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Females prefer carotenoid colored males as mates in the pentamorphic livebearing fish, Poecilia parae.

The first results of female preference and chosen male mating success in a new model organism, the pentamorphic livebearing fish, Poecilia parae, are presented. Poecilia parae is a relative of the guppy, P. reticulata, and is assumed to have similar reproductive behavior. We tested the hypothesis that P. parae females, like female guppies, prefer carotenoid colored males as mates. Here we show that the time a female spent with males was significantly greater for carotenoid coloration in red and yellow melanzona, but time with these two morphs did not differ. The preferred red and yellow males mated significantly more often with their choosing females than did the non-preferred blue and parae males. The few blue melanzona and parae males that mated did so without performing courtship displays. Some females mated with all phenotypes including immaculata males during open group trials. Female P. parae clearly preferred males with carotenoid coloration, thereby corroborating the hypothesis. Alternative male mating tactics by blue melanzona, parae, and immaculata morphs and promiscuous mating by females also resembled features of reproductive behaviors exhibited by guppies.

Animals↗

Does exaggerated morphology preclude plasticity to cuckoldry in the midshipman fish ( Porichthys notatus)?

In species with more than one male reproductive morph, there typically exists a larger morph with exaggerated secondary sexual characters, and a smaller morph with reduced secondary sexual characters. These "exaggerated" and "reduced" morphologies are commonly thought to represent specializations to alternative behavioral reproductive tactics-large body size and exaggerated secondary sexual characters should both facilitate territoriality, courtship, and pair-spawning; while small body size and reduced secondary sexual characters should facilitate "sneaky" cuckoldry. Given this postulated relationship between morphology and behavior, we examined the relationship between the morphology of exaggerated males and cuckoldry. In a field and aquarium study of the midshipman fish, a fish with both exaggerated and reduced morphs, we demonstrated cuckoldry in some males of the exaggerated morph. Since the "reduced" morphology is thought to be an adaptation towards sneaky cuckoldry, we predicted that, of males with the exaggerated morph, less-exaggerated (smaller) males would be better able to gain proximity to the spawning pair during cuckoldry. In contrast to that prediction, access to the spawning pair during cuckoldry increased with the body size of the cuckolding exaggerated-morph males. This may be related to our observation that exaggerated males often cuckolded aggressively. Thus the "exaggerated" morphology need not preclude adaptive plasticity to cuckoldry, and may even aid it.

Animals↗

When males whistle at females: complex FM acoustic signals in cockroaches.

Male cockroaches of the species Elliptorhina chopardi expel air through a pair of modified abdominal spiracles during courtship. This air expulsion simultaneously produces air and substrate-borne vibrations. We described and compared in details these two types of vibrations. Our analysis of the air-borne signals shows that males can produce three categories of signals with distinct temporal and frequency parameters. "Pure whistles" consist of two independent harmonic series fast frequency modulated with independent harmonics that can cross each other. "Noisy whistles" also possess two independent voices but include a noisy broad-band frequency part in the middle. Hiss sounds are more noise-like, being made of a broad-band frequency spectrum. All three call types are unusually high in dominant frequency (>5 kHz) for cockroaches. The substrate-borne signals are categorised similarly. Some harmonics of the substrate-borne signals were filtered out, however, making the acoustic energy centered on fewer frequency bands. Our analysis shows that cockroach signals are complex, with fast frequency modulations and two distinct voices. These results also readdress the question of what system could potentially receive and decode the information contained within such complex sounds.

Abdomen↗

Salmonid opsin sequences undergo positive selection and indicate an alternate evolutionary relationship in oncorhynchus.

Positive selection can be demonstrated by statistical analysis when non-synonymous nucleotide substitutions occur more frequently than synonymous substitutions (dN>dS). This pattern of sequence evolution has been observed in the rhodopsin gene of cichlids. Mutations in opsin genes resulting in amino acid (AA) replacement appear to be associated with the evolution of specific color patterns and the evolution of courtship behaviors. Within fish, AA replacements in opsin proteins have improved vision at great depths and have occurred in deep-sea species. Salmonids experience diverse photic environments during their life history. Furthermore, sexual selection has resulted in species-specific male and female coloration during spawning. To look for evidence of positive selection in salmonid opsins, we sequenced the RH1, RH2, LWS, SWS1, and SWS2 genes from six Pacific salmon species as well as the Atlantic salmon. These salmonids include landlocked and migratory species and species that vary in their coloration during spawning. In each opsin gene comparison from all species sampled, traditional dN:dS analysis did not indicate positive selection. However, the more sensitive Creevey-McInerney statistical analysis indicates that RH1 and RH2 experienced positive selection early in the evolution and speciation of salmonids.

