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NASA NTRS1977

C-14 in lunar soil - Temperature-release and grain-size dependence

C-14 is released from surface soils by heatings between 600 and 1000 C. The amounts of 600-1000-C C-14 obtained increase inversely with grain diameter for the 10-30 and 74-125-micron size fractions. The inverse relation shows that the 600-1000-C C-14 is located on the surface of grains and probably is of solar-wind origin. The amount released at these temperatures gives 0.002 C-14 ions per sq cm/sec for the implantation rate and corresponds to a C-14/H ratio of the order of 4 hundred-billionths for the solar wind. This result from the grain-size dependence is in accord with C-14 depth-variation studies. The C-14 released above melting temperatures can be attributed to spallation production.

Fireman, E. L.↗

NASA NTRS2001

Reduced T(sub c) Niobium Superconducting HEB Mixers

A reduction in the mixer noise is expected when using superconductors with a lower transition temperature (T(sub c)) since the thermal noise components of the mixer noise should scale with T(sub c). Also, the local oscillator (LO) power required for a diffusion-cooled device should decrease as T(sub c) when T(sub bath) << T(sub c). We previously studied mixing in aluminum based hot-electron bolometers (HEBs) at microwave frequencies (approximately 30 GHz), and observed a significant improvement in noise performance, and a reduction in LO power as predicted. However, the bias voltage range over which good mixer performance was observed was approximately 5 - 10 microV. These devices are thus susceptible to saturation effects, in particular output saturation. In the present work, we have investigated Nb HEBs whose T(sub c) is lowered by applying a magnetic field. The goal is to study a case intermediate between Nb and Al, and hopefully to find properties that will allow use in practical receivers. A 15 kOe perpendicular magnetic field was applied to a Nb HEB (L = 0.16 micrometers, W = 0.08 micrometers, R(sub N) = 90 ohms) to reduce T(sub c) from 5.2 K to 2.4 K. The mixer noise, as inferred from the output noise and the conversion efficiency, decreased from 390 K, DSB to 171 K, DSB. The LO power required for near optimum mixer conversion efficiency (eta(sub mixer) = -9 dB in this device) was 8 nW in zero field, and approximately 2 nW when T(sub c) was reduced to 2.4 K. T(sub bath) = 0.22 K. The conversion bandwidth was previously measured to be 2.4 GHz and the same bandwidth was observed in the presence of a magnetic field. By lowering T(sub c), the voltage range over which good mixing was observed also decreased. However, even with T(sub c) reduced to 2.4 K, the conversion efficiency dropped by 3 dB from its maximum value only when the bias voltage was changed by approximately 90 microV. Saturation effects should thus be much less of a concern in these devices than in Al HEBS. In situations where the application of a large magnetic field is not feasible, we suggest using Ta based HEBS. Ta HEBs should have T(sub c) = 3 - 3.5 K and material properties very similar to Nb.

