Evolution of inclined planets in three-dimensional radiative discs

While planets in the solar system only have a low inclination with respect to the ecliptic there is mounting evidence that in extrasolar systems the inclination can be very high, at least for close-in planets. One process to alter the inclination of a planet is through planet-disc interactions. Recent simulations considering radiative transport have shown that the evolution of migration and eccentricity can strongly depend on the thermodynamic state of the disc. We extend previous studies to investigate the planet-disc interactions of fixed and moving planets on inclined and eccentric orbits. We also analyse the effect of the disc's thermodynamic properties on the orbital evolution of embedded planets in detail. The protoplanetary disc is modelled as a viscous gas where the internally produced dissipation is transported by radiation. For locally isothermal discs, we confirm previous results and find inclination damping and inward migration for planetary cores. For low inclinations i < 2 H/r, the damping is exponential, while di/dt is proportional to i^-2 for larger i. For radiative discs, the planetary migration is very limited, as long as their inclination exceeds a certain threshold. If the inclination is damped below this threshold, planetary cores with a mass up to approximately 33 Earth masses start to migrate outwards, while larger cores migrate inwards right from the start. The inclination is damped for all analysed planet masses. In a viscous disc an initial inclination of embedded planets will be damped for all planet masses. This damping occurs on timescales that are shorter than the migration time. If the inclination lies beneath a certain threshold, the outward migration in radiative discs is not handicapped. Outward migration is strongest for circular and non-inclined orbits

Range of outward migration and influence of the disc's mass on the migration of giant planet cores

The migration of planets plays an important role in the early planet-formation process. An important problem has been that standard migration theories predict very rapid inward migration, which poses problems for population synthesis models. However, it has been shown recently that low-mass planets (20-30 Earth Masses) that are still embedded in the protoplanetary disc can migrate outwards under certain conditions. Simulations have been performed mostly for planets at given radii for a particular disc model. Here, we plan to extend previous work and consider different masses of the disc to quantify the influence of the physical disc conditions on planetary migration. We perform three-dimensional (3D) radiation hydrodynamical simulations of embedded planets in protoplanteary discs. For planets on circular orbits at various locations we measure the radial dependece of the torques. For all considered planet masses (20-30 Earth masses) in this study we find outward migration within a limited radial range of the disc, typically from about 0.5 up to 1.5-2.5 a_Jup. Inside and outside this intervall, migration is inward and given by the Lindblad value for large radii. Because outward migration stops at a certain location in the disc, there exists a zero-torque distance for planetary embryos, where they can continue to grow without moving too fast. For higher disc masses (M_disc > 0.02 M_Sol) convection ensues, which changes the structure of the disc and therefore the torque on the planet as well. Outward migration stops at different points in the disc for different planetary masses, resulting in a quite extended region where the formation of larger cores might be easier. In higher mass discs, convection changes the disc's structure resulting in fluctuations in the surface density, which influence the torque acting on the planet, and therefore its migration rate

Planetesimal Formation In Self-Gravitating Discs

We study particle dynamics in local two-dimensional simulations of self-gravitating accretion discs with a simple cooling law. It is well known that the structure which arises in the gaseous component of the disc due to a gravitational instability can have a significant effect on the evolution of dust particles. Previous results using global simulations indicate that spiral density waves are highly efficient at collecting dust particles, creating significant local over-densities which may be able to undergo gravitational collapse. We expand on these findings, using a range of cooling times to mimic the conditions at a large range of radii within the disc. Here we use the Pencil Code to solve the 2D local shearing sheet equations for gas on a fixed grid together with the equations of motion for solids coupled to the gas solely through aerodynamic drag force. We find that spiral density waves can create significant enhancements in the surface density of solids, equivalent to 1-10cm sized particles in a disc following the profiles of Clarke (2009) around a solar mass star, causing it to reach concentrations several orders of magnitude larger than the particles mean surface density. We also study the velocity dispersion of the particles, finding that the spiral structure can result in the particle velocities becoming highly ordered, having a narrow velocity dispersion. This implies low relative velocities between particles, which in turn suggests that collisions are typically low energy, lessening the likelihood of grain destruction. Both these findings suggest that the density waves that arise due to gravitational instabilities in the early stages of star formation provide excellent sites for the formation of large, planetesimal-sized objects.Comment: 11 pages, 8 figures, accepted for publication in MNRA

