bims-supasi Biomed News
on Sulfation pathways and signalling
Issue of 2025–04–06
sixteen papers selected by
Jonathan Wolf Mueller, University of Birmingham



  1. Carbohydr Polym. 2025 Jun 01. pii: S0144-8617(25)00214-0. [Epub ahead of print]357 123433
      The identification of enzymes involved in biosynthesis of heparan sulfate (HS) and heparin and their successful application in chemoenzymatic synthesis have provided great impetus to rationally design well-defined oligosaccharides as ideal alternatives to animal-derived heterogeneous heparin anticoagulants clinically to treat clotting disorders. Herein, we revisited the substrate specificity of recombinant 2-O-sulfotransferases produced in different expression systems for the highly efficient chemoenzymatic synthesis of HS oligosaccharides containing the rare 2-O-sulfated GlcA (GlcA2S) residues, followed by further assembly into the highly sulfated HS dodecasaccharide (12-mer) and decasaccharide (10-mer) containing the antithrombin-binding domain and the trisulfated disaccharide (GlcA2S-GlcNS6S) units rarely found in natural heparin. The GlcA2S-containing HS 10-mer demonstrated both the effectively reversible anticoagulant activity similar to that of unfractionated heparin and the lower potential risk for life-threatening heparin-induced thrombocytopenia compared with enoxaparin, indicating its promising prospect as the next-generation HS/heparin-like anticoagulant therapeutics.
    Keywords:  2-O-sulfated glucuronic acid; Chemoenzymatic synthesis; Heparan sulfate; Heparin; Heparin-induced thrombocytopenia; Neutralizable anticoagulant activity
    DOI:  https://doi.org/10.1016/j.carbpol.2025.123433
  2. Nat Prod Res. 2025 Mar 31. 1-9
      The extract of Petrosia (Strongylophora) strongylata was separated to trace sulfated terpenoids, which were expected to be present in the crude extract based on its 1H NMR spectrum. As a result, seven sulfated (1-7) sterols and one highly oxygenated steroid (8), including two unreported analogs (4 and 7), were isolated and identified. The MIC measurement of all isolated compounds against six bacterial cell lines demonstrated that compounds 1-7 efficiently inhibit the growth of Gram-positive bacteria. Compound 8, which lacks the sulfate group in the structure, displayed no activity even at a concentration of 128 µM. This study demonstrated that the sulfate groups in the structures of sterols play a critical role in the growth inhibition of Gram-positive bacteria, suggesting the potential of sulfated sterols for use as antimicrobial agents. The modes of action of these compounds and their molecular -targets are currently under investigation.
    Keywords:  Petrosia (Strongylophora) strongylata; Sulfated sterol; halistanol sulfate; ibisterol sulfate
    DOI:  https://doi.org/10.1080/14786419.2025.2484261
  3. Int J Biol Macromol. 2025 Mar 29. pii: S0141-8130(25)03216-7. [Epub ahead of print]308(Pt 3): 142664
      Chondroitin sulfate (CS), a glycosaminoglycan (GAG), plays a pivotal role in various physiological functions and is extensively utilized in medical and clinical applications. This study aimed to explore the enhancing effects and underlying mechanisms of three commonly encountered sulfated glycosaminoglycans CS-A, CS-C and CS-D on α-glucosidase activity. In vitro enzyme kinetic studies demonstrated that all three types of CS promoted α-glucosidase activity, with CS-D exhibiting the most pronounced effect, reaching 124.7 %. Fluorescence and circular dichroism (CD) spectroscopy, along with molecular docking experiments, revealed that CSs spontaneously interacted with the enzyme's active site, forming hydrogen bonds with Arg600 and His674. Additionally, hydrophobic interactions with Trp376 and Trp481 further strengthened these hydrogen bonds. These interactions increased the flexibility of the α-glucosidase polypeptide backbone, leading to greater solvent exposure of Trp residues and alterations in the enzyme's secondary structure composition. Furthermore, trajectory analysis from kinetic simulations indicated that activation of the α-glucosidase active site induced an inward folding and contraction of the region, thereby enlarging the internal cavity and enhancing its hydrophobic nature. This structural reconfiguration not only provided additional space for substrate hydrolysis but also minimized interference from water molecules, collectively contributing to an overall enhancement of α-glucosidase hydrolytic activity. In conclusion, this study identifies CS as an α-glucosidase activator and elucidates its interaction mechanisms through both in vitro and in silico approaches, highlighting its potential applications in the food industry.