Siddiqi, I.↗

NASA NTRS2014

Hi-C Observations of Penumbral Bright Dots: Comparison with the IRIS Results

We observed bright dots (BDs) in a sunspot penumbra by using data acquired by the High Resolution Coronal Imager (Hi-C). The sizes of these BDs are on the order of 1 arcsecond (1') and are therefore hard to identify using the Atmospheric Imaging Assembly's (AIA) 0.6' pixel -1 resolution. These BDs become readily apparent with Hi-C's 0.1' pixel -1 resolution. Tian et al. (2014) found penumbral BDs in the transition region (TR) by using the Interface Region Imaging Spectrograph (IRIS). However, only a few of their dots could be associated with any enhanced brightness in AIA channels. In this work, we examine the characteristics of the penumbral BDs observed by Hi-C in a sunspot penumbra, including their sizes, lifetimes, speeds, and intensity. We also attempt to relate these BDs to the IRIS BDs. There are fewer Hi-C BDs in the penumbra than seen by IRIS, though different sunspots were studied and Hi-C had a short observation time. We use 193 A Hi-C data from July 11, 2012 which observed from 18:52:00 UT{18:56:00 UT and supplement it with data from AIA's 193 A passband to see the complete lifetime of the dots that were born before and/or lasted longer than Hi-C's 5-minute observation period. We use additional AIA passbands and compare the light curves of the BDs at different temperatures to test whether the Hi-C BDs are TR BDs. We find that most Hi-C BDs show clear movement, and of those that do, they move in a radial direction, toward or away from the sunspot umbra, sometimes doing both. BDs interact with other BDs, combining to fade away or brighten. The BDs that do not interact with other BDs tend to move less and last longer. We examine the properties of the Hi-C BDs and compare them with the IRIS BDs. Our BDs are similar to the exceptional values of the IRIS BDs: they move slower on average and their sizes and lifetimes are on the higher end of the distributions of IRIS BDs. We infer that our penumbral BDs are some of the larger BDs observed by IRIS.

Alpert, S. E.↗

DOE OSTI2022

First measurement of the $ {\Lambda}_c^{+} $ → pη ' decay

We present the first measurement of the branching fraction of the singly Cabibbo-suppressed (SCS) decay $ {\Lambda}_c^{+} $ → pη ' with η ' → ηπ + π – , using a data sample corresponding to an integrated luminosity of 981 fb – 1 , collected by the Belle detector at the KEKB e + e – asymmetric-energy collider. A significant $ {\Lambda}_c^{+} $ → pη ' signal is observed for the first time with a signal significance of 5.4 σ . The relative branching fraction with respect to the normalization mode $ {\Lambda}_c^{+} $ → pK – π + is measured to be $$ \frac{\mathcal{B}\left({\Lambda}_c^{+}\to p\eta^{\prime}\right)}{\mathcal{B}\left({\Lambda}_c^{+}\to {pK}^{-}{\pi}^{+}\right)}=\left(7.54\pm 1.32\pm 0.73\right)\times {10}^{-3}, $$ where the uncertainties are statistical and systematic, respectively. Using the world-average value of $ \mathcal{B}\left({\Lambda}_c^{+}\to {pK}^{-}{\pi}^{+}\right) $ = (6 . 28 ± 0 . 32) × 10 – 2 , we obtain $$ \mathcal{B}\left({\Lambda}_c^{+}\to p\eta^{\prime}\right)=\left(4.73\pm 0.82\pm 0.46\pm 0.24\right)\times {10}^{-4}, $$ where the uncertainties are statistical, systematic, and from $ \mathcal{B}\left({\Lambda}_c^{+}\to {pK}^{-}{\pi}^{+}\right) $ , respectively.

72 PHYSICS OF ELEMENTARY PARTICLES AND FIELDS↗

DOE OSTI2022

Partial wave analysis of the charmed baryon hadronic decay $ {\Lambda}_c^{+} $ → Λ π +π 0

Based on e + e - collision samples corresponding to an integrated luminosity of 4.4 fb -1 collected with the BESIII detector at center-of-mass energies between 4.6 GeV and 4.7 GeV, a partial wave analysis of the charmed baryon hadronic decay ${\Lambda}_c^{+}$ → Λπ + π 0 is performed, and the decays ${\Lambda}_c^{+}$ → Λρ(770) + and ${\Lambda}_c^{+}$ → Σ(1385)π are studied for the first time. Making use of the world-average branching fraction $\mathcal{B}$ (${\Lambda}_c^{+}$ → Λπ + π 0 ), their branching fractions are determined to be $\mathcal{B}$(${\Lambda}_c^{+}$ → Λρ(770) + ) = (4.06 ± 0.30 ± 0.35 ± 0.23) x 10 -2 , $\mathcal{B}$(${\Lambda}_c^{+}$ → Σ(1385) + )π 0 ) = (5.86 ± 0.49 ± 0.52 ± 0.35) x 10 -3 , $\mathcal{B}$(${\Lambda}_c^{+}$ → Σ(1385) 0 )π + ) = (6.47 ± 0.59 ± 0.66 ± 0.38) x 10 -3 , where the first uncertainties are statistical, the second are systematic, and the third are from the uncertainties of the branching fractions $\mathcal{B}$ ${\Lambda}_c^{+}$ → Λπ + π 0 and $\mathcal{B}$(Σ(1385) → Λπ). In addition, the decay asymmetry parameters are measured to be αΛρ(770)+ = - 0.763 ± 0.053 ± 0.045, $\alpha$ Σ(1385)+π 0 = -0.917 ± 0.069 ± 0.056, and $\alpha$ Σ(1385) 0 π + = -0.789 ± 0.098 ± 0.056.

