Mating behavior and reproductive morphology predict macroevolution of sex allocation in hermaphroditic flatworms

Background:Sex allocation is the distribution of resources to male or female reproduction. In hermaphrodites, this concerns an individual’s resource allocation to, for example, the production of male or female gametes. Macroevolutionary studies across hermaphroditic plants have revealed that the self-pollination rate and the pollination mode are strong predictors of sex allocation. Consequently, we expect similar factors such as the selfing rate and aspects of the reproductive biology, like the mating behaviour and the intensity of postcopulatory sexual selection, to predict sex allocation in hermaphroditic animals. However, comparative work on hermaphroditic animals is limited. Here, we study sex allocation in 120 species of the hermaphroditic free-living flatworm genus Macrostomum. We ask how hypodermic insemination, a convergently evolved mating behaviour where sperm are traumatically injected through the partner’s epidermis, affects the evolution of sex allocation. We also test the commonly-made assumption that investment into male and female reproduction should trade-off. Finally, we ask if morphological indicators of the intensity of postcopulatory sexual selection (female genital complexity, male copulatory organ length, and sperm length) can predict sex allocation.Results:We find that the repeated evolution of hypodermic insemination predicts a more female-biased sex allocation (i.e., a relative shift towards female allocation). Moreover, transcriptome-based estimates of heterozygosity reveal reduced heterozygosity in hypodermically mating species, indicating that this mating behavior is linked to increased selfing or biparental inbreeding. Therefore, hypodermic insemination could represent a selfing syndrome. Furthermore, across the genus, allocation to male and female gametes is negatively related, and larger species have a more female-biased sex allocation. Finally, increased female genital complexity, longer sperm, and a longer male copulatory organ predict a more male-biased sex allocation.Conclusions:Selfing syndromes have repeatedly originated in plants. Remarkably, this macroevolutionary pattern is replicated in Macrostomum flatworms and linked to repeated shifts in reproductive behavior. We also find a trade-off between male and female reproduction, a fundamental assumption of most theories of sex allocation. Beyond that, no theory predicts a more female-biased allocation in larger species, suggesting avenues for future work. Finally, morphological indicators of more intense postcopulatory sexual selection appear to predict more intense sperm competition

Starburst99: Synthesis Models for Galaxies with Active Star Formation

Starburst99 is a comprehensive set of model predictions for spectrophotometric and related properties of galaxies with active star formation. The models are an improved and extended version of the data set previously published by Leitherer & Heckman (1995). We have upgraded our code by implementing the latest set of stellar evolution models of the Geneva group and the model atmosphere grid compiled by Lejeune et al. (1997). Several predictions which were not included in the previous publication are shown here for the first time. The models are presented in a homogeneous way for five metallicities between Z = 0.040 and 0.001 and three choices of the initial mass function. The age coverage is 10^6 to 10^9 yr. We also show the spectral energy distributions which are used to compute colors and other quantities. The full data set is available for retrieval at http://www.stsci.edu/science/starburst99/. This website allows users to run specific models with non-standard parameters as well. We also make the source code available to the community.Comment: 32 pages, LaTeX. All the Figures and the summary Table are located at http://www.stsci.edu/science/starburst99/, ApJ accepte

New Evolutionary Synthesis code. An application to the irregular galaxy NGC 1560

We have developed a new evolutionary synthesis code, which incorporates the output from chemical evolution models. We compare results of this new code with other published codes, and we apply it to the irregular galaxy NGC 1560 using sophisticated chemical evolution models. The code makes important contributions in two areas: a) the building of synthetic populations with time-dependent star formation rates and stellar populations of different metallicities; b) the extension of the set of stellar tracks from the Geneva group by adding the AGB phases for $m_i/M_\odot \geq 0.8$ as well as the very low mass stars. Our code predicts spectra, broad band colors, and Lick indices by using a spectra library, which cover a more complete grid of stellar parameters. The application of the code with the chemical models to the galaxy NGC 1560 constrain the star formation age for its stellar population around 10.0 Gy.Comment: 10 pages, 15 figures, submited to A&

