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            Effector-Triggered Immunity (ETI) is an important part of the plant immune system, allowing plants to sense and respond to harmful pathogen proteins known as “effectors.” Effectors can be sensed directly or indirectly by NLR (Nucleotide-binding Leucine-rich Repeat) proteins, many of which “guard” the plant proteins targeted by effectors. Although a few effector–target–NLR interactions have been characterized, a general understanding of how these molecular interactions give rise to a functioning immune system is lacking. Here, we present a physics-based model of ETI based on protein–protein interactions. We show that the simplest physical model consistent with the biology gives rise to a robust immune sensor and explains the empirical phenomenon of effector interference as a generic consequence of molecules competing for binding partners. Using the evolutionarily conserved ZAR1 defense gene as a model, we explain how more complex interaction networks integrate multiple pathogen signals into a single response. We then examine alternatives to a guarding architecture, including direct sensing, decoys, and blended “integrated decoy” strategies, and reveal that these sensing architectures obey functional trade-offs between their sensitivity, target protection, and proteomic cost. This allows a quantitative analysis of the trade-offs between different forms of ETI. We discuss these findings in the context of the evolutionary forces shaping the plant immune system.more » « lessFree, publicly-accessible full text available June 17, 2026
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            null (Ed.)Cells possess a multiplicity of non-membrane-bound compartments, which form via liquid-liquid phase separation. These condensates assemble and dissolve as needed to enable central cellular functions. One important class of condensates is those composed of two associating polymer species that form one-to-one specific bonds. What are the physical principles that underlie phase separation in such systems? To address this question, we employed coarse-grained molecular dynamics simulations to examine how the phase boundaries depend on polymer valence, stoichiometry, and binding strength. We discovered a striking phenomenon – for sufficiently strong binding, phase separation is suppressed at rational polymer stoichiometries, which we termed the magic-ratio effect. We further developed an analytical dimer-gel theory that confirmed the magic-ratio effect and disentangled the individual roles of polymer properties in shaping the phase diagram. Our work provides new insights into the factors controlling the phase diagrams of biomolecular condensates, with implications for natural and synthetic systems.more » « less
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            Abstract We analyze a sample of 25 [Nev] (λ3426) emission-line galaxies at 1.4 <z< 2.3 using Hubble Space Telescope/Wide Field Camera 3 G102 and G141 grism observations from the CANDELS LyαEmission at Reionization (CLEAR) survey. [Nev] emission probes extremely energetic photoionization (creation potential of 97.11 eV) and is often attributed to energetic radiation from active galactic nuclei (AGNs), shocks from supernovae, or an otherwise very hard ionizing spectrum from the stellar continuum. In this work, we use [Nev] in conjunction with other rest-frame UV/optical emission lines ([Oii]λλ3726, 3729, [Neiii]λ3869, Hβ, [Oiii]λλ4959, 5007, Hα+[Nii]λλ6548, 6583, [Sii]λλ6716, 6731), deep (2–7 Ms) X-ray observations (from Chandra), and mid-infrared imaging (from Spitzer) to study the origin of this emission and to place constraints on the nature of the ionizing engine. The majority of the [Nev]-detected galaxies have properties consistent with ionization from AGNs. However, for our [Nev]-selected sample, the X-ray luminosities are consistent with local (z≲ 0.1) X-ray-selected Seyferts, but the [Nev] luminosities are more consistent with those fromz∼ 1 X-ray-selected QSOs. The excess [Nev] emission requires either reduced hard X-rays or a ∼0.1 keV excess. We discuss possible origins of the apparent [Nev] excess, which could be related to the “soft (X-ray) excess” observed in some QSOs and Seyferts and/or be a consequence of a complex/anisotropic geometry for the narrow-line region, combined with absorption from a warm, relativistic wind ejected from the accretion disk. We also consider implications for future studies of extreme high-ionization systems in the epoch of reionization (z≳ 6) with the James Webb Space Telescope.more » « less