72 PHYSICS OF ELEMENTARY PARTICLES AND FIELDS↗

DOE OSTI2019

The role of leaf width and conductances to CO 2 in determining water use efficiency in C 4 grasses

C 4 plants achieve higher photosynthesis (An) and intrinsic water use efficiency (iWUE) than C 3 plants, but processes underpinning the variability in An and iWUE across the three C 4 subtypes remain unclear, partly because we lack an integrated framework for quantifying the contribution of diffusional and biochemical limitations to C 4 photosynthesis. We exploited the natural diversity among C 4 grasses to develop an original mathematical approach for estimating eight key processes of C 4 photosynthesis and their relative limitations to An. We also developed a new formulation to estimate mesophyll conductance (gm) based on actual hydration rates of CO 2 by carbonic anhydrases. We found a positive relationship between gm and iWUE and an inverse correlation with gsw among C 4 grasses. We also revealed subtype-specific regulatory processes of iWUE that may be related to known anatomical traits characterising each C 4 subtype. Leaf width was an important determinant of iWUE and showed significant correlations with key limitations of An, especially among NADP-ME species. In conclusion, incorporating leaf width in breeding trials may unlock new opportunities for C 4 crops because the revealed negative relationship between leaf width and iWUE may translate into higher crop and canopy WUE.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2016

C{sub 60} AS A PROBE FOR ASTROPHYSICAL ENVIRONMENTS

The C{sub 60} molecule has been recently detected in a wide range of astrophysical environments through its four active intramolecular vibrational modes ( T {sub 1u}) near 18.9, 17.4, 8.5, and 7.0 μ m. The strengths of the mid-infrared emission bands have been used to infer astrophysical conditions in the fullerene-rich regions. Widely varying values of the relative intrinsic strengths (RIS) of these four bands are reported in laboratory and theoretical papers, which impedes the derivation of the excitation mechanism of C{sub 60} in the astrophysical sources. The spectroscopic analysis of the C{sub 60} samples produced with our method delivers highly reproducible RIS values of 100, 25 ± 1, 26 ± 1 and 40 ± 4. A comparison of the inferred C{sub 60} emission band strengths with the astrophysical data shows that the observed strengths cannot be explained in terms of fluorescent or thermal emission alone. The large range in the observed 17.4 μ m/18.9 μ m emission ratios indicates that either the emission bands contain significant contributions from emitters other than C{sub 60}, or that the population distribution among the C{sub 60} vibrational modes is affected by physical processes other than thermal or UV excitation, such as chemo-luminescence from nascent C{sub 60} or possibly Poincaré fluorescence resulting from an inverse internal energy conversion. We have carefully analyzed the effect of the weakly active fundamental modes and second order modes in the mid-infrared spectrum of C{sub 60}, and propose that neutral C{sub 60} is the carrier of the unidentified emission band at 6.49 μ m which has been observed in fullerene-rich environments.