Animals↗

Diffusive transport in Stokeslet flow and its application to plankton ecology.

In this paper we consider the advective/diffusive transport of a solute near a hovering zooplankter. We approximate the fluid flow with that of a Stokeslet, corresponding to the plankter exerting a point force on the water, and assume that the plankter acts as a point source for the transported solute, located at the same point as the force. We find an analytical expression in closed form for the steady-state concentration of the solute. We also discuss the situation where the plankter performs Brownian motion. Finally we apply the results to the courtship of the marine copepod Pseudocalanus elongatus, where the male performs a mating dance below the hovering female. For this situation, our model supports the hypothesis that the mating dance is guided by the plume of a signalling pheromone.

Animals↗

Immediate early gene response to hearing song correlates with receptive behavior and depends on dialect in a female songbird.

Stimulus-induced expression of the immediate early gene ZENK (egr-1) in the songbird's auditory forebrain presumably depends on the behavioral significance of the stimulus. Few studies, however, have quantified both the ZENK and behavioral responses to a stimulus in the same individuals. We played conspecific male song of either hatch (local) or foreign dialect to female white-crowned sparrows (Zonotrichia leucophrys oriantha) and quantified both the auditory ZENK response and their behavioral response, which is known to depend on dialect. Birds hearing hatch dialect showed greater ZENK induction in the caudomedial hyperstriatum ventrale and the dorsal portion of the caudomedial neostriatum than birds hearing foreign dialect, supporting previous work showing a relationship between ZENK and salience of the stimulus. In the dorsal portion of the caudomedial neostriatum, ZENK induction was correlated with the amount of non-vocal courtship behavior; however, in the caudomedial hyperstriatum ventrale, ZENK induction was more highly correlated with the females' own vocal behavior and thus may have been partly self-induced. Some females sang and showed a male-like pattern of ZENK induction in their song systems. This study provides the first evidence that the ZENK response in a sensory area to a social stimulus is proportional to the animal's preference for the stimulus.

Animals↗

Influence of temperature and reproductive state upon the jamming avoidance response in the pulse-type electric fish Brachyhypopomus pinnicaudatus.

The electric organ discharge (EOD) in Brachyhypopomus pinnicaudatus is modified by temperature and reproductive state. We studied the influence of these variables upon a complex behavior, the jamming avoidance response (JAR). Experiments were performed in non-reproductive fish and in two groups of fish after the induction of reproductive state (by nature or by acclimation at 28 degrees C). JARs were elicited at 20 and 30 degrees C by free-run electric stimuli with different deltaLs (interval difference between stimulus and EOD). In non-reproductive fish, JARs induced by stimuli with +deltaLs showed temperature sensitivity, with smaller responses at 30 degrees C. Conversely, similar JARs were obtained at both temperatures in reproductive fish. These observations were replicated in curarized preparations. Stimuli with -deltaLs were almost ineffective in non-reproductive fish at 30 degrees C, whereas adequate JARs were shown by reproductive fish. Phase-locked stimuli were used to evaluate the duration of the low-threshold electrosensory periods preceding and following the EOD. In non-reproductive fish, the temperature step induced a shortening of these periods. The opposite effect was observed in reproductive fish, probably explaining the differences in JAR capability. A prolongation of the low-threshold periods would favor the perception of electrocommunication signals during courtship. JAR changes would be a consequence of this adaptation.

Animals↗

Juggling with volatiles: exposure of perfumes by displaying male orchid bees.