Instability of Plastid DNA in the Nuclear Genome

Functional gene transfer from the plastid (chloroplast) and mitochondrial genomes to the nucleus has been an important driving force in eukaryotic evolution. Non-functional DNA transfer is far more frequent, and the frequency of such transfers from the plastid to the nucleus has been determined experimentally in tobacco using transplastomic lines containing, in their plastid genome, a kanamycin resistance gene (neo) readymade for nuclear expression. Contrary to expectations, non-Mendelian segregation of the kanamycin resistance phenotype is seen in progeny of some lines in which neo has been transferred to the nuclear genome. Here, we provide a detailed analysis of the instability of kanamycin resistance in nine of these lines, and we show that it is due to deletion of neo. Four lines showed instability with variation between progeny derived from different areas of the same plant, suggesting a loss of neo during somatic cell division. One line showed a consistent reduction in the proportion of kanamycin-resistant progeny, suggesting a loss of neo during meiosis, and the remaining four lines were relatively stable. To avoid genomic enlargement, the high frequency of plastid DNA integration into the nuclear genome necessitates a counterbalancing removal process. This is the first demonstration of such loss involving a high proportion of recent nuclear integrants. We propose that insertion, deletion, and rearrangement of plastid sequences in the nuclear genome are important evolutionary processes in the generation of novel nuclear genes. This work is also relevant in the context of transgenic plant research and crop production, because similar processes to those described here may be involved in the loss of plant transgenes

Recent developments in planet migration theory

Planetary migration is the process by which a forming planet undergoes a drift of its semi-major axis caused by the tidal interaction with its parent protoplanetary disc. One of the key quantities to assess the migration of embedded planets is the tidal torque between the disc and planet, which has two components: the Lindblad torque and the corotation torque. We review the latest results on both torque components for planets on circular orbits, with a special emphasis on the various processes that give rise to additional, large components of the corotation torque, and those contributing to the saturation of this torque. These additional components of the corotation torque could help address the shortcomings that have recently been exposed by models of planet population syntheses. We also review recent results concerning the migration of giant planets that carve gaps in the disc (type II migration) and the migration of sub-giant planets that open partial gaps in massive discs (type III migration).Comment: 52 pages, 18 figures. Review article to be published in "Tidal effects in Astronomy and Astrophysics", Lecture Notes in Physic

Monitoring Voltage-Dependent Charge Displacement of Shaker B-IR K+ Ion Channels Using Radio Frequency Interrogation

Here we introduce a new technique that probes voltage-dependent charge displacements of excitable membrane-bound proteins using extracellularly applied radio frequency (RF, 500 kHz) electric fields. Xenopus oocytes were used as a model cell for these experiments, and were injected with cRNA encoding Shaker B-IR (ShB-IR) K+ ion channels to express large densities of this protein in the oocyte membranes. Two-electrode voltage clamp (TEVC) was applied to command whole-cell membrane potential and to measure channel-dependent membrane currents. Simultaneously, RF electric fields were applied to perturb the membrane potential about the TEVC level and to measure voltage-dependent RF displacement currents. ShB-IR expressing oocytes showed significantly larger changes in RF displacement currents upon membrane depolarization than control oocytes. Voltage-dependent changes in RF displacement currents further increased in ShB-IR expressing oocytes after ∼120 µM Cu2+ addition to the external bath. Cu2+ is known to bind to the ShB-IR ion channel and inhibit Shaker K+ conductance, indicating that changes in the RF displacement current reported here were associated with RF vibration of the Cu2+-linked mobile domain of the ShB-IR protein. Results demonstrate the use of extracellular RF electrodes to interrogate voltage-dependent movement of charged mobile protein domains — capabilities that might enable detection of small changes in charge distribution associated with integral membrane protein conformation and/or drug–protein interactions

Tracking Cats: Problems with Placing Feline Carnivores on δ18O, δD Isoscapes

Several felids are endangered and threatened by the illegal wildlife trade. Establishing geographic origin of tissues of endangered species is thus crucial for wildlife crime investigations and effective conservation strategies. As shown in other species, stable isotope analysis of hydrogen and oxygen in hair (δD(h), δ(18)O(h)) can be used as a tool for provenance determination. However, reliably predicting the spatial distribution of δD(h) and δ(18)O(h) requires confirmation from animal tissues of known origin and a detailed understanding of the isotopic routing of dietary nutrients into felid hair.We used coupled δD(h) and δ(18)O(h) measurements from the North American bobcat (Lynx rufus) and puma (Puma concolor) with precipitation-based assignment isoscapes to test the feasibility of isotopic geo-location of felidae. Hairs of felid and rabbit museum specimens from 75 sites across the United States and Canada were analyzed. Bobcat and puma lacked a significant correlation between H/O isotopes in hair and local waters, and also exhibited an isotopic decoupling of δ(18)O(h) and δD(h). Conversely, strong δD and δ(18)O coupling was found for key prey, eastern cottontail rabbit (Sylvilagus floridanus; hair) and white-tailed deer (Odocoileus virginianus; collagen, bone phosphate).Puma and bobcat hairs do not adhere to expected pattern of H and O isotopic variation predicted by precipitation isoscapes for North America. Thus, using bulk hair, felids cannot be placed on δ(18)O and δD isoscapes for use in forensic investigations. The effective application of isotopes to trace the provenance of feline carnivores is likely compromised by major controls of their diet, physiology and metabolism on hair δ(18)O and δD related to body water budgets. Controlled feeding experiments, combined with single amino acid isotope analysis of diets and hair, are needed to reveal mechanisms and physiological traits explaining why felid hair does not follow isotopic patterns demonstrated in many other taxa

