Soil microbial community response to corrinoids is shaped by a natural reservoir of vitamin B12.

Soil microbial communities perform critical ecosystem services through the collective metabolic activities of numerous individual organisms. Most microbes use corrinoids, a structurally diverse family of cofactors related to vitamin B12. Corrinoid structure influences the growth of individual microbes, yet how these growth responses scale to the community level remains unknown. Analysis of metagenome-assembled genomes suggests corrinoids are supplied to the community by members of the archaeal and bacterial phyla Thermoproteota, Actinobacteria, and Proteobacteria. Corrinoids were found largely adhered to the soil matrix in a grassland soil, at levels exceeding those required by cultured bacteria. Enrichment cultures and soil microcosms seeded with different corrinoids showed distinct shifts in bacterial community composition, supporting the hypothesis that corrinoid structure can shape communities. Environmental context influenced both community and taxon-specific responses to specific corrinoids. These results implicate corrinoids as key determinants of soil microbiome structure and suggest that environmental micronutrient reservoirs promote community stability.

Phylogenetic distribution and experimental characterization of corrinoid production and dependence in soil bacterial isolates.

Soil microbial communities impact carbon sequestration and release, biogeochemical cycling, and agricultural yields. These global effects rely on metabolic interactions that modulate community composition and function. However, the physicochemical and taxonomic complexity of soil and the scarcity of available isolates for phenotypic testing are significant barriers to studying soil microbial interactions. Corrinoids-the vitamin B12 family of cofactors-are critical for microbial metabolism, yet they are synthesized by only a subset of microbiome members. Here, we evaluated corrinoid production and dependence in soil bacteria as a model to investigate the ecological roles of microorganisms involved in metabolic interactions. We isolated and characterized a taxonomically diverse collection of 161 soil bacteria from a single study site. Most corrinoid-dependent bacteria in the collection prefer B12 over other corrinoids, while all tested producers synthesize B12, indicating metabolic compatibility between producers and dependents in the collection. Furthermore, a subset of producers release B12 at levels sufficient to support dependent isolates in laboratory culture at estimated ratios of up to 1000 dependents per producer. Within our isolate collection, we did not find strong phylogenetic patterns in corrinoid production or dependence. Upon investigating trends in the phylogenetic dispersion of corrinoid metabolism categories across sequenced bacteria from various environments, we found that these traits are conserved in 47 out of 85 genera. Together, these phenotypic and genomic results provide evidence for corrinoid-based metabolic interactions among bacteria and provide a framework for the study of nutrient-sharing ecological interactions in microbial communities.

Soil microbial community response to corrinoids is shaped by a natural reservoir of vitamin B12.

Soil microbial communities perform critical ecosystem services through the collective metabolic activities of numerous individual organisms. Most microbes use corrinoids, a structurally diverse family of cofactors related to vitamin B12. Corrinoid structure influences the growth of individual microbes, yet how these growth responses scale to the community level remains unknown. Analysis of metagenome-assembled genomes suggests corrinoids are supplied to the community by members of the archaeal and bacterial phyla Thermoproteota, Actinobacteria, and Proteobacteria. Corrinoids were found largely adhered to the soil matrix in a grassland soil, at levels exceeding those required by cultured bacteria. Enrichment cultures and soil microcosms seeded with different corrinoids showed distinct shifts in bacterial community composition, supporting the hypothesis that corrinoid structure can shape communities. Environmental context influenced both community and taxon-specific responses to specific corrinoids. These results implicate corrinoids as key determinants of soil microbiome structure and suggest that environmental micronutrient reservoirs promote community stability.

Laboratory evolution of E. coli with a natural vitamin B12 analog reveals roles for cobamide uptake and adenosylation in methionine synthase-dependent growth.

The majority of bacteria use cobamides as cofactors for methionine synthesis or other diverse metabolic processes. Cobamides are a structurally diverse family of cofactors related to vitamin B12 (cobalamin), and most bacteria studied to date grow most robustly with particular cobamides. Because different environments contain varying abundances of distinct cobamides, bacteria are likely to encounter cobamides that do not function efficiently for their metabolism. Here, we performed a laboratory evolution of a cobamide-dependent strain of Escherichia coli with pseudocobalamin (pCbl), a cobamide that E. coli uses less effectively than cobalamin for MetH-dependent methionine synthesis, to identify genetic adaptations that lead to improved growth with less-preferred cobamides. After propagating and sequencing nine independent lines and validating the results by constructing targeted mutations, we found that increasing expression of the outer membrane cobamide transporter BtuB is beneficial during growth under cobamide-limiting conditions. Unexpectedly, we also found that overexpression of the cobamide adenosyltransferase BtuR confers a specific growth advantage in pCbl. Characterization of this phenotype revealed that BtuR and adenosylated cobamides contribute to optimal MetH-dependent growth. Together, these findings improve our understanding of how bacteria expand their cobamide-dependent metabolic potential.

