Development of Communication Behaviour: Receiver Ontogeny in Túngara Frogs and a Prospectus for a Behavioural Evolutionary Development

Most studies addressing the development of animal communication have focused on signal production rather than receiver decoding, and similar emphasis has been given to learning over nonlearning. But receivers are an integral part of a communication network, and nonlearned mechanisms appear to be more ubiquitous than learned ones in the communication systems of most animals. Here we review the results of recent experiments and outline future directions for integrative studies on the development of a primarily nonlearned behaviour—recognition of communication signals during ontogeny in a tropical frog. The results suggest that antecedents to adult behaviours might be a common feature of developing organisms. Given the essential role that acoustic communication serves in reproduction for many organisms and that receivers can exert strong influence on the evolution of signals, understanding the evolutionary developmental basis of mate recognition will provide new insights into the evolution of communication systems

Distribution And Abundance Of Glucocorticoid And Mineralocorticoid Receptors Throughout The Brain Of The Great Tit (Parus Major)

The glucocorticoid stress response, regulated by the hypothalamic-pituitary-adrenal (HPA) axis, enables individuals to cope with stressors through transcriptional effects in cells expressing the appropriate receptors. The two receptors that bind glucocorticoids—the mineralocorticoid receptor (MR) and glucocorticoid receptor (GR)—are present in a variety of vertebrate tissues, but their expression in the brain is especially important. Neural receptor patterns have the potential to integrate multiple behavioral and physiological traits simultaneously, including self-regulation of glucocorticoid secretion through negative feedback processes. In the present work, we quantified the expression of GR and MR mRNA throughout the brain of a female great tit (Parus major), creating a distribution map encompassing 48 regions. This map, the first of its kind for P. major, demonstrated a widespread but not ubiquitous distribution of both receptor types. In the paraventricular nucleus of the hypothalamus (PVN) and the hippocampus (HP)—the two brain regions that we sampled from a total of 25 birds, we found high GR mRNA expression in the former and, unexpectedly, low MR mRNA in the latter. We examined the covariation of MR and GR levels in these two regions and found a strong, positive relationship between MR in the PVN and MR in the HP and a similar trend for GR across these two regions. This correlation supports the idea that hormone pleiotropy may constrain an individual’s behavioral and physiological phenotype. In the female song system, we found moderate GR in hyperstriatum ventrale, pars caudalis (HVC), and moderate MR in robust nucleus of the arcopallium (RA). Understanding intra- and interspecific patterns of glucocorticoid receptor expression can inform us about the behavioral processes (e.g. song learning) that may be sensitive to stress and stimulate future hypotheses concerning the relationships between receptor expression, circulating hormone concentrations and performance traits under selection, including behavior

Novelty Induces Behavioural And Glucocorticoid Responses In A Songbird Artificially Selected For Divergent Personalities

Stress physiology is thought to contribute to individual differences in behaviour. In part this reflects the fact that canonical personality measures consist of responses to challenges, including novel objects and environments. Exposure to novelty is typically assumed to induce a moderate increase in glucocorticoids (CORT), although this has rarely been tested. We tested this assumption using great tits, Parus major, selected for divergent personalities (bold-fast and shy-slow explorers), predicting that the shy birds would exhibit higher CORT following exposure to a novel object. We also scored behavioural responses to the novel object, predicting that bold birds would more frequently approach the novel object and exhibit more abnormal repetitive behaviours. We found that the presence of a novel object did induce a moderate CORT response, but selection lines did not differ in the magnitude of this response. Furthermore, although both selection lines showed a robust CORT elevation to a subsequent restraint stressor, the CORT response was stronger in bold birds and this effect was specific to novel object exposure. Shy birds showed a strong positive phenotypic correlation between CORT concentrations following the novel object exposure and the subsequent restraint stress. Behaviourally, the selection lines differed in their response during novel object exposure: as predicted, bold birds more frequently approached the novel object and shy birds more strongly decreased overall locomotion during the novel object trial, but birds from both selection lines showed significant and similar frequencies of abnormal repetitive behaviours during novel object exposure. Our findings support the hypothesis that personality emerges as a result of correlated selection on behaviour and underlying endocrine mechanisms and suggest that the relationship between endocrine stress physiology and personality is context dependent

Social Communication Across Reproductive Boundaries: Hormones And The Auditory Periphery Of Songbirds And Frogs