Giant Planet Formation and Migration

© 2018, The Author(s). Planets form in circumstellar discs around young stars. Starting with sub-micron sized dust particles, giant planet formation is all about growing 14 orders of magnitude in size. It has become increasingly clear over the past decades that during all stages of giant planet formation, the building blocks are extremely mobile and can change their semimajor axis by substantial amounts. In this chapter, we aim to give a basic overview of the physical processes thought to govern giant planet formation and migration, and to highlight possible links to water delivery.S.-J. Paardekooper is supported by a Royal Society University Research Fellowship. A. Johansen is supported by the Knut and Alice Wallenberg Foundation, the Swedish Research Council (grant 2014-5775) and the European Research Council (ERC Starting Grant 278675-PEBBLE2PLANET)

Location, identity, amount and serial entry of chloroplast DNA sequences in crucifer mitochondrial DNAs

Southern blot hybridization techniques were used to examine the chloroplast DNA (cpDNA) sequences present in the mitochondrial DNAs (mtDNAs) of two Brassica species ( B. campestris and B. hirta ), two closely related species belonging to the same tribe as Brassica (Raphanus sativa, Crambe abyssinica) , and two more distantly related species of crucifers (Arabidopsis thaliana, Capsella bursa-pastoris) . The two Brassica species and R. sativa contain roughly equal amounts (12–14 kb) of cpDNA sequences integrated within their 208–242 kb mtDNAs. Furthermore, the 11 identified regions of transferred DNA, which include the 5′ end of the chloroplast psa A gene and the central segment of rpo B, have the same mtDNA locations in these three species. Crambe abyssinica mtDNA has the same complement of cpDNA sequences, plus an additional major region of cpDNA sequence similarity which includes the 16S rRNA gene. Therefore, except for the more recently arrived 16S rRNA gene, all of these cpDNA sequences appear to have entered the mitochondrial genome in the common ancestor of these three genera. The mitochondrial genomes of A. thaliana and Capsella bursa-pastoris contain significantly less cpDNA (5–7 kb) than the four other mtDNAs. However, certain cpDNA sequences, including the central portion of the rbc L gene and the 3′ end of the psa A gene, are shared by all six crucifer mtDNAs and appear to have been transferred in a common ancestor of the crucifer family over 30 million years ago. 1n conclusion, DNA has been transferred sequentially from the chloroplast to the mitochondrion during crucifer evolution and these cpDNA sequences can persist in the mitochondrial genome over long periods of evolutionary time.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/46968/1/294_2004_Article_BF00521276.pd

The evolution of the plastid chromosome in land plants: gene content, gene order, gene function

This review bridges functional and evolutionary aspects of plastid chromosome architecture in land plants and their putative ancestors. We provide an overview on the structure and composition of the plastid genome of land plants as well as the functions of its genes in an explicit phylogenetic and evolutionary context. We will discuss the architecture of land plant plastid chromosomes, including gene content and synteny across land plants. Moreover, we will explore the functions and roles of plastid encoded genes in metabolism and their evolutionary importance regarding gene retention and conservation. We suggest that the slow mode at which the plastome typically evolves is likely to be influenced by a combination of different molecular mechanisms. These include the organization of plastid genes in operons, the usually uniparental mode of plastid inheritance, the activity of highly effective repair mechanisms as well as the rarity of plastid fusion. Nevertheless, structurally rearranged plastomes can be found in several unrelated lineages (e.g. ferns, Pinaceae, multiple angiosperm families). Rearrangements and gene losses seem to correlate with an unusual mode of plastid transmission, abundance of repeats, or a heterotrophic lifestyle (parasites or myco-heterotrophs). While only a few functional gene gains and more frequent gene losses have been inferred for land plants, the plastid Ndh complex is one example of multiple independent gene losses and will be discussed in detail. Patterns of ndh-gene loss and functional analyses indicate that these losses are usually found in plant groups with a certain degree of heterotrophy, might rendering plastid encoded Ndh1 subunits dispensable