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At least 361 records · Page 20

Novel candidate genes for lignin structure identified through genome-wide association study of naturally varying Populus trichocarpa

Populus is a promising lignocellulosic feedstock for biofuels and bioproducts. However, the cell wall biopolymer lignin is a major barrier in conversion of biomass to biofuels. To investigate the variability and underlying genetic basis of the complex structure of lignin, a population of 409 three-year-old, naturally varying Populus trichocarpa genotypes were characterized by heteronuclear single quantum coherence (HSQC) nuclear magnetic resonance (NMR). A subsequent genome-wide association study (GWAS) was conducted using approximately 8.3 million single nucleotide polymorphisms (SNPs), which identified 756 genes that were significantly associated (−log 10 ( p -value)>6) with at least one lignin phenotype. Several promising candidate genes were identified, many of which have not previously been reported to be associated with lignin or cell wall biosynthesis. These results provide a resource for gaining insights into the molecular mechanisms of lignin biosynthesis and new targets for future genetic improvement in poplar.

59 BASIC BIOLOGICAL SCIENCES↗

Orbital Evolution of Binary Black Holes in Active Galactic Nucleus Disks: A Disk Channel for Binary Black Hole Mergers?

We perform a series of high-resolution 2D hydrodynamical simulations of equal-mass binary black holes (BBHs) embedded in active galactic nucleus (AGN) accretion disks to study whether these binaries can be driven to merger by the surrounding gas. We find that the gravitational softening adopted for the BBH has a profound impact on this result. When the softening is less than 10% of the binary separation, we show that, in agreement with recent simulations of isolated equal-mass binaries, prograde BBHs expand in time rather than contract. Eventually, however, the binary separation becomes large enough that the tidal force of the central AGN disrupts them. Only when the softening is relatively large do we find that prograde BBHs harden. We determine through detailed analysis of the binary torque, that this dichotomy is due to a loss of spiral structure in the circum-single disks orbiting each black hole when the softening is a significant fraction of the binary separation. Properly resolving these spirals—both with high resolution and small softening—results in a significant source of binary angular momentum. Only for retrograde BBHs do we find consistent hardening, regardless of softening, as these BBHs lack the important spiral structure in their circum-single disks. This suggests that the gas-driven inspiral of retrograde binaries can produce a population of compact BBHs in the gravitational-wave-emitting regime in AGN disks, which may contribute a large fraction to the observed BBH mergers.

79 ASTRONOMY AND ASTROPHYSICS↗

Studies of Vestibular Neurons in Normal, Hyper- and Hypogravity

During the past year, pre-, in- and postflight studies were conducted in association with the Axon project for Bion 10 (Cosmos 2229). Recordings were made during pre- and postflight studies, from 118 horizontal semicircular canal afferents and 27 vestibular nucleus neurons in 7 rhesus monkeys; 137 pulse rotation protocols alone were executed (548 acceleration and deceleration responses were curve fit). Usable data was obtained from 127 horizontal afferents concerning their spontaneous discharge. Curve fits and analysis was made of sinusoidal and sum of sinusoidal responses from 42 and 35 horizontal afferents, respectively. Also recordings were made from neurons inflight from the two flight animals. The mean spontaneous rate varied from 128 spikes/sec. during preflight to 92 spikes/sec during postflight (day 5) - a change of 28%. In direct contrast to the results of Cosmos 2044, the best fitted neural adaptation operator (k) and the gain of the pulse response were decreased during post flight when compared to preflight. Surprisingly, the best fitted gain and k values for the sum of sines were slightly elevated during post flight tests. The gain and phase of single sine responses were compared for pre- and post flight tests and compared to a larger population of afferents. In contrast to Cosmos 2044 results where on the first day of post flight testing the gains of the best fitted sine response were skewed toward the higher values of the Miles and Braitman distribution, the gain of the best fitted sine responses during the first day of post flight testing (day 2) during Cosmos 2229 were exactly on the mode of the Miles and Braitman distribution. Thus, at least for the periodic stimuli, (pulses and sine waves) we found no change in gain and neural adaptation during post flight testing following Cosmos 2229. This conclusion is different from the one derived following the Cosmos 2044 flight. Cosmos flight 2229 differed from Cosmos flight 2044 in several significant ways: For example, during preflight, (1) The animals preflight training was different (less well trained on the gaze task) and (2) the animals were exposed to more experimental manipulations (surgical and rotational). Inflight, (1) the animals were required to make a pointing gesture (motor response) in association with eye movements to obtain reward, (2) the inflight diet was different (more balanced), (3) the feeder for one of the animals clogged following 9 days of flight resulting in evident dehydration and probably less head motion exposure in that monkey and (4) there was limited video taping of the monkeys in space. During postflight, (1) we were unable to test the flight animals until 26 hours postflight as compared to 14.5 hours during Cosmos 2044, (2) the animals received significantly more exposure to motion stimuli during postflight testing than during Cosmos 2044. These differences in the vestibular environment will require analysis of several parameters other than just neural and eye movement responses. For example, computer programs will have to be written and used to recover and quantify the number of head movements made by each animal during flight. This activity is critical to the production of neural adaptation and increased gain.