Cobalamin Riboswitches Are Broadly Sensitive to Corrinoid Cofactors to Enable an Efficient Gene Regulatory Strategy.

In bacteria, many essential metabolic processes are controlled by riboswitches, gene regulatory RNAs that directly bind and detect metabolites. Highly specific effector binding enables riboswitches to respond to a single biologically relevant metabolite. Cobalamin riboswitches are a potential exception because over a dozen chemically similar but functionally distinct cobalamin variants (corrinoid cofactors) exist in nature. Here, we measured cobalamin riboswitch activity in vivo using a Bacillus subtilis fluorescent reporter system and found, among 38 tested riboswitches, a subset responded to corrinoids promiscuously, while others were semiselective. Analyses of chimeric riboswitches and structural models indicate, unlike other riboswitch classes, cobalamin riboswitches indirectly differentiate among corrinoids by sensing differences in their structural conformation. This regulatory strategy aligns riboswitch-corrinoid specificity with cellular corrinoid requirements in a B. subtilis model. Thus, bacteria can employ broadly sensitive riboswitches to cope with the chemical diversity of essential metabolites. IMPORTANCE Some bacterial mRNAs contain a region called a riboswitch which controls gene expression by binding to a metabolite in the cell. Typically, riboswitches sense and respond to a limited range of cellular metabolites, often just one type. In this work, we found the cobalamin (vitamin B12) riboswitch class is an exception, capable of sensing and responding to multiple variants of B12-collectively called corrinoids. We found cobalamin riboswitches vary in corrinoid specificity with some riboswitches responding to each of the corrinoids we tested, while others responding only to a subset of corrinoids. Our results suggest the latter class of riboswitches sense intrinsic conformational differences among corrinoids in order to support the corrinoid-specific needs of the cell. These findings provide insight into how bacteria sense and respond to an exceptionally diverse, often essential set of enzyme cofactors.

Purification and detection of vitamin B12 analogs.

Cobamides are a family of enzyme cofactors that are required by organisms in all domains of life. Over a dozen cobamides exist in nature although only cobalamin (vitamin B12), the cobamide required by humans, has been studied extensively. Cobamides are exclusively produced by a subset of prokaryotes. Importantly, the bacteria and archaea that synthesize cobamides de novo typically produce a single type of cobamide, and furthermore, organisms that use cobamides are selective for certain cobamides. Therefore, a detailed understanding of the cobamide-dependent metabolism of an organism or microbial community of interest requires experiments performed with a variety of cobamides. A notable challenge is that cobalamin is the only cobamide that is commercially available at present. In this chapter, we describe methods to extract, purify, and quantify various cobamides from bacteria for use in laboratory experiments.

Regulation of proton partitioning in kinase-activating acute myeloid leukemia and its therapeutic implication.

Gain-of-function kinase mutations are common in AML and usually portend an inferior prognosis. We reported a novel mechanism whereby kinase mutants induced intracellular alkalization characteristic in oncogenesis. Thirteen kinases were found to activate sodium/hydrogen exchanger (NHE1) in normal hematopoietic progenitors, of which FLT3-ITD, KRASG12D, and BTK phosphorylated NHE1 maintained alkaline intracellular pH (pHi) and supported survival of AML cells. Primary AML samples with kinase mutations also showed increased NHE1 phosphorylation and evidence of NHE1 addiction. Amiloride enhanced anti-leukemic effects and intracellular distribution of kinase inhibitors and chemotherapy. Co-inhibition of NHE1 and kinase synergistically acidified pHi in leukemia and inhibited its growth in vivo. Plasma from patients taking amiloride for diuresis reduced pHi of leukemia and enhanced cytotoxic effects of kinase inhibitors and chemotherapy in vitro. NHE1-mediated intracellular alkalization played a key pathogenetic role in transmitting the proliferative signal from mutated-kinase and could be exploited for therapeutic intervention in AML.

Impact of cold plasma on the techno-functional and sensory properties of whey dairy beverage added with xylooligosaccharide.