Most animals experience reproductive transitions in their lives; for instance, reaching reproductive maturity or cycling in and out of breeding condition. Some reproductive transitions are abrupt, while others are more gradual. In most cases, changes in communication between the sexes follow the time course of these reproductive transitions and are typically thought to be coordinated by steroid hormones. We know a great deal about hormonal control of communication behaviors in birds and frogs, as well as the central neural control of these behaviors. There has also been significant interest in the effects of steroid hormones on central nervous system structures that control both the production and reception of communication signals associated with reproductive behaviors. However, peripheral sensory structures have typically received less attention, although there has been growing interest in recent years. It is becoming clear that peripheral sensory systems play an important role in reproductive communication, are plastic across reproductive conditions, and, in some cases, this plasticity may be mediated by steroid hormones. In this paper, we discuss recent evidence for the role of peripheral auditory structures in reproductive communication in birds and frogs, the plasticity of the peripheral auditory system, and the role of steroid hormones in mediating the effects of the peripheral auditory system on reproductive communication. We focus on both seasonal and acute reproductive transitions, introduce new data on the role of hormones in modulating seasonal patterns, and make recommendations for future work

The Waiting And Mating Game: Condition Dependent Mate Sampling In Female Gray Treefrogs (Hyla Versicolor)

Strong sexual selection by receivers can lead to the evolution of elaborate courtship behaviors in signalers. However the process by which receivers sample signalers and execute mate choice under complex signaling conditions—and thus the realized strength of sexual section—is poorly understood. Moreover, receivers can vary in condition, which can further influence mate sampling strategies. Using wild female frogs we tested two hypotheses at the intersection of these important problems: that some of the individual variation in mate sampling is explained by (1) the reproductive urgency hypothesis, which predicts that receivers in a more urgent reproductive state will sample mates less and/or (2) the reproductive investment hypothesis, which predicts that receivers that have invested less in the current reproductive effort will sample mates less. Eastern gray treefrogs, Hyla versicolor, were collected in amplexus and repeatedly tested for phonotaxis behavior using a dynamic playback assay. To evaluate if hormonal mechanisms explained variation in the mate sampling, three steroid hormones, estradiol, progesterone, and corticosterone, were collected using a noninvasive water-borne hormone assay, validated for this species in the present study. Finally, we measured clutch size (investment) and the duration of time required for each female to oviposit after being reunited with their male mate (urgency). We found repeatability in many of the behaviors, including mate sampling. We found that females with higher concentrations estradiol and corticosterone made quicker choices, and that females with higher progesterone sampled mates more. We also found that female frogs in a more urgent reproductive state had lower concentrations of progesterone and estradiol, thereby providing the first evidence of a relationship between gonadal hormones and reproductive urgency. Collectively we found some support for the reproductive urgency but not the investment hypothesis. Thus, even though a female frog\u27s reproductive readiness is a highly transient life history stage, fine scale variation in her reproductive timeline could mitigate the strength of directional selection

Progesterone And Prostaglandin F2α Induce Species-Typical Female Preferences For Male Sexual Displays In Cope\u27s Gray Treefrog (Hyla Chrysoscelis)

Endocrine systems play critical roles in facilitating sexual behavior in seasonally breeding vertebrates. Much of the research exploring this topic has focused on the endocrine correlates of signaling behavior in males and sexual proceptivity in females. What is less understood is how hormones promote the expression of the often complex and highly selective set of stimulus–response behaviors that are observed in naturally breeding animals. In female frogs, phonotaxis is a robust and sensitive bioassay of mate choice and is exhibited by gravid females during the breeding season. In stark contrast, females exhibit low phonotactic responsiveness outside the breeding season, but the administration of hormones can induce sexual proceptivity. Here we test the hypothesis that manipulation of a minimal set of reproductive hormones—progesterone and prostaglandin F2α—are capable of evoking not only proceptive behavior in non-breeding females, but also the patterns of intraspecific selectivity for male sexual displays observed in gravid females tested during the breeding season. Specifically, we investigated whether preferences for faster call rates, longer call durations, and higher call efforts were similar between breeding and hormone-treated females of Cope\u27s gray treefrog (Hyla chrysoscelis). Hormone injections induced patterns of selective phonotaxis in non-breeding females that were remarkably similar to those observed in breeding females. These results suggest that there may be an important contribution of hormonal pleiotropy in regulating this complex, acoustically-guided sexual behavior. Our findings also support the idea that hormonal induction could be used to evaluate hypotheses about selective mate choice, and its underlying mechanisms, using non-breeding females

First Measurement of the Clustering Evolution of Photometrically-Classified Quasars