    Keywords:  Chondroitin sulfate; Interactions; Α-Glucosidase
    DOI:  https://doi.org/10.1016/j.ijbiomac.2025.142664
  4. J Neuropathol Exp Neurol. 2025 Mar 29. pii: nlaf018. [Epub ahead of print]
      Cerebral amyloid angiopathy (CAA) is a common feature of Alzheimer's disease in which amyloid-β (Aβ) deposits in cerebral and leptomeningeal vessel walls, predisposing vessels to micro- and macro-hemorrhages. The vessel walls contain distinct proteins and heparan sulfate (HS), yet how vascular proteins and HS jointly associate with Aβ is unknown. We conducted the first multi-omics study to systematically characterize the proteins as well as the HS abundance, sulfation level, and disaccharide composition of leptomeninges from 23 moderate to severe CAA cases and controls. We then analyzed the associations between Aβ and other proteins, HS, and apolipoprotein E genotype. We found an increase in a minor HS disaccharide containing unsubstituted glucosamine, as well as 6-O sulfated disaccharides; Aβ40 levels positively correlated with unsubstituted glucosamine. There was also an increase in extracellular proteins derived from brain parenchyma or plasma, including olfactomedin-like protein 3, fibrinogen, serum amyloid protein, apolipoprotein E, and secreted frizzled related protein-3. Our findings of vascular HS and protein alterations specific to CAA-affected leptomeningeal vessels provide molecular insight into the extracellular remodeling that co-occurs with Aβ deposits and may indicate a basis for antemortem diagnostic assay development and therapeutic strategies to impede Aβ-HS interactions.
    Keywords:  Alzheimer's disease; glycomics; heparan sulfate; neurodegeneration; protein misfolding; proteomics
    DOI:  https://doi.org/10.1093/jnen/nlaf018
  5. Carbohydr Polym. 2025 Jun 01. pii: S0144-8617(25)00196-1. [Epub ahead of print]357 123415
      A novel sulfated glycosaminoglycan-like polysaccharide (SIP), with a molecular weight of 13.78 kDa, was isolated from Sepia recurvirostra ink. SIP displayed a sulfate content of 6.82 % and consisted of GlcNAc, GalNAc, GlcA and Fuc. The distinct structural sequence of SIP was identified by 1D/2D NMR assignments of the backbone released by deaminative treatment and the oligosaccharides generated by mild acid hydrolysis. The backbone was built up of disaccharide repeating blocks structured as [-D-GlcA-β1,4-L-Fuc-α1,3-]. Notably, the non-sulfated GalNAc was attached to C-3 of Fuc, while the 4, 6-O-disulfated GlcNAc was pendant at C-4 in 20 % of GlcA residues, as revealed by the well-defined structures of trisaccharide, tetrasaccharides, heptasaccharides and decasaccharides. Except for the trisaccharide, the non-reducing ends of the other oligosaccharides were GlcNAc6S or GlcNAc4S6S, suggesting selectivity of glycosidic bond cleavage and desulfation during mild acid hydrolysis. SIP showed negligible anticoagulant and heparanase inhibitory activity. Moreover, the effects of SIP on the generation of short-chain fatty acids (SCFAs) by colonic microbiota were evaluated. The addition of SIP resulted in a significant increase in levels of acetate, propionate, isobutyric acid, and isovaleric acid, offering insights into the contribution of the colonic microbiota to the health benefits of dietary squid ink.