79 ASTRONOMY AND ASTROPHYSICS↗

DOE OSTI2024

Observation of χ c J → 3 ( K + K − )

By analyzing ( 27.12 ± 0.14 ) × 10 8 ψ ( 3686 ) events collected with the BESIII detector operating at the BEPCII collider, the decay processes χ c J → 3 ( K + K − ) ( J = 0 , 1, 2) are observed for the first time with statistical significances of 8.2 σ , 8.1 σ , and 12.4 σ , respectively. The product branching fractions of ψ ( 3686 ) → γ χ c J , χ c J → 3 ( K + K − ) are presented and the branching fractions of χ c J → 3 ( K + K − ) decays are determined to be B χ c 0 → 3 ( K + K − ) = ( 10.7 ± 1.8 ± 1.1 ) × 10 − 6 , B χ c 1 → 3 ( K + K − ) = ( 4.2 ± 0.9 ± 0.5 ) × 10 − 6 , and B χ c 2 → 3 ( K + K − ) = ( 7.2 ± 1.1 ± 0.8 ) × 10 − 6 , where the first uncertainties are statistical and the second are systematic. Published by the American Physical Society 2024

Astronomy & Astrophysics↗

DOE OSTI2024

Search for the light hadron decay χ c 1 ( 3872 ) → π + π − η

With a data sample corresponding to an integrated luminosity of 11.5 fb − 1 collected with the BESIII detector operating at the BEPCII storage ring, for the first time the light hadron decay χ c 1 ( 3872 ) → π + π − η is searched for. While no significant signal is observed, the upper limits at the 90% confidence level for σ [ e + e − → γ χ c 1 ( 3872 ) ] B [ χ c 1 ( 3872 ) → π + π − η ] at center of mass energies from 4.13 GeV to 4.34 GeV are determined. By normalizing to the χ c 1 ( 3872 ) → π + π − J / ψ decay channel, a 90% confidence level upper limit for the branching fraction ratio R = B [ χ c 1 ( 3872 ) → π + π − η ] / B [ χ c 1 ( 3872 ) → π + π − J / ψ ] < 0.12 is given. These measurements provide important inputs for understanding the internal structure of the χ c 1 ( 3872 ) resonance. Published by the American Physical Society 2024

Astronomy & Astrophysics↗

DOE OSTI2023

Iron‐Catalyzed Intermolecular C−H Amination Assisted by an Isolated Iron‐Imido Radical Intermediate

Here we report the use of a base metal complex [( tBu pyrpyrr 2 )Fe(OEt 2 )] (1-OEt 2 ) ( tBu pyrpyrr 2 2− =3,5-tBu 2 -bis(pyrrolyl)pyridine) as a catalyst for intermolecular amination of C sp3 −H bonds of 9,10-dihydroanthracene (2 a) using 2,4,6-trimethyl phenyl azide (3 a) as the nitrene source. The reaction is complete within one hour at 80 °C using as low as 2 mol % 1-OEt 2 with control in selectivity for single C−H amination versus double C−H amination. Catalytic C−H amination reactions can be extended to other substrates such as cyclohexadiene and xanthene derivatives and can tolerate a variety of aryl azides having methyl groups in both ortho positions. Under stoichiometric conditions the imido radical species [( tBu pyrpyrr 2 )Fe{=N(2,6-Me 2 -4-tBu-C 6 H 2 )] (1-imido) can be isolated in 56 % yield, and spectroscopic, magnetometric, and computational studies confirmed it to be an S = 1 Fe IV complex. Complex 1-imido reacts with 2 a to produce the ferrous aniline adduct [( tBu pyrpyrr 2 )Fe{NH(2,6-Me 2 -4-tBu-C 6 H 2 )(C 14 H 11 )}] (1-aniline) in 45 % yield. Lastly, it was found that complexes 1-imido and 1-aniline are both competent intermediates in catalytic intermolecular C−H amination.