This study aimed to evaluate the impact of cold plasma (0, 5, 10, or 15 min) on the techno-functional and sensory properties of whey dairy beverages added with xylooligosaccharide (XOS, 1.5% p/v). Untreated and pasteurized whey beverages were also evaluated. The products were evaluated for physicochemical characteristics, bioactive compounds, XOS stability, rheological properties, and sensory characteristics. Cold plasma and pasteurized products presented lower color intensity (L*=87.4-87.9, a*=-0.24- -0.60, b*=2.41-5.19), reduced consistency (K = 4.31-42.21 mPa.sn and N = 0.57-0.95), and similar apparent viscosity, XOS chemical stability, and sensory characteristics compared with the untreated product. However, the cold plasma-treated beverages presented lower heat load indicators (hydroxymethylfurfural [HMF] values of 1.91-2.10 µmol/L and whey protein nitrogen index [WPNI] of 6.09-6.66 µmol/L) and a higher concentration of bioactive compounds (antioxidant activity [5.31-9.30%], and inhibition of ACE [14.17-22.53%], α-amylase [18.52-25.67%] and α-glucosidase [22.50-27.50%] activities) than the pasteurized product, being the effects more pronounced for the higher exposure times. Overall, cold plasma has important advantages for the processing of whey beverages added with XOS.

A multidisciplinary evaluation of suspected, non-confirmed cases of COVID-19 including chest CT, as compared to World Health Organization recommendations.

AIM: To report an audit of the evaluation of suspected, unconfirmed cases of COVID-19 including chest computed tomography (CT), as compared to World Health Organization recommendations. METHODS: A clinical audit was undertaken examining the evaluation of patients with suspected COVID-19 with negative SARS-CoV-2 reverse transcriptase polymerase chain reaction (RT-PCR) results, with comparison to WHO recommendations. A retrospective chart review was undertaken for 90 patients examining investigations, in particular CT, used to clarify the diagnosis. RESULTS: Ninety patients underwent additional investigation. Seventy-five per cent adherence to WHO recommendations was observed. Fifty-two men (57.78%) and 38 (42.22%) women were investigated, with a median age of 69 years (range 20-96 years). Seventy-nine chest CT examinations demonstrated positive, indeterminate, and negative rates for COVID-19 of 3.79%, 24.1%, and 72.15% respectively. Three patients had discordant swab results with initially negative and subsequently positive results for SARS-CoV-2, resulting in false-negative rates of 5.1% for those retested. Combining discordant RT-PCR swab results, positive radiology, and patients treated as COVID-19-positive due to indeterminate radiology and highly consistent symptoms, resulted in a false-negative rate for initial SARS-CoV-2 RT-PCR swabs of 16.67%. CONCLUSION: Seventy-five per cent compliance with relevant WHO guidance and a false-negative rate for initial swabs of 16.67% was demonstrated. Further evidence is needed to fully determine the utility of chest CT in the diagnosis of COVID-19 in the context of initial false-negative RT-PCR results.

Identification of a Novel Cobamide Remodeling Enzyme in the Beneficial Human Gut Bacterium Akkermansia muciniphila.

The beneficial human gut bacterium Akkermansia muciniphila provides metabolites to other members of the gut microbiota by breaking down host mucin, but most of its other metabolic functions have not been investigated. A. muciniphila strain MucT is known to use cobamides, the vitamin B12 family of cofactors with structural diversity in the lower ligand. However, A. muciniphila MucT is unable to synthesize cobamides de novo, and the specific forms that can be used by A. muciniphila have not been examined. We found that the levels of growth of A. muciniphila MucT were nearly identical with each of seven cobamides tested, in contrast to nearly all bacteria that had been studied previously. Unexpectedly, this promiscuity is due to cobamide remodeling-the removal and replacement of the lower ligand-despite the absence of the canonical remodeling enzyme CbiZ in A. muciniphila We identified a novel enzyme, CbiR, that is capable of initiating the remodeling process by hydrolyzing the phosphoribosyl bond in the nucleotide loop of cobamides. CbiR does not share similarity with other cobamide remodeling enzymes or B12-binding domains and is instead a member of the apurinic/apyrimidinic (AP) endonuclease 2 enzyme superfamily. We speculate that CbiR enables bacteria to repurpose cobamides that they cannot otherwise use in order to grow under cobamide-requiring conditions; this function was confirmed by heterologous expression of cbiR in Escherichia coli Homologs of CbiR are found in over 200 microbial taxa across 22 phyla, suggesting that many bacteria may use CbiR to gain access to the diverse cobamides present in their environment.IMPORTANCE Cobamides, comprising the vitamin B12 family of cobalt-containing cofactors, are required for metabolism in all domains of life, including most bacteria. Cobamides have structural variability in the lower ligand, and selectivity for particular cobamides has been observed in most organisms studied to date. Here, we discovered that the beneficial human gut bacterium Akkermansia muciniphila can use a diverse range of cobamides due to its ability to change the cobamide structure via a process termed cobamide remodeling. We identify and characterize the novel enzyme CbiR that is necessary for initiating the cobamide remodeling process. The discovery of this enzyme has implications for understanding the ecological role of A. muciniphila in the gut and the functions of other bacteria that produce this enzyme.