Male orchid bees (Euglossini) pollinate 10% of the neotropical orchid flora while collecting floral scents, which they store and accumulate in hind tibial pouches. The purpose of these fragrances is unclear, as is the context, timing and mechanism of their possible exposure. Here we show for the first time that males expose and relocate their fragrances during courtship display. We present high-speed video analyses revealing an intricate and repetitive leg movement performed by displaying male Euglossa cognata. The behavior involves several morphological structures of hitherto unknown function and suggests transfer of substances from the hind tibia to a contralateral mid-tibial tuft of hairs. Body-side-specific fluorescent dye application and consecutive detection of signals on males after display confirmed this transfer. Deposited on the mid-tibial tufts, the fragrances are ideally placed in order to become ventilated by jugal combs on the wing bases, as previously suggested by Bembé (in Apidologie 35:288-291, 2004). Being clearly distinct from motor patterns involved in fragrance collection, the described movement is continuously performed by displaying males, suggesting an equally continuous exposure of volatiles. Although the findings strengthen, the view that the volatiles serve as attractants in the context of mating behavior, the signal addressee, conspecific males or females, has yet to be found.

Animal Communication↗

Measuring and quantifying dynamic visual signals in jumping spiders.

Animals emit visual signals that involve simultaneous, sequential movements of appendages that unfold with varying dynamics in time and space. Algorithms have been recently reported (e.g. Peters et al. in Anim Behav 64:131-146, 2002) that enable quantitative characterization of movements as optical flow patterns. For decades, acoustical signals have been rendered by techniques that decompose sound into amplitude, time, and spectral components. Using an optic-flow algorithm we examined visual courtship behaviours of jumping spiders and depict their complex visual signals as "speed waveform", "speed surface", and "speed waterfall" plots analogous to acoustic waveforms, spectrograms, and waterfall plots, respectively. In addition, these "speed profiles" are compatible with analytical techniques developed for auditory analysis. Using examples from the jumping spider Habronattus pugillis we show that we can statistically differentiate displays of different "sky island" populations supporting previous work on diversification. We also examined visual displays from the jumping spider Habronattus dossenus and show that distinct seismic components of vibratory displays are produced concurrently with statistically distinct motion signals. Given that dynamic visual signals are common, from insects to birds to mammals, we propose that optical-flow algorithms and the analyses described here will be useful for many researchers.

Algorithms↗

Female sex pheromone of a wandering spider (Cupiennius salei): identification and sensory reception.

Females of the wandering spider Cupiennius salei attach a sex pheromone to their dragline. Males encountering the female dragline examine the silk thread with their pedipalps and then typically initiate reciprocal vibratory courtship with the sexual partner. The female pheromone was identified as (S)-1,1'-dimethyl citrate. The male pheromone receptive sensory cells are located in tip pore sensilla and respond to touching the sensillum tip with female silk or pieces of filter paper containing the synthetic pheromone.

Animal Communication↗

Electrical stimulation of the preoptic area in Eigenmannia: evoked interruptions in the electric organ discharge.

The functional role of the basal forebrain and preoptic regions in modulating the normally regular electric organ discharge was determined by focal brain stimulation in the weakly electric fish, Eigenmannia. The rostral preoptic area, which is connected with the diencephalic prepacemaker nucleus, was examined physiologically by electrical stimulation in a curarized fish. Electrical stimulation of the most rostral region of the preoptic area with trains of relatively low intensity current elicits discrete bursts of electric organ discharge interruptions in contrast to other forebrain loci. These responses were observed primarily as after-responses following the termination of the stimulus train and were relatively immune to variations in the stimulus parameters. As the duration and rate of these preoptic-evoked bursts of electric organ discharge interruptions (approximately 100 ms at 2 per s) are similar to duration and rate of natural interruptions, it is proposed that these bursts might be precursors to natural interruptions. These data suggest that the preoptic area, consistent with its role in controlling reproductive behaviors in vertebrates, may be influencing the occurrence of electric organ discharge courtship signals by either direct actions on the prepacemaker nucleus or through other regions that are connected with the diencephalic pre-pacemaker nucleus.

Action Potentials↗

Acoustic communication underground: vocalization characteristics in subterranean social mole-rats (Cryptomys sp., Bathyergidae).