37 INORGANIC, ORGANIC, PHYSICAL, AND ANALYTICAL CH↗

DOE OSTI2015

Dissociation of glucocerebrosidase dimer in solution by its co-factor, saposin C

Mutations in the gene for the lysosomal enzyme glucocerebrosidase (GCase) cause Gaucher disease and are the most common risk factor for Parkinson disease (PD). Analytical ultracentrifugation of 8 μM GCase shows equilibrium between monomer and dimer forms. However, in the presence of its co-factor saposin C (Sap C), only monomer GCase is seen. Isothermal calorimetry confirms that Sap C associates with GCase in solution in a 1:1 complex (K d = 2.1 ± 1.1 μM). Saturation cross-transfer NMR determined that the region of Sap C contacting GCase includes residues 63–66 and 74–76, which is distinct from the region known to enhance GCase activity. Because α-synuclein (α-syn), a protein closely associated with PD etiology, competes with Sap C for GCase binding, its interaction with GCase was also measured by ultracentrifugation and saturation cross-transfer. Unlike Sap C, binding of α-syn to GCase does not affect multimerization. However, adding α-syn reduces saturation cross-transfer from Sap C to GCase, confirming displacement. To explore where Sap C might disrupt multimeric GCase, GCase x-ray structures were analyzed using the program PISA, which predicted stable dimer and tetramer forms. In conclusion, for the most frequently predicted multimer interface, the GCase active sites are partially buried, suggesting that Sap C might disrupt the multimer by binding near the active site.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2025

Quantifying spatial and vertical variations in soil C:N relationships in permafrost-affected landscapes

Permafrost regions are experiencing rapid changes that affect carbon (C) and nitrogen (N) cycles, with implications for vegetation dynamics and gas exchanges with the atmosphere. Soil C:N ratio is a key indicator of organic matter quality, yet spatial estimates of N stocks and C:N ratios lag behind those for C. We used quantile regression forests to compare direct and indirect digital soil mapping approaches for predicting soil C:N ratios at 0–30, 30–60, and 60–100 cm depths across a latitudinal transect in Alaska. The indirect approach – deriving C:N from separately predicted C and N stocks – outperformed direct mapping for the surface layer (0–30 cm), while direct mapping was marginally better at greater depths. However, prediction accuracy decreased with depth for both methods. Temperature and topography were the most important predictors. Both approaches overestimated low and underestimated high C:N ratios, with direct mapping showing greater bias. Our results underscore the challenges of modeling C:N ratios in heterogeneous, data-sparse permafrost soils, but also suggest that indirect mapping holds promise if supported by more extensive datasets.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2021

Long-term leaf C:N ratio change under elevated CO 2 and nitrogen deposition in China: Evidence from observations and process-based modeling

Climate change, elevating atmosphere CO 2 (eCO 2 ) and increased nitrogen deposition (iNDEP) are altering the biogeochemical interactions between plants, microbes and soils, which further modify plant leaf carbon-nitrogen (C:N) stoichiometry and their carbon assimilation capability. Many field experiments have observed large sensitivity of leaf C:N ratio to eCO 2 and iNDEP. However, the large-scale pattern of this sensitivity is still unclear, because eCO 2 and iNDEP drive leaf C:N ratio toward opposite directions, which are further compounded by the complex processes of nitrogen acquisition and plant-and-microbial nitrogen competition. Here, we attempt to map the leaf C:N ratio spatial variation in the past 5 decades in China with a combination of data-driven model and process-based modeling. These two approaches showed consistent results. Over different regions, we found that leaf C:N ratio had significant but uneven changes between 2 time periods (1960-1989 and 1990-2015): a 5% ± 8% increase for temperate grasslands in northern China, a 3% ± 6% increase for boreal grasslands in western China, and by contrast, a 7% ± 6% decrease for temperate forests in southern China, and a 3% ± 5% decrease for boreal forests in northeastern China. Additionally, the structural equation models indicated that the leaf C:N change was sensitive to ΔNDEP, ΔCO 2 and ΔMAT rather than ΔMAP and ecosystem types. In this work, process-based modeling suggested that iNDEP was the main source of soil mineral nitrogen change, dominating leaf C:N ratio change in most areas in China, while eCO 2 led to leaf C:N ratio increase in low iNDEP area. This study also indicates that the long-term leaf C:N ratio acclimation was dominated by climate constraint, especially temperature, but was constrained by soil N availability over decade scale.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2013