    Keywords:  Anticoagulant; Glycosaminoglycan; Heparanase; Oligosaccharide structure; SCFA; Squid ink polysaccharide
    DOI:  https://doi.org/10.1016/j.carbpol.2025.123415
  6. Int J Biol Macromol. 2025 Mar 26. pii: S0141-8130(25)03129-0. [Epub ahead of print]308(Pt 3): 142577
      Sulfation at the 4-OH position of galactose or its derivatives is a common and significant modification for polysaccharides. It confers different biological activities to polysaccharides and can be obtained by sulfonate group transfer, which is catalyzed by galactose 4-O-sulfotransferase (G4ST). Consequently, G4STs serves as a valuable tool for investigating the impact of sulfation on biological activity and for the synthesis of novel sulfated sugars. Despite its significance, only a few vertebrate-derived G4STs have been characterized, highlighting the need for exploring G4STs from diverse sources and for understanding of their structure-function relationships. In this study, we attempted for the first time to identify G4STs derived from microbes. Amino acid sequence screening was conducted, followed by protein structure prediction and clustering, resulting in 100 candidate proteins, which were classified into four clades. Notably, we successfully confirmed G4ST activity in one protein from clade1, named BST61. Structural insight into BST61 revealed the key residues involved in substrate binding, and, more importantly, a three residues-surrounded and conserved spatial position functioning in galactose recognition was determined. Using the conserved spatial position as a robust filter to re-screen the 100 candidate proteins, new sulfotransferases towards galactose were discovered as expected. These findings provided precise methods for mining G4STs, greatly expanding the application of G4STs for the analysis and preparation of sulfated sugars.
    Keywords:  Carbohydrate sulfotransferases; Catalytic mechanism; Galactose 4-O-sulfotransferase (G4ST); Structure-based protein clustering; Sulfated polysaccharides; Sulfation modification
    DOI:  https://doi.org/10.1016/j.ijbiomac.2025.142577
  7. FASEB J. 2025 Apr 15. 39(7): e70505
      Elastogenesis is a cell surface-located hierarchical process that requires the core components tropoelastin and fibrillins and several accessory proteins, including fibulin-4 (FBLN4) and latent TGF-β binding protein-4 (LTBP4). FBLN4 and LTBP4 interact with cells, but their cell receptors and associated molecular elastogenic mechanisms remain unknown. Primary skin fibroblasts and several vascular smooth muscle cells bound strongly to FBLN4 multimers and LTBP4 monomers. We identified two cell interaction epitopes on FBLN4 located in cbEGF2-3 and the C-terminal domain, whereas FBLN4 multimerization sites were mapped to cbEGF4-5 and the C-terminal domain. We also determined a novel cell interaction site in the N-terminal half of LTBP4. Cell binding to FBLN4 and LTBP4 was strongly inhibited in the presence of heparin, heparan sulfate, or after enzymatic removal of heparan sulfate, suggesting heparan sulfate proteoglycans as relevant cell surface receptors. siRNA knockdown experiments identified syndecan (SDC)2 and SDC3 as cell receptors for FBNL4 and SDC3 for LTBP4. Direct protein interactions between FBLN4 and the recombinant ectodomains of SDC2 and SDC3, and between LTBP4 and SDC3 validated these results. Interaction of the elastogenic cells with FBLN4 and LTBP4 enhanced elastogenesis, whereas SDC2 and/or SDC3 knockdowns led to reduced elastic fiber formation. The cell interactions with FBLN4 and LTBP4 significantly enhanced focal adhesion formation, induced cell contraction, and led to activation of focal adhesion kinase (FAK), Erk1/2, and RhoA. Pharmacological inhibition of these effectors markedly attenuated elastic fiber formation, and siRNA knockdown of SDC2 and SDC3 led to reduced levels of pFAK, pERK, and active RhoA. Together, these data demonstrate that FBLN4 and LTBP4 cell interactions through SDC2 and SDC3 promote elastogenesis by enhancing focal adhesion formation, leading to cell contractility through FAK, Erk1/2, and RhoA activation, underscoring the significance of these pathways in elastogenesis.