The Massive Stellar Content in the Starburst NGC3049: A Test for Hot-Star Mode

We have obtained high-spatial resolution ultraviolet and optical STIS spectroscopy and imaging of the metal-rich nuclear starburst in NGC3049. The stellar continuum and the absorption line spectrum in the ultraviolet are used to constrain the massive stellar population. The strong, blueshifted stellar lines of CIV and SiIV detected in the UV spectra indicate a metal-rich, compact, massive (1E6 Msol) cluster of age 3--4 Myr emitting the UV-optical continuum. We find strong evidence against a depletion of massive stars in this metal-rich cluster. The derived age and the upper mass-limit cut-off of the initial mass function are also consistent with the detection of Wolf-Rayet (WR) features at optical wavelengths. As a second independentconstraint on the massive stellar content, the nebular emission-line spectrum is modeled with photoionization codes using stellar spectra from evolutionary synthesis models. However, the nebular lines are badly reproduced by 3--4 Myr instantaneous bursts, as required by the UV line spectrum, when unblanketed WR and/or Kurucz stellar atmospheres are used. The corresponding number of photons above 24 and 54 eV in the synthetic models is too high in comparison with values suggested by the observed line ratios. Since the ionizing spectrum in this regime is dominated by emission from WR stars, this discrepancy between observations and models is most likely the result of incorrect assumptions about the WR stars. Thus we conclude that the nebular spectrum of high-metallicity starbursts is poorly reproduced by models for WR dominated populations. However, the new model set of Smith et al. (2002) with blanketed WR and O atmospheres and adjusted WR temperatures predicts a softer far-UV radiation field, providing a better match to the data.Comment: To be published in ApJ, Dec. issue 17 figures, 3 in gif forma

HST's view of the youngest massive stars in the Magellanic Clouds

Accurate physical parameters of newborn massive stars are essential ingredients to shed light on their formation, which is still an unsolved problem. The rare class of compact H II regions in the Magellanic Clouds (MCs), termed ``high-excitation blobs'' (HEBs), presents a unique opportunity to acquire this information. These objects (~ 4" to 10", ~ 1 to 3 pc, in diameter) harbor the youngest massive stars of the OB association/molecular cloud complexes in the MCs accessible through high-resolution near-IR and optical techniques. We present a brief overview of the results obtained with HST mainly on two HEBs, one in the LMC (N159-5) and the other in the SMC (N81).Comment: 5 pages, to appear in the Proceedings of the 41st ESLAB Symposium "The Impact of HST on European Astronomy", 29 May to 1 June 2007, ESTEC, Noordwijk, Netherlands; eds. Guido De Marchi and Duccio Macchett

New Models for Wolf-Rayet and O Star Populations in Young Starbursts

Using the latest stellar evolution models, theoretical stellar spectra, and a compilation of observed emission line strengths from Wolf-Rayet (WR) stars, we construct evolutionary synthesis models for young starbursts. We explicitly distinguish between the various WR subtypes (WN, WC, WO), and we treat O and Of stars separately. We provide detailed predictions of UV and optical emission line strengths for both the WR stellar lines and the major nebular hydrogen and helium emission lines, as a function of several input parameters related to the starburst episode. We also derive the theoretical frequency of WR-rich starbursts. We then discuss: nebular HeII 4686 emission, the contribution of WR stars to broad Balmer line emission, techniques used to derive the WR and O star content from integrated spectra, and explore the implications of the formation of WR stars through mass transfer in close binary systems in instantaneous bursts. The observational features predicted by our models allow a detailed quantitative determination of the massive star population in a starburst region (particularly in so-called "WR galaxies") from its integrated spectrum and provide a means of deriving the burst properties (e.g., duration, age) and the parameters of the initial mass function of young starbursts. (Abridged abstract)Comment: Accepted by ApJ Supplements. LaTeX using aasmp4, psfigs macros. 49 pages including 23 figures. Paper (full, or text/figures separated) and detailed model results available at http://www.stsci.edu/ftp/science/starburst/sv97.htm

Revisiting the Modified Eddington Limit for Massive Stars

We have determined the location of the line-opacity modified Eddington limit for stars in the LMC using the most recent atmosphere models combined with a precise mapping to the HR Diagram through up-to-date stellar evolution calculations. While we find, in agreement with previous studies, that the shape of the modified Eddington limit qualitatively corresponds to the Humphreys-Davidson (HD) limit defined by the most luminous supergiants, the modified limit is actually a full magnitude higher than the upper luminosity limit observed for LMC stars. The observed limit is consistent with atmosphere models in which the maximum value of the ratio of the radiation force outwards to the gravitational force inwards, Y_max, is 0.9, i.e., the photospheres of stars at the observed luminosity limit are bound. As massive stars evolve, they move to higher, and therefore less stable values of Y_max, so mass loss, either sporadic or continuous, may halt their natural redward evolution as they approach the Y_max = 0.9 limit. We assess the metallicity dependence of this limit. If the limit does determine the most luminous stars, and the value of Y_max corresponding to the luminosity limit in the LMC is universal, then the brightest supergiants the SMC should be only marginally brighter (0.3 mag) than those of the LMC, in agreement with observations. Moreover, the brightest supergiants in M31 should be 0.75~mag fainter than those in the LMC.Comment: 13 pages with 4 figures, AAS Latex, ApJ Submitted (August