Correia, Manning J.↗

Titanospirillum velox: a huge, speedy, sulfur-storing spirillum from Ebro Delta microbial mats

A long (20-30 micrometer), wide (3-5 micrometer) microbial-mat bacterium from the Ebro Delta (Tarragona, Spain) was grown in mixed culture and videographed live. Intracellular elemental sulfur globules and unique cell termini were observed in scanning-electron-microprobe and transmission-electron micrographs. A polar organelle underlies bundles of greater than 60 flagella at each indented terminus. These Gram-negative bacteria bend, flex, and swim in a spiral fashion; they translate at speeds greater than 10 body lengths per second. The large size of the spirillum permits direct observation of cell motility in single individual bacteria. After desiccation (i.e., absence of standing water for at least 24 h), large populations developed in mat samples remoistened with sea water. Ultrastructural observations reveal abundant large sulfur globules irregularly distributed in the cytoplasm. A multilayered cell wall, pliable and elastic yet rigid, distends around the sulfur globules. Details of the wall, multiflagellated termini, and large cytoplasmic sulfur globules indicate that these fast-moving spirilla are distinctive enough to warrant a genus and species designation: Titanospirillum velox genus nov., sp. nov. The same collection techniques at a similar habitat in the United States (Plum Island, northeast Essex County, Massachusetts) also yielded large populations of the bacterium among purple phototrophic and other inhabitants of sulfurous microbial-mat muds. The months-long survival of T. velox from Spain and from the United States in closed jars filled with mud taken from both localities leads us to infer that this large spirillum has a cosmopolitan distribution.

Non-NASA Center↗

Direct Observation of the Dynamics of Single-Cell Metabolic Activity during Microbial Diauxic Growth

Understanding how genetic information is realized as the behavior of individual cells is a long-term goal of biology but represents a significant technological challenge. In clonal microbial populations, variation in gene regulation is often interpreted as metabolic heterogeneity. This follows the central dogma of biology, in which information flows from DNA to RNA to protein and ultimately manifests as activity. At present, DNA and RNA can be characterized in single cells, but the abundance and activity of proteins cannot. Inferences about metabolic activity usually therefore rely on the assumption that transcription reflects activity. By tracking the atoms from which they build their biomass, we make direct observations of growth rate and substrate specialization in individual cells throughout a period of growth in a changing environment. This approach allows the flow of information from DNA to be constrained from the distal end of the regulatory cascade and will become an essential tool in the rapidly advancing field of single-cell metabolism.

59 BASIC BIOLOGICAL SCIENCES↗

Dynamic and single cell characterization of a CRISPR-interference toolset in Pseudomonas putida KT2440 for β-ketoadipate production from p -coumarate

We report Pseudomonas putida KT2440 is a well-studied bacterium for the conversion of lignin-derived aromatic compounds to bioproducts. The development of advanced genetic tools in P. putida has reduced the turnaround time for hypothesis testing and enabled the construction of strains capable of producing various products of interest. Here, we evaluate an inducible CRISPR-interference (CRISPRi) toolset on fluorescent, essential, and metabolic targets. Nuclease-deficient Cas9 (dCas9) expressed with the arabinose (8K)-inducible promoter was shown to be tightly regulated across various media conditions and when targeting essential genes. In addition to bulk growth data, single cell time lapse microscopy was conducted, which revealed intrinsic heterogeneity in knockdown rate within an isoclonal population. The dynamics of knockdown were studied across genomic targets in exponentially-growing cells, revealing a universal 1.75 ± 0.38 hour quiescent phase after induction where 1.5 ± 0.35 doublings occur before a phenotypic response is observed. To demonstrate application of this CRISPRi toolset, β-ketoadipate, a monomer for performance-advantaged nylon, was produced at a 4.39 ± 0.5 g/L and yield of 0.76 ± 0.10 mol/mol from p-coumarate, a hydroxycinnamic acid that can be derived from grasses. These cultivation metrics were achieved by using the higher strength IPTG (1K)-inducible promoter to knockdown the pcaIJ operon in the βKA pathway during early exponential phase. This allowed the majority of the carbon to be shunted into the desired product while eliminating the need for a supplemental carbon and energy source to support growth and maintenance.

