Phenotypic plasticity as a clue for the invasion success of the submerged aquatic plant Elodea nuttallii

Two closely related alien submerged aquatic plants were introduced into Europe. The new invader (Elodea nuttallii) gradually displaced E. canadensis even at sites where the latter was well established. The aim of the study was to evaluate the combined effects of environmental factors on several phenotypic characteristics of the two Elodea species, and to relate these phenotypic characteristics to the invasion success of E. nuttallii over E. canadensis. In a factorial design, Elodea plants were grown in aquaria containing five different nitrogen concentrations and incubated at five different light intensities. We used six functional traits (apical shoot RGR), total shoot RGR, relative elongation, root length, lateral spread, branching degree) to measure the environmental response of the species. We calculated plasticity indices to express the phenotypic differences between species. Light and nitrogen jointly triggered the development of phenotypic characteristics that make E. nuttallii a more successful invader in eutrophic waters than E. canadensis. The stronger invader showed a wider range of phenotypic plasticity. The apical elongation was the main difference between the two species, with E. nuttallii being more than two times longer than E. canadensis. E. canadensis formed dense side shoots even under high shade and low nitrogen levels, whereas E. nuttallii required higher light and nitrogen levels. We found that under more eutrophic conditions, E. nuttallii reach the water surface sooner than E. canadensis and through intensive branching outcompetes all other plants including E. canadensis. Our findings support the theory that more successful invaders have wider phenotypic plasticit

The Physics of Crystallization from Globular Cluster White Dwarf Stars in NGC 6397

We explore the physics of crystallization in the deep interiors of white dwarf stars using the color-magnitude diagram and luminosity function constructed from proper motion cleaned Hubble Space Telescope photometry of the globular cluster NGC 6397. We demonstrate that the data are consistent with the theory of crystallization of the ions in the interior of white dwarf stars and provide the first empirical evidence that the phase transition is first order: latent heat is released in the process of crystallization as predicted by van Horn (1968). We outline how this data can be used to observationally constrain the value of Gamma = E_{Coulomb}/E_{thermal} near the onset of crystallization, the central carbon/oxygen abundance, and the importance of phase separation.Comment: 5 pages, 5 figures, accepted for publication in the Astrophysical Journal Letter

New phase diagrams for dense carbon-oxygen mixtures and white dwarf evolution

Cool white dwarfs are reliable and independent stellar chronometers. The most common white dwarfs have carbon-oxygen dense cores. Consequently, the cooling ages of very cool white dwarfs sensitively depend on the adopted phase diagram of the carbon-oxygen binary mixture. A new phase diagram of dense carbon-oxygen mixtures appropriate for white dwarf interiors has been recently obtained using direct molecular dynamics simulations. In this paper, we explore the consequences of this phase diagram in the evolution of cool white dwarfs. To do this we employ a detailed stellar evolutionary code and accurate initial white dwarf configurations, derived from the full evolution of progenitor stars. We use two different phase diagrams, that of Horowitz et al. (2010), which presents an azeotrope, and the phase diagram of Segretain & Chabrier (1993), which is of the spindle form. We computed the evolution of 0.593 and 0.878M_sun white dwarf models during the crystallization phase, and we found that the energy released by carbon-oxygen phase separation is smaller when the new phase diagram of Horowitz et al. (2010) is used. This translates into time delays that are on average a factor about 2 smaller than those obtained when the phase diagram of Segretain & Chabrier (1993) is employed. Our results have important implications for white dwarf cosmochronology, because the cooling ages of very old white dwarfs are different for the two phase diagrams. This may have a noticeable impact on the age determinations of very old globular clusters, for which the white dwarf color-magnitude diagram provides an independent way of estimating their age.Comment: 7 pages, 7 figures, accepted for publication in Astronomy and Astrophysic