The Stanley Foundation Bipolar Network: Results of the naturalistic follow-up study after 2.5 years of follow-up in the German centres

The Stanley Foundation Bipolar Network (SFBN) is an international, multisite network investigating the characteristics and course of bipolar disorder. Methods (history, ratings and longitudinal follow-up) are standardized and equally applied in all 7 centres. This article describes demographics and illness characteristics of the first 152 German patients enrolled in them SFBN as well as the results of 2.5 years of follow-up. Patients in Germany were usually enrolled after hospitalisation. More than 72% of the study population suffered from bipolar I disorder and 25% from bipolar 11 disorder. The mean +/- SD age of the study participants was 42.08 +/- 13.5 years, and the mean SD age of onset 24.44 +/- 10.9 years. More than 40% of the sample reported a rapid-cycling course in history, and even more a cycle acceleration overtime. 37% attempted suicide at least once. 36% had an additional Axis I disorder, with alcohol abuse being the most common one, followed by anxiety disorders. During the follow-up period, only 27% remained stable, 56% had a recurrence, 12.8% perceived subsyndromal symptoms despite treatment and regular visits. 27% suffered from a rapid-cycling course during the follow-up period. Recurrences were significantly associated with bipolar I disorder, an additional comorbid Axis I disorder, rapid cycling in history, a higher number of mood stabilizers and the long-term use of typical antipsychotics. Rapid cycling during follow-up was only associated with a rapidcycling course in history, a higher number of mood stabilizers and at least one suicide attempt in history. Copyright (c) 2003 S. Karger AG, Basel

A Spectroscopic Search for the non-nuclear Wolf-Rayet Population of the metal-rich spiral galaxy M83

We present a catalogue of non-nuclear regions containing Wolf-Rayet stars in the metal-rich spiral galaxy M83 (NGC5236). From a total of 283 candidate regions identified using HeII 4686 imaging with VLT-FORS2, Multi Object Spectroscopy of 198 regions was carried out, confirming 132 WR sources. From this sub-sample, an exceptional content of 1035 +/- 300 WR stars is inferred, with N(WC)/N(WN) approx 1.2, continuing the trend to larger values at higher metallicity amongst Local Group galaxies, and greatly exceeding current evolutionary predictions at high metallicity. Late-type stars dominate the WC population of M83, with N(WC8-9)/N(WC4-7)=9 and WO subtypes absent, consistent with metallicity dependent WC winds. Equal numbers of late to early WN stars are observed, again in contrast to current evolutionary predictions. Several sources contain large numbers of WR stars. In particular, #74 (alias region 35 from De Vaucouleurs et al. contains 230 WR stars, and is identified as a Super Star Cluster from inspection of archival HST/ACS images. Omitting this starburst cluster would result in revised statistics of N(WC)/N(WN) approx 1 and N(WC8-9)/N(WC4-7) approx 6 for the `quiescent' disk population. Including recent results for the nucleus and accounting for incompleteness in our spectroscopic sample, we suspect the total WR population of M83 may exceed 3000 stars.Comment: 39 pages, 13 figures, 17 finding charts, accepted for Astronomy & Astrophysics. Version will full resolution images available at ftp://astro1.shef.ac.uk/pub/pac/m83.ps.g

The First Stars

The formation of the first generations of stars at redshifts z > 15-20 signaled the transition from the simple initial state of the universe to one of increasing complexity. We here review recent progress in understanding the assembly process of the first galaxies, starting with cosmological initial conditions and modelling the detailed physics of star formation. In particular, we study the role of HD cooling in ionized primordial gas, the impact of UV radiation produced by the first stars, and the propagation of the supernova blast waves triggered at the end of their brief lives. We conclude by discussing how the chemical abundance patterns observed in extremely low-metallicity stars allow us to probe the properties of the first stars.Comment: 12 pages, 9 figures, proceedings of the IAU Symposium 250 "Massive stars as cosmic engines