59 BASIC BIOLOGICAL SCIENCES↗

Reduction of spectroscopic overlap across the Z = 8 shell in neutron-rich nuclei

The recent discovery and spectroscopic measurements of 27 O and 28 O suggests the disappearance of the N = 20 shell structure in these neutron-rich oxygen isotopes. We measured one- and two-proton removal cross sections from 27 F and 29 N, respectively, extracting spectroscopic factors and comparing them to shell model overlap functions coupled with eikonal reaction model calculations. The invariant mass technique was used to reconstruct the two-body ( 24 O + n) and three-body ( 24 O + 2n) decay energies from knockout reactions of 27 F (106.2 MeV/u) and 29 Ne (112.8 MeV/u) beams impinging on a 9 Be target. The one-proton removal from 27 F strongly populated the ground state of 26 O and the extracted cross section of $3.4^{+0.3}_{-1.5}$ mb agrees with eikonal model calculations that are normalized by the shell model spectroscopic factors and account for the systematic reduction factor observed for single nucleon removal reactions within the models used. For the two-proton removal reaction from 29 Ne an upper limit of 0.08 mb was extracted for populating states in 27 O decaying though the ground state of 26 O. The measured upper limit for the population of the ground state of 26 O in the two-proton removal reaction from 29 Ne indicates a significant difference in the underlying nuclear structure of 27 F and 29 Ne.

73 NUCLEAR PHYSICS AND RADIATION PHYSICS↗

Surfactant-assisted one-pot sample preparation for label-free single-cell proteomics

Large numbers of cells are generally required for quantitative global proteome profiling due to surface adsorption losses associated with sample processing. Such bulk measurement obscures important cell-to-cell variability (cell heterogeneity) and makes proteomic profiling impossible for rare cell populations (e.g., circulating tumor cells (CTCs)). Here we report a surfactant-assisted one-pot sample preparation coupled with mass spectrometry (MS) method termed SOP-MS for label-free global single-cell proteomics. SOP-MS capitalizes on the combination of a MS-compatible nonionic surfactant, n-Dodecyl-β-D-maltoside, and hydrophobic surface-based low-bind tubes or multi-well plates for ‘all-in-one’ one-pot sample preparation. This ‘all-in-one’ method including elimination of all sample transfer steps maximally reduces surface adsorption losses for effective processing of single cells, thus improving detection sensitivity for single-cell proteomics. This method allows convenient label-free quantification of hundreds of proteins from single human cells and ~1200 proteins from small tissue sections (close to ~20 cells). When applied to a patient CTC-derived xenograft (PCDX) model at the single-cell resolution, SOP-MS can reveal distinct protein signatures between primary tumor cells and early metastatic lung cells, which are related to the selection pressure of anti-tumor immunity during breast cancer metastasis. The approach paves the way for routine, precise, quantitative single-cell proteomics.

36 MATERIALS SCIENCE↗

High-throughput Single-Cell Proteomics and Transcriptomics from the Same Cells with a Nanoliter-Scale Spin-Transfer Approach

Single-cell multiomic platforms provide a comprehensive snapshot of cellular states and cell types by offering critical insights into the spatiotemporal regulation of biomolecular networks at a systems level, thereby defining the basis of multicellularity. Here, we introduce nanoSPINS, an advanced platform that enables high-throughput profiling and integrative analysis of the transcriptome and proteome from the same single cells using RNA sequencing and isobaric labeling LC-MS-based proteomics, respectively. NanoSPINS can efficiently transfer mRNA-containing droplets across two microarrays via a centrifugation-based approach, while proteins are retained on the initial platform. Benchmarking of nanoSPINS on two cell lines demonstrates its ability to generate global proteomic and transcriptomic profiles that align well with previously established methodologies/platforms. The incorporation of isobaric TMTpro labeling into this single-cell multiomics platform significantly enhances the throughput of single-cell proteomic analyses. Through the high-throughput quantification of the proteome and transcriptome, nanoSPINS not only facilitates the identification of molecular features at both mRNA and protein level but also provides larger sample sizes for improved statistical power in clustering and differential abundance. Given the broad applicability of single-cell multiomics in biological research and clinical settings, we believe nanoSPINS represents a powerful platform for the characterization of heterogeneous cell populations.