Molecular composition of recycled organic wastes, as determined by solid-state {sup 13}C NMR and elemental analyses

Highlights: • Model estimated the molecular C components well for most RO wastes. • Molecular nature of organic matter in RO wastes varied widely. • Molecular composition by NMR modelling preferable to extraction techniques. • Some model shortcomings in estimating molecular composition of biochars. • Waste molecular composition important for carbon/nutrient outcomes in soil. - Abstract: Using solid state {sup 13}C NMR data and elemental composition in a molecular mixing model, we estimated the molecular components of the organic matter in 16 recycled organic (RO) wastes representative of the major materials generated in the Sydney basin area. Close correspondence was found between the measured NMR signal intensities and those predicted by the model for all RO wastes except for poultry manure char. Molecular nature of the organic matter differed widely between the RO wastes. As a proportion of organic C, carbohydrate C ranged from 0.07 to 0.63, protein C from <0.01 to 0.66, lignin C from <0.01 to 0.31, aliphatic C from 0.09 to 0.73, carbonyl C from 0.02 to 0.23, and char C from 0 to 0.45. This method is considered preferable to techniques involving imprecise extraction methods for RO wastes. Molecular composition data has great potential as a predictor of RO waste soil carbon and nutrient outcomes.

12 MANAGEMENT OF RADIOACTIVE AND NON-RADIOACTIVE W↗

DOE OSTI2024

Mechanism of Ni-Catalyzed Photochemical Halogen Atom-Mediated C(sp 3 )–H Arylation

Within the context of Ni photoredox catalysis, halogen atom photoelimination from Ni has emerged as a fruitful strategy for enabling hydrogen atom transfer (HAT)-mediated C(sp 3 )–H functionalization. Despite the numerous synthetic transformations invoking this paradigm, a unified mechanistic hypothesis that is consistent with experimental findings on the catalytic systems and accounts for halogen radical formation and facile C(sp 2 )–C(sp 3 ) bond formation remains elusive. We employ kinetic analysis, organometallic synthesis, and computational investigations to decipher the mechanism of a prototypical Ni-catalyzed photochemical C(sp 3 )–H arylation reaction. Our findings revise the previous mechanistic proposals, first by examining the relevance of SET and EnT processes from Ni intermediates relevant to the HAT-based arylation reaction. Our investigation highlights the ability for blue light to promote efficient Ni–C(sp 2 ) bond homolysis from cationic Ni III and C(sp 2 )–C(sp 3 ) reductive elimination from bipyridine Ni II complexes. However interesting, the rates and selectivities of these processes do not account for the productive catalytic pathway. Instead, our studies support a mechanism that involves halogen atom evolution from in situ generated Ni II dihalide intermediates, radical capture by a Ni II (aryl)(halide) resting state, and key C–C bond formation from Ni III . Oxidative addition to Ni I , as opposed to Ni 0 , and rapid Ni III /Ni I comproportionation play key roles in this process. Furthermore, the findings presented herein offer fundamental insight into the reactivity of Ni in the broader context of catalysis.

37 INORGANIC, ORGANIC, PHYSICAL, AND ANALYTICAL CH↗

DOE OSTI2020

Three Decades of Divergent Land Use and Plant Community Change Alters Soil C and N Content in Tallgrass Prairie