In captive adult Zambian mole-rats 14 different sounds (13 true vocalizations) have been recorded during different behavioural contexts. The sound analysis revealed that all sounds occurred in a low and middle frequency range with main energy below 10 kHz. The majority of calls contained components of 1.6-2 kHz, 0.63-0.8 kHz, and/or 5-6.3 kHz. The vocalization range thus matched well the hearing range as established in other studies. The frequency content of courtship calls in two species of Zambian Cryptomys was compared with that in naked mole-rats (Heterocephalus glaber) and blind mole-rats (Spalax ehrenbergi) as described in the literature. The frequency range of maximum sound energy is negatively correlated with the body weight and coincides with the frequencies of best hearing in the respective species. In general, the vocalization range in subterranean mammals is shifted towards low frequencies which are best propagated in underground burrows.

Acoustics↗

Diversity of sexual dimorphism in electrocommunication signals and its androgen regulation in a genus of electric fish, Apteronotus.

Gymnotiform electric fish emit an electric organ discharge that, in several species, is sexually dimorphic and functions in gender recognition. In addition, some species produce frequency modulations of the electric organ discharge, known as chirps, that are displayed during aggression and courtship. We report that two congeneric species (Apteronotus leptorhynchus and A. albifrons) differ in the expression of sexual dimorphism in these signals. In A. leptorhynchus, males chirp more than females, but in A. albifrons chirping is monomorphic. The gonadosomatic index and plasma levels of 11-ketotestosterone were equivalent in both species, suggesting that they were in similar reproductive condition. Corresponding to this difference in dimorphism, A. leptorhynchus increases chirping in response to androgens, but chirping in A. albifrons is insensitive to implants of testosterone, dihydrotestosterone or 11-ketotestosterone. Species also differ in the sexual dimorphism and androgen sensitivity of electric organ discharge frequency. In A. leptorhynchus, male discharge at higher frequencies than females, and androgens increase electric organ discharge frequency. In A. albifrons, males discharge at lower frequencies than females, and androgens decrease electronic organ discharge frequency. Thus, in both chirping and electric organ discharge frequency, evolutionary changes in the presence or direction of sexual dimorphism have been accompanied and perhaps caused by changes in the androgen regulation of the electric organ discharge.

Androgens↗

"Allohormones": a class of bioactive substances favoured by sexual selection.

During close bodily contact, many species transfer substances that influence the behaviour or physiology of conspecifics. Such transfer is especially common during courtship and copulation. When this is the case the involved bioactive substances are favoured by sexual selection because their effects include increased egg production, inhibited remating, and changed sperm transport or storage in the partner. The direct mode of action of these substances is fundamentally different from that of pheromones and nuptial gifts. Therefore, the term allohormone is introduced here. An allohormone is defined as a substance that is transferred from one individual to another free-living member of the same species and that induces a direct behavioural or physiological response, bypassing external sensory organs. Although we emphasise the importance of allohormones in reproductive processes, allohormones may also have important functions outside of copulation.

Animals↗

Functional conservation and divergence of intersex, a gene required for female differentiation in Drosophila melanogaster.

In Drosophila melanogaster, somatic sexual differentiation is regulated by a well characterized genetic hierarchy, by which the ratio of X chromosomes to autosomes (X:A) ultimately directs the deployment of sex-specific transcription factors encoded by doublesex (dsx) and fruitless (fru). In other dipterans, the X:A ratio is not the primary sex-determination signal. Correspondingly, the Drosophila hierarchy is not fully conserved. In all non-drosophilid fly species examined, Sex-lethal (Sxl), the master switch at the top of the Drosophila hierarchy, does not control somatic sex. This rapid divergence contrasts with the apparently deep conservation of dsx, which in Drosophila controls virtually all aspects of somatic sex except for male courtship behavior (which is controlled by fru). Sex-specific dsx mRNAs have been reported in Diptera, Lepidoptera and Hymenoptera, and dsx homologs in nematodes and mammals are required for aspects of male differentiation. Thus, it seems that the bottom of the hierarchy is rather ancient, especially compared with the top. To test this, we cloned insect and vertebrate homologs of the Drosophila gene intersex (ix), which functions together with dsx at the bottom of the hierarchy in females. When expressed in D. melanogaster females mutant at the endogenous ix gene, dipteran and lepidopteran ix homologs restore proper sexual differentiation, substantiating the hypothesis that ix, like dsx, is broadly conserved. When the mouse homolog is expressed it produces a dominant-negative phenotype suggesting partial functional divergence. Our results raise the possibility that a functional association between ix- and dsx-related gene products existed before the origin of the bifunctional dsx gene used in insect sex determination.