multi 'omics↗

Quenching of Single-Particle Strength in A=15 Nuclei

Here, absolute cross sections for the addition of s- and d-wave neutrons to 14 C and 14 N have been determined simultaneously via the (d,p) reaction at 10 MeV/u. The difference between the neutron and proton separation energies, ΔS, is around -20 MeV for the 14 C+n system and +8 MeV for 14 N+n. The population of the 1s 1/2 and 0d 5/2 orbitals for both systems is reduced by a factor of approximately 0.5 compared with the independent single-particle model, or about 0.6 when compared with the shell model. This finding strongly contrasts with results deduced from intermediate-energy knockout reactions between similar nuclei on targets of 9 Be and 12 C. The simultaneous technique used removes many systematic uncertainties.

73 NUCLEAR PHYSICS AND RADIATION PHYSICS↗

Post-AGB Stars in Nearby Galaxies as Calibrators for HST

This report summarizes activities carried out with support from the NASA Ultraviolet, Visible, and Gravitational Astrophysics Research and Analysis Program under Grant NAG 5-6821. The Principal Investigator is Howard E. Bond (Space Telescope Science Institute). STScI Postdoctoral Associates Laura K. Fullton (1998), David Alves (1998-2001), and Michael Siegel (2001) were partially supported by this grant. The aim of the program is to calibrate the absolute magnitudes of post-asymptotic- giant-branch (post-AGB or PAGB) stars, which we believe will be an excellent new "standard candle" for measuring extragalactic distances. The argument is that, in old populations, the stars that are evolving through the PAGB region of the HR diagram arise from only a single main-sequence turnoff mass. In addition, theoretical PAGB evolutionary tracks show that they evolve through this region at constant luminosity; hence the PAGB stars should have an extremely narrow luminosity function. Moreover, as the PAGB stars evolve through spectral types F and A (en route from the AGB to hot stellar remnants and white dwarfs), they have the highest luminosities attained by old stars (both bolometrically and in the visual band). Finally, PAGB stars of these spectral types are very easily identified. because of their large Balmer jumps, which are due to their very low surface gravities. Our approach is first to identify PAGB stars in Milky Way globular clusters and in other Local Group galaxies, which are at known distances, and thus to measure accurate absolute magnitudes for the PAGB stars. With this Milky Way and Local Group luminosity calibration, we will then be in a position to find PAGB stars in more distant galaxies from the ground, and ultimately from the Hubble Space Telescope. and thus derive distances. These PAGB stars are, as noted above, the visually brightest members of Population II, and hence will allow distance measurements to galaxies that do not contain Cepheids, such as elliptical galaxies, as well as distances to spirals using PAGB stars in their halos. Moreover, the method is entirely independent of Cepheids. and thus provides a direct test of the Cepheid distance scale. The program will also provide information on the evolutionary lifetimes of PAGB stars.

Bond, Howard E.↗

Constraints on the formation history of the elliptical galaxy NGC 3923 from the colors of its globular clusters

