux, indicate that the various visually controlled functional categories of behavior can be assigned to the following spectral regions (Figs. 6, 8): 1. The open-space reaction corresponds to the UV and violet region, ca. 320–420 nm; 2. The feeding reaction corresponds to the blue region, ca. 420–500 nm, and the orange-red region, ca. 590–610 nm; 3. Egg-laying and drumming correspond to the green-yellow region, ca. 520–580 or 590 nm, respectively. The intensity dependence of the individual responses is again apparent in these experiments with monochromatic light stimuli (Figs. 7, 11, 12a).Even at very high intensities and when the content of the relevant wavelength is high, white light is practically ineffective for the feeding reaction (Fig. 9), drumming and egg-laying (cf. Results), regardless of its UV content. The open-space reaction, however, can be elicited by white light according to its UV content (Fig. 12 b).P. brassicae cannot be trained to give a feeding response to monochromatic light stimuli (Fig. 10).Experiments with mixtures of wavelengths have shown that the combination of the two maxima in the spectral sensitivity curve for the feeding reaction (=600 plus 447 nm) is just as effective as =447 nm alone (Fig. 13, left). Moreover, the mixture producing the hypothetical Pieris purple (=600 plus 370 nm) is no more or less effective in eliciting the feeding and open-space reactions than the more effective component for each of these reactions when presented alone (Fig. 13, right). With the mixture of =600 plus 558 nm, both the feeding reaction and drumming are distinctly reduced (Fig. 13, middle). This mixed color, unlike the other two mixtures tested, has a color quality different from that of the component colors.That the behavior ofP. brassicae is exclusively wavelength-specific can thus be ruled out. There are indications that wavelength-specific behavior and color vision are both present.Abbreviation RNQ relative number of quanta This publication is dedicated to Professor Dr. Dr.h.c. H. Autrum on the occasion of his 80th birthday     

Two auditory filter systems determine the calling behavior of the fire-bellied toad: a behavioral and neurophysiological characterization

Summary Male fire-bellied toads,Bombina bombina, are stimulated into calling by conspecific mating calls. They tend to synchronize their activity with other calling individuals, which leads to antiphonal calling. Behavioral physiological experiments revealed the existence of two different auditory filter systems which influence this acoustic behavior. One filter system is tuned to the recognition of conspecific mating calls, the other is part of a circuit that controls an acoustic jamming avoidance reaction. The two filter systems differ in several properties: filter bandwidth, threshold, and time constant. Neurophysiological recordings in the midbrain and the thalamus show that the filter characteristics can be explained by the coding properties of auditory neurons in these two brain nuclei.     

Anuran mating calling circuits: inhibition by prostaglandin

Male American toads (Bufo americanus) were induced to mating call in response to electronically simulated, conspecific mating calls. The injection of prostaglandin (PG) F2 alpha caused suppression of mating call answering. Neural correlates of mating calling were triggered by electrical stimulation of the anterior preoptic nucleus in the isolated brainstem of male, Northern leopard frogs (Rana p. pipiens). The addition of PGF2 alpha to the bath completely abolished the correlates of mating calling without changing the correlates of pulmonary respiration. The suppression of mating calling shown here, along with the suppression of release signalling described by Diakow and Nemiroff (1981), supports the hypothesis of a close interrelation between the neural circuits of these two calls. The suppression of the neural correlates of mating calling in an isolated preparation shows a central site of action of the PG. The retention of normal correlates of pulmonary respiration, even after suppression of mating calling correlates, suggests that the generation of mating calling patterns involves the extension and pulsing of the expiratory phase of breathing.     

Age-dependent pattern of calling behavior in <Emphasis Type="Italic">Atheloca subrufella</Emphasis> (Hulst) (Lepidoptera: Phycitidae)

Changes in the signaling effort of virgin females are known to occur until their first mating. Young females tend to increase the calling effort (number and time duration of bouts) as they age. In this context, the calling behavior of virgin females of Atheloca subrufella, an important pest of coconut palms (Cocos nucifera), was studied under laboratory conditions. Thirty four virgin females were observed during 16 scotophases (25 ± 2 °C, 75 ± 5 % RH and 10 h scotophase), until they stopped calling in the following photophase. The calling position, pattern and frequency were evaluated. Females of A. subrufella showed only one calling position, and calling peaked between the second and fifth scotophases. Most females did not start calling immediately after emergence. Neither the individual duration of calling nor the number of calling bouts changed over time. However, the time that calling started was earlier in older females, probably to avoid competition with younger females and increase the probability of mating. These data support the hypothesis that there is an age-dependent pattern of pheromone emission in virgin females of A. subrufella.     

The Timing of Calling and Mating in <Emphasis Type="Italic">Heterolocha jinyinhuaphaga</Emphasis> and the Influence of Environmental Determinants

We investigated the periodicities of calling and mating of Heterolocha jinyinhuaphaga (Lepidoptera: Geometridae) and the effects of temperature, relative humidity, and photoperiod on the timing of these behaviors. Females called on the first night after emergence, mainly during the second half of scotophase. The calling percentage increased on the second night and peaked on the third night, before decreasing. Mating had similar rhythms but peaked 1 h later than calling. As the moth aged, the mean onset times of calling and mating were advanced. Temperature, relative humidity, and photoperiod all affected the calling and mating. We found that 25°C was the most suitable temperature for calling and mating, whereas at higher or lower temperatures, the total calling percentage and the total mating percentage decreased, the onset time of calling was delayed, and mating duration decreased. Increased the relative humidity facilitated calling and mating. However, lengthened photoperiods inhibited the calling and mating.     