    Keywords:  cell contraction; cell interactions; elastic fibers; fibulin‐4; focal adhesion; latent TGF‐β binding protein‐4; syndecan‐2; syndecan‐3
    DOI:  https://doi.org/10.1096/fj.202402767R
  8. Carbohydr Res. 2025 Mar 28. pii: S0008-6215(25)00099-0. [Epub ahead of print]552 109473
      Sulfated polysaccharide (SP) was successfully extracted from rhodophyta Gracilaria birdiae cultivated in northeast Brazil. For the first time, edible SP films from G. birdiae were formulated by the casting method using an experimental design 32, in which the variables were SP concentration and glycerol concentration. A polysaccharide extraction yield of 26.4 % was obtained through hot aqueous extraction and purification by dialysis. The chemical composition indicated the quality of the SP due to the high total carbohydrate (85.1 %) and sulfate (8.1 %) contents and the absence of proteins. Structural characterization showed the presence, besides the sulfate group, of the methylated monosaccharide residues, namely 3,6-α-l-anhydrogalactose-2-O-methyl and β-d-galactose-6-O-methyl with methylation degree of 0.257. In addition, the polysaccharide chain was mainly composed of 3,6-α-l-anhydrogalactose → β-d-galactose, α-l-galactose-6-sulfate → β-d-galactose, β-d-galactose → 3,6-α-l-anhydrogalactose, and β-d-galactose → α-l-galactose-6-sulfato. SP was also considered thermostable, showing a thermal degradation pattern similar to that of galactans. Through experimental design, it was observed that the film formulation with 2 % SP and 0.2 % glycerol presented the best mechanical, physico-chemical and optical properties for application in food packaging.
    Keywords:  Edible films; Gracilaria birdiae; Structural chemistry; Sulfated polysaccharide
    DOI:  https://doi.org/10.1016/j.carres.2025.109473
  9. Anal Chem. 2025 Mar 29.
      Heparin is a vital macromolecule that regulates blood coagulation, while protamine is an essential polypeptide clinically used to counteract heparin overdose. Detecting both heparin and its antidote protamine under physiological conditions is crucial for biological and clinical applications. This report introduces a cucurbituril[8] (CB[8])-based phosphorescent probe for their detection. The method employs a nanoassembly induced phosphorescence switch-on mechanism for heparin sensing and a disassembly induced phosphorescence switch-off approach for protamine detection. An arginine-rich guest forms a supramolecular complex with heparin, enhancing phosphorescence under secondary confinement and enabling its detection. Conversely, protamine sulfate, as a stronger competitor for heparin, disrupts the probe-heparin aggregates, leading to emission quenching and protamine sensing. This sensor demonstrated high selectivity in detecting both analytes in biological samples, such as human blood serum and urine. The detection limits for heparin and protamine were determined to be 61 and 82 ng/mL in 10% HBS, respectively.
    DOI:  https://doi.org/10.1021/acs.analchem.5c00382
  10. Chembiochem. 2025 Apr 04. e2500024
      Sulfation is a widely used strategy in nature to modify the solubility, polarity, and biological activities of molecules. The enzymes catalyzing sulfation, sulfotransferases (STs), are typically highly specific to a single sulfation site in a molecule. Herein, the identification and characterization of sulfated adipostatins is reported and reveals a novel sulfotransferase, AdpST, which is responsible for di-sulfation at two sites of adipostatins. The initial bioinformatic analysis in search of adipostatin analogs from Streptomyces davaonensis DSM101723 identifies adpST and a 3'-phosphoadenosine-5'-phosphosulfate (PAPS) biosynthetic cassette, which are co-clustered with the adipostatin-encoding type III polyketide synthase. Mono- and di-sulfated adipostatin analogs are discovered in the extracts of S. davaonensis DSM101723, whereas di-sulfated bacterial natural products has not been reported. Using a series of in vivo and in vitro experiments, it is confirmed that AdpST is solely responsible for both mono- and di-sulfation of adipostatins, a catalytic activity which has not been identified in bacterial PAPS-dependent STs to date. It is further demonstrated that the dedicated PAPS biosynthetic cassette improves di-sulfation capacity. Lastly, it is determined that AdpST shares similarity with a small group of uncharacterized STs, suggesting the presence of additional unique bacterial STs in nature, and that AdpST is phylogenetically distant from many characterized STs.