Evolutionary calculations of phase separation in crystallizing white dwarf stars

We present an exploration of the significance of Carbon/Oxygen phase separation in white dwarf stars in the context of self-consistent evolutionary calculations. Because phase separation can potentially increase the calculated ages of the oldest white dwarfs, it can affect the age of the Galactic disk as derived from the downturn in the white dwarf luminosity function. We find that the largest possible increase in ages due to phase separation is 1.5 Gyr, with a most likely value of approximately 0.6 Gyr, depending on the parameters of our white dwarf models. The most important factors influencing the size of this delay are the total stellar mass, the initial composition profile, and the phase diagram assumed for crystallization. We find a maximum age delay in models with masses of 0.6 solar masses, which is near the peak in the observed white dwarf mass distribution. We find that varying the opacities (via the metallicity) has little effect on the calculated age delays. In the context of Galactic evolution, age estimates for the oldest Galactic globular clusters range from 11.5 to 16 Gyr, and depend on a variety of parameters. In addition, a 4 to 6 Gyr delay is expected between the formation of the globular clusters and that of the Galactic thin disk, while the observed white dwarf luminosity function gives an age estimate for the thin disk of 9.5 +/-1.0 Gyr, without including the effect of phase separation. Using the above numbers, we see that phase separation could add between 0 to 3 Gyr to the white dwarf ages and still be consistent with the overall picture of Galaxy formation. Our calculated maximum value of 1.5 Gyr fits within these bounds, as does our best guess value of 0.6 Gyr.Comment: 13 total pages, 8 figures, 3 tables, accepted for publication in the Astrophysical Journal on May 25, 199

Above- and below-ground vertebrate herbivory may each favour a different subordinate species in an aquatic plant community

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species

Experimental Assessment of the Water Quality Influence on the Phosphorus Uptake of an Invasive Aquatic Plant: Biological Responses throughout Its Phenological Stage

International audienceUnderstanding how an invasive plant can colonize a large range of environments is still a great challenge in freshwater ecology. For the first time, we assessed the relative importance of four factors on the phosphorus uptake and growth of an invasive macrophyte Elodea nuttallii (Planch.) St. John. This study provided data on its phenotypic plasticity, which is frequently suggested as an important mechanism but remains poorly investigated. The phosphorus uptake of two Elodea nuttallii subpopulations was experimentally studied under contrasting environmental conditions. Plants were sampled in the Rhine floodplain and in the Northern Vosges mountains, and then maintained in aquaria in hard (Rhine) or soft (Vosges) water. Under these conditions, we tested the influence of two trophic states (eutrophic state, 100 mu g.l(-1) P-PO43- and hypertrophic state, 300 mu g.l(-1) P-PO43-) on the P metabolism of plant subpopulations collected at three seasons (winter, spring and summer). Elodea nuttallii was able to absorb high levels of phosphorus through its shoots and enhance its phosphorus uptake, continually, after an increase of the resource availability (hypertrophic > eutrophic). The lowest efficiency in nutrient use was observed in winter, whereas the highest was recorded in spring, what revealed thus a storage strategy which can be beneficial to new shoots. This experiment provided evidence that generally, the water trophic state is the main factor governing P uptake, and the mineral status (softwater > hardwater) of the stream water is the second main factor. The phenological stage appeared to be a confounding factor to P level in water. Nonetheless, phenology played a role in P turnover in the plant. Finally, phenotypic plasticity allows both subpopulations to adapt to a changing environment

Water velocity limits the temporal extent of herbivore effects on aquatic plants in a lowland river

The role of herbivores in regulating aquatic plant dynamics has received growing recognition from researchers and managers. However, the evidence for herbivore impacts on aquatic plants is largely based on short-term exclosure studies conducted within a single plant growing season. Thus, it is unclear how long herbivore impacts on aquatic plant abundance can persist for. We addressed this knowledge gap by testing whether mute swan (Cygnus olor) grazing on lowland river macrophytes could be detected in the following growing season. Furthermore, we investigated the role of seasonal changes in water current speed in limiting the temporal extent of grazing. We found no relationship between swan biomass density in 1 year and aquatic plant cover or biomass in the following spring. No such carry-over effects were detected despite observing high swan biomass densities in the previous year from which we inferred grazing impacts on macrophytes. Seasonal increases in water velocity were associated with reduced grazing pressure as swans abandoned river habitat. Furthermore, our study highlights the role of seasonal changes in water velocity in determining the length of the mute swan grazing season in shallow lowland rivers and thus in limiting the temporal extent of herbivore impacts on aquatic plant abundance