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            Abstract We use Paschen- β (Pa β ; 1282 nm) observations from the Hubble Space Telescope G141 grism to study the star formation and dust-attenuation properties of a sample of 29 low-redshift ( z < 0.287) galaxies in the CANDELS Ly α Emission at Reionization survey. We first compare the nebular attenuation from Pa β /H α with the stellar attenuation inferred from the spectral energy distribution, finding that the galaxies in our sample are consistent with an average ratio of the continuum attenuation to the nebular gas of 0.44, but with a large amount of excess scatter beyond the observational uncertainties. Much of this scatter is linked to a large variation between the nebular dust attenuation as measured by (space-based) Pa β to (ground-based) H α to that from (ground-based) H α /H β . This implies there are important differences between attenuation measured from grism-based/wide-aperture Pa β fluxes and the ground-based/slit-measured Balmer decrement. We next compare star formation rates (SFRs) from Pa β to those from dust-corrected UV. We perform a survival analysis to infer a census of Pa β emission implied by both detections and nondetections. We find evidence that galaxies with lower stellar mass have more scatter in their ratio of Pa β to attenuation-corrected UV SFRs. When considering our Pa β detection limits, this observation supports the idea that lower-mass galaxies experience “burstier” star formation histories. Together, these results show that Pa β is a valuable tracer of a galaxy’s SFR, probing different timescales of star formation and potentially revealing star formation that is otherwise missed by UV and optical tracers.more » « less
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            null (Ed.)In most environments, organisms compete for limited resources. The number and relative abundance of species that an ecosystem can host is referred to as ‘species diversity’. The competitive-exclusion principle is a hypothesis which proposes that, in an ecosystem, competition for resources results in decreased diversity: only species best equipped to consume the available resources thrive, while their less successful competitors die off. However, many natural ecosystems foster a wide array of species despite offering relatively few resources. Researchers have proposed many competing theories to explain how this paradox can emerge, but they have mainly focused on ecosystems where nutrients are steadily supplied. By contrast, less is known about the way species diversity is maintained when nutrients are only intermittently available, for example in ecosystems that have seasons. To address this question, Erez, Lopez et al. modeled communities of bacteria in which nutrients were repeatedly added and then used up. Depending on conditions, a variety of relationships between the amount of nutrient supplied and community diversity could emerge, suggesting that ecosystems do not follow a simple, universal rule that dictates species diversity. In particular, the resulting communities displayed a higher diversity of microbes than the limit imposed by the competitive-exclusion principle. Further observations allowed Erez, Lopez et al. to suggest guiding principles for when diversity in ecosystems will be maintained or lost. In this framework, ‘early-bird’ species, which rapidly use a subset of the available nutrients, grow to dominate the ecosystem. Even though ‘late-bird’ species are more effective at consuming the remaining resources, they cannot compete with the increased sheer numbers of the ‘early-birds’, leading to a ‘rich-get-richer’ phenomenon. Oceanic plankton, arctic permafrost and many other threatened, resource-poor ecosystems across the world can dramatically influence our daily lives. Closer to home, shifts in the microbe communities that live on the surface of the human body and in the digestive system are linked to poor health. Understanding how species diversity emerges and changes will help to protect our external and internal environments.more » « less
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            Abstract We report on the gas-phase metallicity gradients of a sample of 238 star-forming galaxies at 0.6 < z < 2.6, measured through deep near-infrared Hubble Space Telescope slitless spectroscopy. The observations include 12 orbit depth Hubble/WFC3 G102 grism spectra taken as a part of the CANDELS Ly α Emission at Reionization (CLEAR) survey, and archival WFC3 G102+G141 grism spectra overlapping the CLEAR footprint. The majority of galaxies in this sample are consistent with having a zero or slightly positive metallicity gradient ( dZ / dR ≥ 0, i.e., increasing with radius) across the full mass range probed (8.5 < log M * / M ⊙ < 10.5). We measure the intrinsic population scatter of the metallicity gradients, and show that it increases with decreasing stellar mass—consistent with previous reports in the literature, but confirmed here with a much larger sample. To understand the physical mechanisms governing this scatter, we search for correlations between the observed gradient and various stellar population properties at fixed mass. However, we find no evidence for a correlation with the galaxy properties we consider—including star formation rates, sizes, star formation rate surface densities, and star formation rates per gravitational potential energy. We use the observed weakness of these correlations to provide material constraints for predicted intrinsic correlations from theoretical models.more » « less