    Keywords:   Streptomyces ; PAPS; adipostatin; di‐sulfation; sulfotransferases
    DOI:  https://doi.org/10.1002/cbic.202500024
  11. Small. 2025 Mar 31. e2500150
      Corneal transplantation remains the prevailing treatment for corneal defects, which is always restricted by donor shortages and numerous postoperative complications accompanying suturing. Photocurable hydrogels have emerged as alternative therapeutic strategies for the repair of corneal defects, but most hydrogels focus on repairing focal corneal defects and still suffer from low transparency and poor mechanical properties. Herein, photocurable hydrogel GelMA/OCS composed of gelatin methacryloyl (GelMA) and oxidized chondroitin sulfate (OCS) is developed for sutureless repair of large corneal defects (6 mm). This injectable hybridized hydrogel demonstrates excellent transparency, low swelling rate, enhanced mechanical properties, and superior adhesion properties. In vitro experiments reveal that GelMA/OCS hydrogel can support the proliferation and migration, and adhesion growth of human corneal epithelial cells (HCECs), demonstrating satisfactory cytocompatibility and cell affinity. In addition, GelMA/OCS hydrogel is capable of accurately filling the large corneal defects in rabbits and forming hydrogel grafts with smooth surfaces. Postoperative slit lamp, histological evaluation, and transcriptomic analysis reveal that GelMA/OCS hydrogel can significantly facilitate corneal re-epithelialization and the integration and reconstruction of stromal structures, as well as reduce inflammation responses and scar formation. Therefore, GelMA/OCS hydrogel may provide a promising alternative for the sutureless treatment of large corneal defects.
    Keywords:  dual network; large corneal defects; photocurable hydrogels; sutureless repair
    DOI:  https://doi.org/10.1002/smll.202500150
  12. Theriogenology. 2025 Mar 18. pii: S0093-691X(25)00122-0. [Epub ahead of print]240 117396
      The present study investigated the effects of treatments with sulfatase to desulfate the sulfated carbohydrate chain residues of the zona pellucida (ZP) and arylsulfatase A-blocking peptide (ARSA-BP) for the specific masking of sperm ARSA on the interaction of sperm ARSA with ZP, which plays an important role in the in vitro fertilization (IVF) of porcine oocytes. The pretreatment of oocytes with sulfatase for 10 min before the onset of insemination or the addition of 5 ng/mL ARSA-BP to IVF medium significantly reduced the incidence of polyspermy; however, these treatments did not affect sperm penetrability or male pronucleus formation. Combined treatments with sulfatase plus ARSA-BP significantly reduced the incidence of polyspermy to the range of 23-26 % under conditions maintaining a high sperm penetration rate from that of untreated oocytes (56 %). Although the treatments with sulfatase or ARSA-BP alone and sulfatase plus ARSA-BP during IVF did not affect the number of sperm that bound to the ZP or ZP solubility, the incidence of sperm with the acrosome reaction that bound to the ZP was efficiently reduced by the treatment. In oocytes treated with ARSA-BP but not sulfatase, developmental potency to the blastocyst stage was enhanced by decreasing the number of apoptotic blastomeres. Therefore, the present results clearly indicate that interference with the interaction between sperm ARSA and sulfated ZP glycoprotein residues reduced the induction of the acrosome reaction in sperm that bound to the ZP, effectively suppressing polyspermy during porcine IVF.
