The genetic architecture of colonization resistance in Brachypodium distachyon to non-adapted stripe rust (Puccinia striiformis) isolates

Multilayered defense responses ensure that plants are hosts to only a few adapted pathogens in the environment. The host range of a plant pathogen depends on its ability to fully overcome plant defense barriers, with failure at any single step sufficient to prevent life cycle completion of the pathogen. Puccinia striiformis, the causal agent of stripe rust (=yellow rust), is an agronomically important obligate biotrophic fungal pathogen of wheat and barley. It is generally unable to complete its life cycle on the non-adapted wild grass species Brachypodium distachyon, but natural variation exists for the degree of hyphal colonization by Puccinia striiformis. Using three B. distachyon mapping populations, we identified genetic loci conferring colonization resistance to wheat-adapted and barley-adapted isolates of P. striiformis. We observed a genetic architecture composed of two major effect QTLs (Yrr1 and Yrr3) restricting the colonization of P. striiformis. Isolate specificity was observed for Yrr1, whereas Yrr3 was effective against all tested P. striiformis isolates. Plant immune receptors of the nucleotide binding, leucine-rich repeat (NB-LRR) encoding gene family are present at the Yrr3 locus, whereas genes of this family were not identified at the Yrr1 locus. While it has been proposed that resistance to adapted and non-adapted pathogens are inherently different, the observation of (1) a simple genetic architecture of colonization resistance, (2) isolate specificity of major and minor effect QTLs, and (3) NB-LRR encoding genes at the Yrr3 locus suggest that factors associated with resistance to adapted pathogens are also critical for non-adapted pathogens

The collapse of protoplanetary clumps formed through disc instability: 3D simulations of the pre-dissociation phase

We present 3D smoothed particle hydrodynamics simulations of the collapse of clumps formed through gravitational instability in the outer part of a protoplanetary disc. The initial conditions are taken directly from a global disc simulation, and a realistic equation of state is used to follow the clumps as they contract over several orders of magnitude in density, approaching the molecular hydrogen dissociation stage. The effects of clump rotation, asymmetries, and radiative cooling are studied. Rotation provides support against fast collapse, but non-axisymmetric modes develop and efficiently transport angular momentum outward, forming a circumplanetary disc. This transport helps the clump reach the dynamical collapse phase, resulting from molecular hydrogen dissociation, on a thousand-year timescale, which is smaller than timescales predicted by some previous spherical 1D collapse models. Extrapolation to the threshold of the runaway hydrogen dissociation indicates that the collapse timescales can be shorter than inward migration timescales, suggesting that clumps could survive tidal disruption and deliver a proto-gas giant to distances of even a few AU from the central star.Comment: Accepted for publication in MNRA

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

The diverse lives of massive protoplanets in self-gravitating discs

Gas giant planets may form early-on during the evolution of protostellar discs, while these are relatively massive. We study how Jupiter-mass planet-seeds (termed protoplanets) evolve in massive, but gravitationally stable (Q ∼> 1.5), discs using radiative hydrodynamic simulations. We find that the protoplanet initially migrates inwards rapidly, until it opens up a gap in the disc. Thereafter, it either continues to migrate inwards on a much longer timescale or starts migrating outwards. Outward migration occurs when the protoplanet resides within a gap with gravitationally unstable edges, as a high fraction of the accreted gas is high angular momentum gas from outside the protoplanet’s orbit. The effect of radiative heating from the protoplanet is critical in determining the direction of the migration and the eccentricity of the protoplanet. Gap opening is facilitated by efficient cooling that may not be captured by the commonly used β-cooling approximation. The protoplanet initially accretes at a high rate (∼ 10^(−3)Mj yr−1), and its accretion luminosity could be a few tenths of the host star’s luminosity, making the protoplanet easily observable (albeit only for a short time). Due to the high gas accretion rate, the protoplanet generally grows above the deuterium-burning mass-limit. Protoplanet radiative feedback reduces its mass growth so that its final mass is near the brown dwarf-planet boundary. The fate of a young planet-seed is diverse and could vary from a gas giant planet on a circular orbit at a few AU from the central star to a brown dwarf on an eccentric, wide orbit

Components of Brachypodium distachyon resistance to nonadapted wheat stripe rust pathogens are simply inherited

Phytopathogens have a limited range of host plant species that they can successfully parasitise ie. that they are adapted for. Infection of plants by nonadapted pathogens often results in an active resistance response that is relatively poorly characterised because phenotypic variation in this response often does not exist within a plant species, or is too subtle for genetic dissection. In addition, complex polygenic inheritance often underlies these resistance phenotypes and mutagenesis often does not impact upon this resistance, presumably due to genetic or mechanistic redundancy. Here it is demonstrated that phenotypic differences in the resistance response of Brachypodium distachyon to the nonadapted wheat stripe rust pathogen Puccinia striiformis f. sp. tritici (Pst) are genetically tractable and simply inherited. Two dominant loci were identified on B. distachyon chromosome 4 that each reduce attempted Pst colonisation compared with sib and parent lines without these loci. One locus (Yrr1) is effective against diverse Australian Pst isolates and present in two B. distachyon mapping families as a conserved region that was reduced to 5 candidate genes by fine mapping. A second locus, Yrr2, shows Pst race-specificity and encodes a disease resistance gene family typically associated with host plant resistance. These data indicate that some components of resistance to nonadapted pathogens are genetically tractable in some instances and may mechanistically overlap with host plant resistance to avirulent adapted pathogens

Outward migration of a giant planet with a gravitationally unstable gap edge

We present numerical simulations of disc-planet interactions where the planet opens a gravitationally unstable gap in an otherwise gravitationally stable disc. In our disc models, where the outer gap edge can be unstable to global spiral modes, we find that as we increase the surface density scale the gap becomes more unstable and the planet migrates outwards more rapidly. We show that the positive torque is provided by material brought into the planet's coorbital region by the spiral arms. This material is expected to execute horseshoe turns upon approaching the planet and hence torque it. Our results suggest that standard type II migration, applicable to giant planets in non-self-gravitating viscous discs, is likely to be significantly modified in massive discs when gravitational instabilities associated with the gap occur.Comment: 9 pages, 11 figures, accepted by MNRA

The wheat Sr22, Sr33, Sr35 and Sr45 genes confer resistance against stem rust in barley

In the last 20 years, stem rust caused by the fungus Puccinia graminis f. sp. tritici (Pgt), has re-emerged as a major threat to wheat and barley production in Africa and Europe. In contrast to wheat with 60 designated stem rust (Sr) resistance genes, barley’s genetic variation for stem rust resistance is very narrow with only ten resistance genes genetically identified. Of these, only one complex locus consisting of three genes is effective against TTKSK, a widely virulent Pgt race of the Ug99 tribe which emerged in Uganda in 1999 and has since spread to much of East Africa and parts of the Middle East. The objective of this study was to assess the functionality, in barley, of cloned wheat Sr genes effective against race TTKSK. Sr22, Sr33, Sr35 and Sr45 were transformed into barley cv. Golden Promise using Agrobacterium-mediated transformation. All four genes were found to confer effective stem rust resistance. The barley transgenics remained susceptible to the barley leaf rust pathogen Puccinia hordei, indicating that the resistance conferred by these wheat Sr genes was specific for Pgt. Furthermore, these transgenic plants did not display significant adverse agronomic effects in the absence of disease. Cloned Sr genes from wheat are therefore a potential source of resistance against wheat stem rust in barley