Acoustic communication in the trilling field cricket,Gryllus rubens (Orthoptera: Gryllidae)

Male field crickets produce calling songs that are used for attracting conspecific females for mating. Acoustic communication was studied in the trilling field cricket, Gryllus rubens.A trilled calling song consists of a continuous train of sound pulses, each of which was repeated at a stereotyped rate. Singing males were recorded at different temperatures. The pulse period of the calling song decreased with increasing temperature; there was less effect of temperature on pulse duration. Female phonotaxis was studied on a noncompensating, spherical treadmill. In two-stimulus (choice) playback experiments, females preferred the conspecific trilled song over the chirped calling song of a sympatric species, Gryllus fultoni.This preference persisted even when the song of G. fultoniwas 6 dB louder. Females also discriminated between synthetic trills having different pulse periods; females chose trills with the conspecific pulse period over trills having lower and higher pulse periods.     

Genetic interactions indicate a role for Mdg1p and the SH3 domain protein Bem1p in linking the G-protein mediated yeast pheromone signalling pathway to regulators of cell polarity

The pheromone signal in the yeastSaccharomyces cerevisiae is transmitted by the and subunits of the mating response G-protein. TheSTE20 gene, encoding a protein kinase required for pheromone signal transduction, has recently been identified in a genetic screen for high-gene-dosage suppressors of a partly defective G mutation. The same genetic screen identifiedBEM1, which encodes an SH3 domain protein required for polarized morphogenesis in response to pheromone, and a novel gene, designatedMDG1 (multicopy suppressor ofdefectiveG-protein). TheMDG1 gene was independently isolated in a search for multicopy suppressors of abem1 mutation. TheMDG1 gene encodes a predicted hydrophilic protein of 364 amino acids with a molecular weight of 41 kDa that has no homology with known proteins. A fusion of Mdg1p with the green fluorescent protein fromAequorea victoria localizes to the plasma membrane, suggesting that Mdg1p is an extrinsically bound membrane protein. Deletion ofMDG1 causes sterility in cells in which the wild-type G has been replaced by partly defective G derivatives but does not cause any other obvious phenotypes. The mating defect of cells deleted forSTE20 is partially suppressed by multiple copies ofBEM1 andCDC42, which encodes a small GTP-binding protein that binds to Ste20p and is necessary for the development of cell polarity. Elevated levels ofSTE20 andBEM1 are capable of suppressing a temperature-sensitive mutation inCDC42. This complex network of genetic interactions points to a role for Bem1p and Mdg1p in G-protein mediated signal transduction and indicates a functional linkage between components of the pheromone signalling pathway and regulators of cell polarity during yeast mating.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

The Energetics of Male Mating Strategies in Field Crickets (Orthoptera: Gryllinae: Gryllidae)

Male field crickets (Gryllinae: Gryllidae: Orthoptera) vary in their mating strategies, particularly in whether they call from defended sites to attract phonotactic females or roam silently in search of potential mates. To better understand the economics underlying these alternative strategies, respirometry was employed to examine the energetic costs of each strategy's component behaviors for a representative species, Acheta domesticus. Advertisement calling in this species, composed of low pulse rate chirps, is an order of magnitude less energetically costly than walking per unit time. However, for gryllids that advertise call with higher pulse rate trills, calling and walking appear to be of equivalent energetic cost. Thus, if energetic costs are important in determining grylline mating strategies, trillers and chirpers should have different sensitivities to change in factors affecting the relative payoffs of each strategy. Field studies of changes in mating behavior with increased population density support this argument. This study also determined that courtship calling by A. domesticus was over twice as energetically costly as advertisement calling per unit time, suggesting its importance as a more reliable indicator of signaler, or mate, quality.     

Territory size as a main driver of male-mating success in an Amazonian nurse frog (<Emphasis Type="Italic">Allobates paleovarzensis</Emphasis>, Dendrobatoidea)

In polygamous mating systems, it is most often males that compete for the opposite sex, using strategies to provide access to as many females as possible. Females, on the other hand, constitute the sex that exerts the choice and so require a means of accessing the quality of a potential partner in comparison to its competitors. A common challenge in sexual selection studies is to identify the most relevant trait for mating success, since many are correlated with each other. In addition, little is known about how the female accesses the aspects related to male quality. In this context, we tested the role of different male characteristics on mating success in a natural environment using the Amazonian frog Allobates paleovarzensis as a model. A multiple linear regression model showed a positive relationship between territory size and number of male matings, while calling persistence was slightly related to the mating success. We did not detect a relation of the number of matings with the distance to the nearest body of water nor with male body size. Additionally, we observed that territory size was not related to calling persistence, but had a positive relation with the duration of the couple’s courting process. Thus, we conclude that: (1) territory size is the main determinant of male-mating success, and this is not correlated with the other attributes tested; and (2) females access the size of the males’ territory through the courting process that precedes oviposition.