Frequent fire and grazing by megafauna are important determinants of tallgrass prairie plant community structure. However, fire suppression and removal of native grazers have altered these natural disturbance regimes and changed grassland plant communities with potential long-term consequences for soil carbon (C) and nitrogen (N) storage. We investigated multidecade changes in soil C and N pools in response to contrasting long-term burning and grazing treatments. Fire suppression with or without grazers and exclusion of grazers in annually burned prairie increased soil C content and shifted the δ 13 C signature of soil C over time, concomitant with changes in plant community composition. Soil δ 13 C values indicated that increased soil C content was associated with an increased contribution from plants using a C 3 photosynthetic pathway (i.e., woody shrubs) under fire suppression. Soil N content also increased when fire was suppressed, relative to frequently burned grassland, but the rate of increase was slower when grazers were present. Additionally, changes in δ 15 N values suggested that grazing increased the openness of the N cycle, presumably due to greater N losses. By coupling long-term fire and grazing treatments with plant community data and soil samples archived over three decades, we demonstrate that human-caused changes to natural disturbance regimes in a tallgrass prairie significantly alter soil C and N cycles through belowground changes associated with shifts in the plant community. Since natural disturbance regimes have been altered in grasslands across the world, our results are relevant for understanding the long-term biogeochemical consequences of these ongoing land use changes.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2024

Myosin-binding protein C regulates the sarcomere lattice and stabilizes the OFF states of myosin heads

Muscle contraction is produced via the interaction of myofilaments and is regulated so that muscle performance matches demand. Myosin-binding protein C (MyBP-C) is a long and flexible protein that is tightly bound to the thick filament at its C-terminal end (MyBP-C C8C10 ), but may be loosely bound at its middle- and N-terminal end (MyBP-C C1C7 ) to myosin heads and/or the thin filament. MyBP-C is thought to control muscle contraction via the regulation of myosin motors, as mutations lead to debilitating disease. We use a combination of mechanics and small-angle X-ray diffraction to study the immediate and selective removal of the MyBP-C C1C7 domains of fast MyBP-C in permeabilized skeletal muscle. We show that cleavage leads to alterations in crossbridge kinetics and passive structural signatures of myofilaments that are indicative of a shift of myosin heads towards the ON state, highlighting the importance of MyBP-C C1C7 to myofilament force production and regulation.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2021

Evolution of a σ–(c-di-GMP)–anti-σ switch

Filamentous actinobacteria of the genus Streptomyces have a complex lifecycle involving the differentiation of reproductive aerial hyphae into spores. We recently showed c-di-GMP controls this transition by arming a unique anti-σ, RsiG, to bind the sporulation-specific σ, WhiG. The Streptomyces venezuelae RsiG–(c-di-GMP) 2 –WhiG structure revealed that a monomeric RsiG binds c-di-GMP via two E(X) 3 S(X) 2 R(X) 3 Q(X) 3 D repeat motifs, one on each helix of an antiparallel coiled-coil. Here we show that RsiG homologs are found scattered throughout the Actinobacteria. Strikingly, RsiGs from unicellular bacteria descending from the most basal branch of the Actinobacteria are small proteins containing only one c-di-GMP binding motif, yet still bind their WhiG partners. Our structure of a Rubrobacter radiotolerans (RsiG) 2 –(c-di-GMP) 2 –WhiG complex revealed that these single-motif RsiGs are able to form an antiparallel coiled-coil through homodimerization, thereby allowing them to bind c-di-GMP similar to the monomeric twin-motif RsiGs. Further data show that in the unicellular actinobacterium R. radiotolerans , the (RsiG) 2 –(c-di-GMP) 2 –WhiG regulatory switch controls type IV pilus expression. Phylogenetic analysis indicates the single-motif RsiGs likely represent the ancestral state and an internal gene-duplication event gave rise to the twin-motif RsiGs inherited elsewhere in the Actinobacteria. Thus, these studies show how the anti-σ RsiG has evolved through an intragenic duplication event from a small protein carrying a single c-di-GMP binding motif, which functions as a homodimer, to a larger protein carrying two c-di-GMP binding motifs, which functions as a monomer. Consistent with this, our structures reveal potential selective advantages of the monomeric twin-motif anti-σ factors.

59 BASIC BIOLOGICAL SCIENCES↗