Effect of pollutant source location on air pollutant dispersion around a high-rise building.

This article investigates the dispersion of airborne pollutants emitted from different locations near a high-rise building. A Computational Fluid Dynamics (CFD) model for simulating the wind flow field and the pollutant dispersion was developed and validated by wind tunnel data. Then the spreading of the pollutant emitted from different locations to a rectangular-shaped high-rise residential (HRR) building was numerically studied. The pollutant source location was set in a wide range of the position angle and distance between the source and the building. It was found that the pollutant concentration on the building decreases with an increase in the emission distance whereas the effect of the position angle is more complicated. Interestingly, there is a critical range of the position angle from which the emitted pollutants will not spread to the building in a significant way. The effect of the source location was linked to the wind flow field around the building, particularly with several major flows. The vertical distributions of the pollutant concentration on different faces were also investigated, and it was found that these are more affected by the vertical flow near each face. Finally, a mathematical model was developed to evaluate the pollutant concentration as a function of the emission distance and position angle. These findings are helpful to the understanding of the dispersion of airborne pollutants around high-rise buildings and the related hazard management in urban design.

Capacity building for dementia care in community care services: a mixed methods approach.

BACKGROUND: The prevalence of dementia is surging that results in huge service demand in the community care services. Dementia care competence of staff working in these settings is fundamental of the care quality. This project aims to examine the effects of staff training on their competence for the anticipated challenges in dementia care and explore how the training influence their care practices. METHODS: This study adopted a mixed methods triangulation design, including a prospective multi-center study with pre-test post-test evaluations and a narrative analysis of the participants' reflective essays. Seventeen experienced health and social care professionals were trained as trainers at the Dementia Services Development Centre of the University of Stirling, UK. The trainers provided local facilitator training to staff members by using training materials that were culturally adapted to the local context. The facilitators were required to deliver 12 two-hour in-service training sessions for 6 months to their colleagues in a small group format in their respective workplace. Eventually a total of 1347 staff members from community care centers, day care centers, outreach teams and care homes of 70 non-government organizations in Hong Kong participated in the study between April 2017 and December 2018. Validated instruments were used to measure knowledge, attitude, sense of competence in dementia care and job satisfaction at the baseline and at 12-month follow-up. All participants were required to write a reflective essay to describe their experiences in dementia care by the end of the training. RESULTS: A total of 1264 participants, including 195 facilitators and 1069 learners, completed all assessment were included for analysis. Significant improvements were observed in all outcomes at the 12-month follow-up assessment (Ps ≤ .001). The magnitude of improvements in attitudes was the largest. The findings also showed that the effects of the training program significantly varied across different groups of learners in terms of age, occupation, work and training experience. CONCLUSIONS: This community-wide large-scale project provided evidence that the train-the-trainer model and reflective learning are effective means to facilitate situated learning that promote awareness and understanding of dementia, and consequently enhance sustainability of changes in care practices.

Usefulness of the Bristol Stool Form Chart scoring system for the laboratory processing of faecal samples in suspected Clostridioides difficile cases.

Clostridioides difficile infection (CDI) remains a threat to hospitalized patients. All patients with diarrhoea [defined as Bristol Stool Form Chart (BSFC) types 5-7] not attributable to another cause should be tested for CDI. Samples should quarter fill and take the shape of the specimen container. Use of the BSFC definition of diarrhoea to determine which samples should be tested for CDI in the laboratory was assessed. In this study, two-thirds of glutamate-dehydrogenase-positive, toxin-positive samples were rated BSFC type <5. Therefore, the BSFC should not be used by laboratories to determine which samples are tested for CDI.

Cofactor Selectivity in Methylmalonyl Coenzyme A Mutase, a Model Cobamide-Dependent Enzyme.