We present a study of the colors of globular clusters associated with the elliptical galaxy NGC 3923. Our final sample consists of Wasington system C and T(sub 1) photometry for 143 globular cluster candidates with an expected contamination of no more than 10%. We find that the color distribution of the NGC 3923 globular cluster system (GCS) is broad and appears to have at least two peaks. A mixture modeling analysis of the color distribution indicates that a two-component model is favored over a single-component one at a high level of confidence (greater than 99%). This evidence for more than one population in the GCS of NGC 3923 is similar to that previously noted for the four other elliptical galaxies for which similar data have been published. Furthermore, we find that the NGC 3923 GCS is redder than the GCSs of previously studed elliptical galaxies of similar luminosity. The median metallicity inferred from our (C-(T(sub 1)))(sub 0) colors is (Fe/H)(sub med) = -0.56, with an uncertainty of 0.14 dex arising from all sources of uncertainty in the mean color. This is more metal rich than the median metallicity found for the GCS of M87 using the same method, (Fe/H)(sub med) = -0.94. Since M87 is more luminous than NGC 3923, this result points to significant scatter about any trend of higher GCS metallicity with increasing galaxy luminosity. We also show that there is a color gradient in the NGC 3923 GCS corresponding to about -0.5 dex in Delta(Fe/H)/Delta(log r). We conclude that the shape of the color distribution of individual GCSs and the variation in mean color among the GCSs of ellipticals are difficult to understand if elliptical galaxies are formed in a single protogalactic collapse. Models in which ellipticals and their globular clusters are formed in more than one event, such as a merger scenario, are more successful in accounting for these observations.

Zepf, Stephen E.↗

Charged- and Multi-Exciton Dynamics in Colloidal Quantum Dot Molecules

Multicarrier states in quantum dots are confined to small volumes, resulting in increased nonradiative Auger recombination rates with implications for different optoelectronic applications. Recently, the fusion of two core-shell quantum dots into a dimer has provided a new physical landscape for multiexciton states, since the excitons may share a core (intradot, localized) or occupy different cores (interdot, segregated). Here, in this study, we employ transient absorption spectroscopy to investigate the multiexciton dynamics in coupled quantum dot dimers. We observe that multiexciton populations in the dimers live significantly longer in comparison to the parent monomers, in contrast to the single exciton regime. A kinetic model that accounts for the statistical differences between monomers and dimers reveals that, while intradot multiexcitons show Auger rates similar to the monomers, interdot states have reduced Auger rates. These results pave the way for the rational design of new quantum dot molecules with tailored multiexciton properties.

Auger recombination↗

Identification of a parasitic symbiosis between respiratory metabolisms in the biogeochemical chlorine cycle

A key step in the chlorine cycle is the reduction of perchlorate (ClO 4 - ) and chlorate (ClO 3 - ) to chloride by microbial respiratory pathways. Perchlorate-reducing bacteria and chlorate-reducing bacteria differ in that the latter cannot use perchlorate, the most oxidized chlorine compound. However, a recent study identified a bacterium with the chlorate reduction pathway dominating a community provided only perchlorate. Here we confirm a metabolic interaction between perchlorate- and chlorate-reducing bacteria and define its mechanism. Perchlorate-reducing bacteria supported the growth of chlorate-reducing bacteria to up to 90% of total cells in communities and co-cultures. Chlorate-reducing bacteria required the gene for chlorate reductase to grow in co-culture with perchlorate-reducing bacteria, demonstrating that chlorate is responsible for the interaction, not the subsequent intermediates chlorite and oxygen. Modeling of the interaction suggested that cells specialized for chlorate reduction have a competitive advantage for consuming chlorate produced from perchlorate, especially at high concentrations of perchlorate, because perchlorate and chlorate compete for a single enzyme in perchlorate-reducing cells. We conclude that perchlorate-reducing bacteria inadvertently support large populations of chlorate-reducing bacteria in a parasitic relationship through the release of the intermediate chlorate. An implication of these findings is that undetected chlorate-reducing bacteria have likely negatively impacted efforts to bioremediate perchlorate pollution for decades.

59 BASIC BIOLOGICAL SCIENCES↗

Counting generations in birth and death processes with competing Erlang and exponential waiting times

Lymphocyte populations, stimulated in vitro or in vivo, grow as cells divide. Stochastic models are appropriate because some cells undergo multiple rounds of division, some die, and others of the same type in the same conditions do not divide at all. If individual cells behave independently, then each cell can be imagined as sampling from a probability density of times to division and death. The exponential density is the most mathematically and computationally convenient choice. It has the advantage of satisfying the memoryless property, consistent with a Markov process, but it overestimates the probability of short division times. With the aim of preserving the advantages of a Markovian framework while improving the representation of experimentally-observed division times, we consider a multi-stage model of cellular division and death. We use Erlang-distributed (or, more generally, phase-type distributed) times to division, and exponentially distributed times to death. We classify cells into generations, using the rule that the daughters of cells in generation n are in generation n + 1. In some circumstances, our representation is equivalent to established models of lymphocyte dynamics. We find the growth rate of the cell population by calculating the proportions of cells by stage and generation. The exponent describing the late-time cell population growth, and the criterion for extinction of the population, differs from what would be expected if N steps with rate λ were equivalent to a single step of rate λ/N. We link with a published experimental dataset, where cell counts were reported after T cells were transferred to lymphopenic mice, using Approximate Bayesian Computation. In the comparison, the death rate is assumed to be proportional to the generation and the Erlang time to division for generation 0 is allowed to differ from that of subsequent generations. The multi-stage representation is preferred to a simple exponential in posterior distributions, and the mean time to first division is estimated to be longer than the mean time to subsequent divisions.