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            Many ecosystems, from vegetation to biofilms, are composed of territorial populations that compete for both nutrients and physical space. What are the implications of such spatial organization for biodiversity? To address this question, we developed and analyzed a model of territorial resource competition. In the model, all species obey trade-offs inspired by biophysical constraints on metabolism; the species occupy nonoverlapping territories, while nutrients diffuse in space. We find that the nutrient diffusion time is an important control parameter for both biodiversity and the timescale of population dynamics. Interestingly, fast nutrient diffusion allows the populations of some species to fluctuate to zero, leading to extinctions. Moreover, territorial competition spontaneously gives rise to both multistability and the Allee effect (in which a minimum population is required for survival), so that small perturbations can have major ecological effects. While the assumption of trade-offs allows for the coexistence of more species than the number of nutrients—thus violating the principle of competitive exclusion—overall biodiversity is curbed by the domination of “oligotroph” species. Importantly, in contrast to well-mixed models, spatial structure renders diversity robust to inequalities in metabolic trade-offs. Our results suggest that territorial ecosystems can display high biodiversity and rich dynamics simply due to competition for resources in a spatial community.more » « less
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            Abstract We present high-cadence photometric and spectroscopic observations of SN 2023axu, a classical Type II supernova with an absoluteV-band peak magnitude of –17.2 ± 0.1 mag. SN 2023axu was discovered by the Distance Less Than 40 Mpc (DLT40) survey within 1 day of the last nondetection in the nearby galaxy NGC 2283 at 13.7 Mpc. We modeled the early light curve using a recently updated shock cooling model that includes the effects of line blanketing and found the explosion epoch to be MJD 59971.48 ± 0.03 and the probable progenitor to be a red supergiant. The shock cooling model underpredicts the overall UV data, which point to a possible interaction with circumstellar material. This interpretation is further supported by spectral behavior. We see a ledge feature around 4600 Å in the very early spectra (+1.1 and +1.5 days after the explosion), which can be a sign of circumstellar interaction. The signs of circumstellar material are further bolstered by the presence of absorption features blueward of Hαand Hβat day >40, which is also generally attributed to circumstellar interaction. Our analysis shows the need for high-cadence early photometric and spectroscopic data to decipher the mass-loss history of the progenitor.more » « less
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            Abstract We use Hubble Space Telescope Wide Field Camera 3 G102 and G141 grism spectroscopy to measure rest-frame optical emission-line ratios of 533 galaxies atz∼ 1.5 in the CANDELS LyαEmission at Reionization survey. We compare [Oiii]/Hβversus [Sii]/(Hα+ [Nii]) as an “unVO87” diagram for 461 galaxies and [Oiii]/Hβversus [Neiii]/[Oii] as an “OHNO” diagram for 91 galaxies. The unVO87 diagram does not effectively separate active galactic nuclei (AGN) and [Nev] sources from star-forming galaxies, indicating that the unVO87 properties of star-forming galaxies evolve with redshift and overlap with AGN emission-line signatures atz> 1. The OHNO diagram does effectively separate X-ray AGN and [Nev]-emitting galaxies from the rest of the population. We find that the [Oiii]/Hβline ratios are significantly anticorrelated with stellar mass and significantly correlated with , while [Sii]/(Hα+ [Nii]) is significantly anticorrelated with . Comparison with MAPPINGS V photoionization models indicates that these trends are consistent with lower metallicity and higher ionization in low-mass and high-star formation rate (SFR) galaxies. We do not find evidence for redshift evolution of the emission-line ratios outside of the correlations with mass and SFR. Our results suggest that the OHNO diagram of [Oiii]/Hβversus [Neiii]/[Oii] will be a useful indicator of AGN content and gas conditions in very high-redshift galaxies to be observed by the James Webb Space Telescope.more » « less
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