Cobamides, a uniquely diverse family of enzyme cofactors related to vitamin B12, are produced exclusively by bacteria and archaea but used in all domains of life. While it is widely accepted that cobamide-dependent organisms require specific cobamides for their metabolism, the biochemical mechanisms that make cobamides functionally distinct are largely unknown. Here, we examine the effects of cobamide structural variation on a model cobamide-dependent enzyme, methylmalonyl coenzyme A (CoA) mutase (MCM). The in vitro binding affinity of MCM for cobamides can be dramatically influenced by small changes in the structure of the lower ligand of the cobamide, and binding selectivity differs between bacterial orthologs of MCM. In contrast, variations in the lower ligand have minor effects on MCM catalysis. Bacterial growth assays demonstrate that cobamide requirements of MCM in vitro largely correlate with in vivo cobamide dependence. This result underscores the importance of enzyme selectivity in the cobamide-dependent physiology of bacteria.IMPORTANCE Cobamides, including vitamin B12, are enzyme cofactors used by organisms in all domains of life. Cobamides are structurally diverse, and microbial growth and metabolism vary based on cobamide structure. Understanding cobamide preference in microorganisms is important given that cobamides are widely used and appear to mediate microbial interactions in host-associated and aquatic environments. Until now, the biochemical basis for cobamide preferences was largely unknown. In this study, we analyzed the effects of the structural diversity of cobamides on a model cobamide-dependent enzyme, methylmalonyl-CoA mutase (MCM). We found that very small changes in cobamide structure could dramatically affect the binding affinity of cobamides to MCM. Strikingly, cobamide-dependent growth of a model bacterium, Sinorhizobium meliloti, largely correlated with the cofactor binding selectivity of S. meliloti MCM, emphasizing the importance of cobamide-dependent enzyme selectivity in bacterial growth and cobamide-mediated microbial interactions.

How to foster informed decision making about food supplements: results from an international Delphi study.

Food supplement use can have beneficial and detrimental effects, making informed decisions about supplement use important. How these decisions are made and which communication strategies can stimulate informed decision making is unclear. This study identified the important characteristics of (i) informed decision making about food supplement use and (ii) important factors indicating how to communicate about food supplements to foster informed decision making. An online three-round Delphi study was conducted. International experts within the field of (risk) communication about food supplements or related fields were recruited via email. The participants' age ranged from 25 to 69 years, and sample sizes for the three rounds were 38, 89 and 51, respectively. Experts indicated that for making an informed decision about food supplement use one needs to have knowledge of their positive and negative effects, the ability to compare these effects, knowing alternatives besides supplements, feeling informed, and feeling able (self-efficacious) to make the decision and making the decision voluntarily. Important communication strategies mentioned were: provision of information about positive and negative effects and the nature of these effects including scientific evidence, ensuring information is easily accessible, well ordered, tailored and provided by a trustworthy, credible and independent source.

Enlarging biopsy-proven fibroadenoma: Is surgical excision necessary?

PURPOSE: Core biopsy-proven fibroadenomas that enlarge on clinical or imaging follow-up are often surgically excised to exclude an associated malignancy. The purpose of this study was to assess how often malignancy is detected upon excision, and to determine whether excision of enlarging, biopsy-proven, uncomplicated fibroadenomas is still warranted. MATERIALS AND METHODS: Review of our institutional pathology database from 2000 to 2010 identified 1117 cases of fibroadenoma, and retrospective chart review, including review of pathology and imaging findings of all these records, was performed. RESULTS: 1117 cases of fibroadenoma were identified in a population of women ranging from ages 17 to 78. Of these, 378 (33.8%) were diagnosed by ultrasound core needle biopsy and formed the study population. Of the 378 cases, 24 (6.3%) had co-existent atypia and were immediately excised; these cases were excluded. An additional 107 (28%) were lost to follow-up. Of the remaining 247 cases, 201 (81%) showed stability on follow-up imaging (mean 31.5 months), and 46 (18.6%) enlarged on follow-up. Of the 46 biopsy proven fibroadenomas that enlarged, 19 had a biopsy at initial presentation and 27 underwent biopsy after they enlarged. Seventeen of the 19 were excised after enlargement, and pathology confirmed fibroadenoma in all cases (100%); two enlarged on initial follow-up imaging but remained stable for at least three years on continued follow-up. Of the 27 cases which were biopsied after enlargement, 23 revealed fibroadenoma on core biopsy, 3 had fibroadenoma with associated atypia with subsequent surgery revealing fibroadenoma and no associated malignancy, and one showed fibroadenoma with smooth muscle with subsequent surgery showing phyllodes tumor. CONCLUSION: Based on this study, enlarging biopsy proven fibroadenomas are not associated with malignancy; therefore, surgical excision does not seem warranted. For presumed enlarging fibroadenomas on imaging, core biopsy should be performed to exclude associated atypia or phyllodes tumor. Finally, surgical excision is indicated for lesions with associated atypia or suspected phyllodes and for symptomatic lesions or cosmetic reasons.