59 BASIC BIOLOGICAL SCIENCES↗

First 𝛽-Delayed Two-Neutron Spectroscopy of the 𝑟-Process Nucleus 134 In and Observation of the 𝑖 13/2 Single-Particle Neutron State in 133 Sn

This manuscript reports on the direct observation of a 𝛽-delayed two-neutron emission in a study of 134 In at the ISOLDE Decay Station using neutron spectroscopy. We also report on the first measurement in 𝛽 − decay of the long-sought 13/2 + excited state in 133 Sn, attributed to be the neutron single-particle 𝑖 13/2 orbital. The observation of sequential neutron emission is used to extract the relative population of the 𝑖 13/2 state, which was found to be much smaller than the predictions of the statistical model. The experiment was possible because of the innovative use of a neutron array with neutron discrimination and interaction tracking capabilities. This is the first study of the details of the two-neutron emission for a nucleus, which belongs to the 𝑟-process path. Understanding 𝛽-delayed two-neutron emission probabilities is essential to validate models used in astrophysical 𝑟-process nucleosynthesis calculations. Observing two-neutron emissions in 𝛽 − decay paves the way for new experiments to study energy and angular correlations for 𝛽-delayed multineutron emitters.

Beta decay↗

Evidence for Cosmic Acceleration Is Robust to Observed Correlations between Type Ia Supernova Luminosity and Stellar Age

Type Ia supernovae (SNe Ia) are powerful standardizable candles for constraining cosmological models and provided the first evidence of the accelerated expansion of the universe. Their precision derives from empirical correlations, now measured from >1000 SNe Ia, between their luminosities, light-curve shapes, colors, and most recently with the stellar mass of their host galaxy. As mass correlates with other galaxy properties, alternative parameters have been investigated to improve SN Ia standardization though none have been shown to significantly alter the determination of cosmological parameters. We re-examine a recent claim, based on 34 SN Ia in nearby passive host galaxies, of a 0.05 mag Gyr -1 dependence of standardized SN Ia luminosity on host age, which, if extrapolated to higher redshifts, would be a bias up to 0.25 mag, challenging the inference of dark energy. We reanalyze this sample of hosts using both the original method and a Bayesian hierarchical model and find after a fuller accounting of the uncertainties the significance of a dependence on age to be ≤2σ and ~1σ after the removal of a single poorly sampled SN Ia. Here, to test the claim that a trend seen in old stellar populations can be applied to younger ages, we extend our analysis to a larger sample that includes young hosts. We find the residual dependence of host age (after all standardization typically employed for cosmological measurements) to be consistent with zero for 254 SNe Ia from the Pantheon sample, ruling out the large but low significance trend seen in passive hosts.

79 ASTRONOMY AND ASTROPHYSICS↗

Hydride structures in Ti-aluminides subjected to high temperature and hydrogen pressure charging conditions

The distribution and chemistry of hydrides produced in single and dual phase alloys with a composition near TiAl have been investigated by using a combination of TEM and X-ray diffraction techniques. The alloys were exposed at 650 C to 13.8 MPa of gaseous H2 for 100 h. In the single-phase gamma alloy, large hydrides preferentially nucleated on the grain boundaries and matrix dislocations and a population of small hydrides was distributed throughout the matrix. X-ray and electron diffraction patterns from these hydrides indicated that they have an fcc structure with a lattice parameter of 0.45 nm. EDAX analysis of the hydrides showed that they were enriched in Ti. The hydrides were mostly removed by vacuum annealing at 800 C for 24 h. On dissolution of the hydrides, the chemistry of hydride-free regions of the grain boundary returned to the matrix composition, suggesting that Ti segregation accompanied the hydride formation rather than Ti enrichment causing the formation of the hydride.

Legzdina, D.↗