We present new measurements of the quasar autocorrelation from a sample of \~80,000 photometrically-classified quasars taken from SDSS DR1. We find a best-fit model of $\omega(\theta) = (0.066\pm^{0.026}_{0.024})\theta^{-(0.98\pm0.15)}$ for the angular autocorrelation, consistent with estimates from spectroscopic quasar surveys. We show that only models with little or no evolution in the clustering of quasars in comoving coordinates since z~1.4 can recover a scale-length consistent with local galaxies and Active Galactic Nuclei (AGNs). A model with little evolution of quasar clustering in comoving coordinates is best explained in the current cosmological paradigm by rapid evolution in quasar bias. We show that quasar biasing must have changed from b_Q~3 at a (photometric) redshift of z=2.2 to b_Q~1.2-1.3 by z=0.75. Such a rapid increase with redshift in biasing implies that quasars at z~2 cannot be the progenitors of modern L* objects, rather they must now reside in dense environments, such as clusters. Similarly, the duration of the UVX quasar phase must be short enough to explain why local UVX quasars reside in essentially unbiased structures. Our estimates of b_Q are in good agreement with recent spectroscopic results, which demonstrate the implied evolution in b_Q is consistent with quasars inhabiting halos of similar mass at every redshift. Treating quasar clustering as a function of both redshift and luminosity, we find no evidence for luminosity dependence in quasar clustering, and that redshift evolution thus affects quasar clustering more than changes in quasars' luminosity. We provide a new method for quantifying stellar contamination in photometrically-classified quasar catalogs via the correlation function.Comment: 34 pages, 10 figures, 1 table, Accepted to ApJ after: (i) Minor textual changes; (ii) extra points added to Fig.

Nineteen Years Of Consistently Positive And Strong Female Mate Preferences Despite Individual Variation

Sexual selection driven by mate choice has generated some of the most astounding diversity in nature, suggesting population-level preferences should be strong and consistent over many generations. On the other hand, mating preferences are among the least repeatable components of an individual animal’s phenotype, suggesting low consistency across an animal’s lifetime. Despite decades of intensive study of sexual selection there is almost no information about the strength and consistency of preferences across many years. In this study we present the results of over 5000 mate choice tests with a species of wild frog conducted over 19 consecutive years. Results show that preferences are positive and strong and vary little across years. This consistency is despite the fact that there are substantial differences among females in their strength of preference. We also suggest mate preferences in populations that are primarily the result of sensory exploitation might be more stable over time compared to preferences that are primarily involved in assessing male quality

How to distinguish starbursts and quiescently star-forming galaxies: The `bimodal' submillimetre galaxy population as a case study

In recent work (arXiv:1101.0002) we have suggested that the high-redshift (z ~ 2-4) bright submillimetre galaxy (SMG) population is heterogeneous, with major mergers contributing both at early stages, where quiescently star-forming discs are blended into one submm source (`galaxy-pair SMGs'), and late stages, where mutual tidal torques drive gas inflows and cause strong starbursts. Here we combine hydrodynamic simulations of major mergers with 3-D dust radiative transfer calculations to determine observational diagnostics that can distinguish between quiescently star-forming SMGs and starburst SMGs via integrated data alone. We fit the far-IR SEDs of the simulated galaxies with the optically thin single-temperature modified blackbody, the full form of the single-temperature modified blackbody, and a power-law temperature-distribution model. The effective dust temperature, T_dust, and power-law index of the dust emissivity in the far-IR, \beta, derived can significantly depend on the fitting form used, and the intrinsic \beta\ of the dust is not recovered. However, for all forms used here, there is a T_dust above which almost all simulated galaxies are starbursts, so a T_dust cut is very effective at selecting starbursts. Simulated merger-induced starbursts also have higher L_IR/M_gas and L_IR/L_FUV than quiescently star-forming galaxies and lie above the star formation rate-stellar mass relation. These diagnostics can be used to test our claim that the SMG population is heterogeneous and to observationally determine what star formation mode dominates a given galaxy population. We comment on applicability of these diagnostics to ULIRGs that would not be selected as SMGs. These `hot-dust ULIRGs' are typically starburst galaxies lower in mass than SMGs, but they can also simply be SMGs observed from a different viewing angle.Comment: 21 pages, 11 figures. Accepted for publication in MNRAS. Minor changes to text but otherwise identical to v

Present-day scaling relations of submm galaxies: origin of spheroidal sytems

We analyse the spectral energy distributions (SEDs) of 23 submm galaxies and 3 ISO-detected EROs, all of which have the spectroscopic redshifts, by using an evolutionary SED model of starbursts. This SED model allows us to investigate intrinsic properties of starbursts, such as the starburst age and the mean stellar metallicity, as it takes into account the chemical evolution. Also, the intrinsic size of the starburst region is estimated from observed SEDs. Using this SED model, we predict colour, magnitude, and size of present-day descendants of submm galaxies, and derive scaling relations, such as the present-day colour-magnitude and size-magnitude relations. We argue that submm galaxies are the progenitors of present-day elliptical galaxies, provided that the initial mass function (IMF) of submm galaxies is slightly flatter than the Salpeter IMF. In this case, we find that 1) the mean present-day magnitude of submm galaxies is similar to that of L* elliptical galaxies, 2) the present-day colour-magnitude relation is consistent with that of elliptical galaxies, 3) the present-day size-magnitude relation of elliptical galaxies can be reproduced if massive submm galaxies consist of multiple starburst regions. We estimate the effect of feedback in submm galaxies as a function of starburst age. It is found that starburst regions in submm galaxies are likely to be self-regulated; i.e. the effect of feedback is nearly balanced with the self-gravity of starburst regions.Comment: 27 pages, submitted to MNRA