Psychological outcomes of life story work for community-dwelling seniors: A randomised controlled trial.

BACKGROUND AND OBJECTIVE: Life story work has a long tradition in the caring sciences and has been found to serve a number of psychological functions. The effects of life story work on the psychological well-being of community-dwelling older people were examined in this study. DESIGN AND METHODS: For this randomised controlled trial, 244 community-dwelling participants in 17 social centres run by a non-governmental organisation were recruited. The participants were randomly allocated to intervention (n = 124) and control (n = 120) groups. Each member of the intervention group worked with a volunteer to prepare his/her life storybook, while those in the control group participated in a social program. Data were collected at baseline, immediately postintervention, and at three and six months postintervention. The outcomes included measures of life satisfaction, self-esteem, general mental well-being and depressive symptoms. RESULTS: No significant interaction effect was observed between the groups over time, except for an improvement in the general mental well-being of the control group at three months postintervention. CONCLUSIONS: A comparison of the findings in the literature showed that some positive results were reported for LSW studies conducted in nursing homes, whereas in community studies, the results were not always positive. Life story work for seniors in the community did not have the same positive outcomes as previously observed among nursing home residents. It is possible that the intervention had a greater effect on more deprived individuals. Community-dwelling seniors can be encouraged to participate in social activities, which apparently can lead to similar outcomes. IMPLICATIONS FOR PRACTICE: Clinicians should not assume that similar interventions can have similar effects when delivered in a different setting. Community-dwelling seniors can be encouraged to participate in social activities, which can also promote psychological wellbeing similar to to the effects of activities related to life story work.

Uneven distribution of cobamide biosynthesis and dependence in bacteria predicted by comparative genomics.

The vitamin B12 family of cofactors known as cobamides are essential for a variety of microbial metabolisms. We used comparative genomics of 11,000 bacterial species to analyze the extent and distribution of cobamide production and use across bacteria. We find that 86% of bacteria in this data set have at least one of 15 cobamide-dependent enzyme families, but only 37% are predicted to synthesize cobamides de novo. The distribution of cobamide biosynthesis and use vary at the phylum level. While 57% of Actinobacteria are predicted to biosynthesize cobamides, only 0.6% of Bacteroidetes have the complete pathway, yet 96% of species in this phylum have cobamide-dependent enzymes. The form of cobamide produced by the bacteria could be predicted for 58% of cobamide-producing species, based on the presence of signature lower ligand biosynthesis and attachment genes. Our predictions also revealed that 17% of bacteria have partial biosynthetic pathways, yet have the potential to salvage cobamide precursors. Bacteria with a partial cobamide biosynthesis pathway include those in a newly defined, experimentally verified category of bacteria lacking the first step in the biosynthesis pathway. These predictions highlight the importance of cobamide and cobamide precursor salvaging as examples of nutritional dependencies in bacteria.

Evidence for a New Component of High-Energy Solar Gamma-Ray Production.

The observed multi-GeV γ-ray emission from the solar disk-sourced by hadronic cosmic rays interacting with gas and affected by complex magnetic fields-is not understood. Utilizing an improved analysis of the Fermi-LAT data that includes the first resolved imaging of the disk, we find strong evidence that this emission is produced by two separate mechanisms. Between 2010 and 2017 (the rise to and fall from solar maximum), the γ-ray emission was dominated by a polar component. Between 2008 and 2009 (solar minimum) this component remained present, but the total emission was instead dominated by a new equatorial component with a brighter flux and harder spectrum. Most strikingly, although six γ rays above 100 GeV were observed during the 1.4 yr of solar minimum, none were observed during the next 7.8 yr. These features, along with a 30-50 GeV spectral dip which will be discussed in a companion paper, were not anticipated by theory. To understand the underlying physics, Fermi-LAT and HAWC observations of the imminent cycle 25 solar minimum are crucial.