Amino Acid Sequence↗

The transformer2 gene in Musca domestica is required for selecting and maintaining the female pathway of development.

We present the isolation and functional analysis of a transformer2 homologue Mdtra2 in the housefly Musca domestica. Compromising the activity of this gene by injecting dsRNA into embryos causes complete sex reversal of genotypically female individuals into fertile males, revealing an essential function of Mdtra2 in female development of the housefly. Mdtra2 is required for female-specific splicing of Musca doublesex (Mddsx) which structurally and functionally corresponds to Drosophila dsx, the bottom-most regulator in the sex-determining pathway. Since Mdtra2 is expressed in males and females, we propose that Mdtra2 serves as an essential co-factor of F, the key sex-determining switch upstream of Mddsx. We also provide evidence that Mdtra2 acts upstream as a positive regulator of F supporting genetic data which suggest that F relies on an autocatalytic activity to select and maintain the female path of development. We further show that repression of male courtship behavior by F requires Mdtra2. This function of F and Mdtra2 appears not to be mediated by Mddsx, suggesting that bifurcation of the pathway at this level is a conserved feature in the genetic architecture of Musca and Drosophila.

Amino Acid Sequence↗

Characterization of the dCaMKII-GAL4 driver line whose expression is controlled by the Drosophila Ca2+/calmodulin-dependent protein kinase II promoter.

Transgenic flies that can drive GAL4 expression under the control of the 7 kb 5'-region of the Drosophila Ca(2+)/calmodulin-dependent protein kinase II (dCaMKII) gene (dCaMKII-GAL4) were established. Characteristic features of this dCaMKII-GAL4 driven reporter expression were compatible with the endogenous dCaMKII expression pattern: The dCaMKII-GAL4 driven reporter gene was expressed preferentially in the central nervous system of the embryo and larvae. Reporter expression was also observed in the brain, thoracic ganglion, and gut of the adult. The whole-brain distribution and projections of dCaMKII-GAL4-expressing cells in the adults were visualized three-dimensionally by using UAS-linked reporter genes. Prominent signals of nuclear-localized beta-Gal reporter gene expression were found in extensive brain regions, especially in the Kenyon cells of the mushroom body (MB), cells in the pars intercerebralis, and subesophageal ganglion (SOG). tau reporter gene expression highlighting neurite projections was detected in the MB lobes, median bundle, antennal lobe glomeruli, and fibers of clusters in the SOG, ventrolateral protocerebrum and superior lateral protocerebrum. These observations agree with those of a previous study mapping the dCaMKII-dependent memory circuits in courtship conditioning. Interestingly, green fluorescent protein reporter gene expression in adult MB lobes was predominantly observed in the alpha and beta lobes with a core-deficient pattern, but not in the alpha' and beta' lobes, similar to Fasciclin II immunoreactivity.

Animals↗

Localization of prolactin receptor in the newt brain.

In the male newt Cynops pyrrhogaster, prolactin (PRL) acts directly on the central nervous system and induces courtship behavior. As a step to elucidate the localization of neurons on which PRL acts, we developed a polyclonal antibody against an oligopeptide having a sequence completely identical with a part of the sequence of PRL receptors (PRLRs) of two species of newts, C. pyrrhogaster and C. ensicauda, and performed an immunohistochemical study with this antibody. PRLR-immunoreactive cells were observed in the medial amygdala, anterior preoptic area, magnocellular preoptic nucleus, suprachiasmatic nucleus, nucleus of the periventricular organ, ventral hypothalamic nucleus, and choroid plexus. We also performed in situ hybridization with a (35)S-labeled newt PRLR antisense RNA probe and detected signals in the preoptic area and choroid plexus. Colocalization of both PRLR-like immunoreactivity and arginine vasotocin-like or mesotocin-like immunoreactivity was demonstrated in the magnocellular preoptic nucleus. This is the first report of PRLR localization in the amphibian brain.

Amygdala↗