    Keywords:  Acrosome reaction; Arylsulfatase A; In vitro fertilization; Polyspermy; Porcine oocytes; Sulfatase
    DOI:  https://doi.org/10.1016/j.theriogenology.2025.117396
  13. J Med Chem. 2025 Apr 02.
      Phenol is produced by β-elimination of l-tyrosine (Tyr) catalyzed by tyrosine phenol-lyase (TPL) during intestinal bacterial metabolism. Phenol and its conjugate, phenyl sulfate (PhS), are protein-bound uremic toxins (PBUTs). Elevated levels of phenol and PhS are strongly implicated in the etiology and outcomes of uremia. Because hemodialysis is insufficient in removing phenol and PhS, novel methods are necessary for inhibiting phenol production during bacterial metabolism. We explored TPL inhibitors and found that dietary polyphenols, particularly gallic acid (GA), strongly inhibited TPL-catalyzed phenol production. A GA derivative, 3,5-dihydroxybenzoic acid (3,5DHBA), competitively inhibited TPL and significantly decreased phenol levels in TPL-expressing bacteria (Morganella morganii and Citrobacter koseri) and Tyr-rich-diet-fed C57BL/6J mouse feces. Our findings suggested that 3,5DHBA was the most promising polyphenol in decreasing phenol levels. Therefore, dietary intake of 3,5DHBA or its phenolic precursors may be useful in minimizing PBUT levels by targeting intestinal bacteria.
    DOI:  https://doi.org/10.1021/acs.jmedchem.5c00418
  14. Kidney360. 2025 Apr 01.
       BACKGROUND: The intricate interplay between chronic kidney disease (CKD) and intestinal microbiota has gained increasing attention, with gut dysbiosis being implicated in uremic toxin accumulation and CKD progression. Sodium-glucose cotransporter 2 (SGLT2) inhibitors are now transforming CKD management but pose uncertain effects on shaping gut microbiota. This study aimed to elucidate the impact of SGLT2 inhibitors on perturbations of gut microbial composition and metabolic responses in CKD patients.
    METHODS: Analysis of fecal microbiota and targeted profiling of serum short-chain fatty acids (SCFAs) and gut-derived uremic toxins were conducted in a matched case-control study, including 60 CKD patients (treated: n=30; untreated: n=30) and 30 non-CKD controls.
    RESULTS: Gut microbial composition differed significantly among three study groups. CKD patients receiving SGLT2 inhibitors exhibited distinctive taxonomic profiles, such as enrichment of Bacteroides stercoris and Bacteroides coprocola. Surveys of metabolomic profiles revealed a reduction of two uremic solutes, indoxyl sulfate (IS) and p-cresyl sulfate (pCS), and several SCFAs (formic, acetic, propionic, valeric, and 2-methylbutanoic acid) in SGLT2 inhibitor-treated CKD patients. Co-occurrence analysis demonstrated a set of intestinal microbes that is positively or negatively correlated with the levels of pCS, and the abundance of these pCS-associated intestinal microorganisms was correlated with the levels of IS and isovaleric acids in the same and opposite direction, respectively. Further functional prediction indicated attenuated pathways related to protein and carbohydrate metabolism.
    CONCLUSIONS: Treatment with SGLT2 inhibitors in CKD patients is associated with distinct gut microbial composition and metabolite profiles, suggesting potential modulation of gut dysbiosis and metabolic pathways. Further studies are warranted to elucidate the clinical implications of these findings in CKD management.
    DOI:  https://doi.org/10.34067/KID.0000000792
  15. Handb Exp Pharmacol. 2025 Apr 02.
      Glycosaminoglycans (GAGs), linear anionic periodic polysaccharides, play pivotal roles in various biologically relevant processes within the extracellular matrix (ECM). These processes encompass cell development, proliferation, signaling, ECM assembly, coagulation, and angiogenesis. GAGs perform their functions through their interactions with specific protein partners, rendering them attractive targets for regenerative medicine and drug design. However, the molecular mechanisms governing protein-GAG interactions remain unclear. Classical structure determination techniques face significant challenges when dealing with protein-GAG complexes. This is due to GAGs' unique properties, including their extensive length, flexibility, periodicity, symmetry, multipose binding, and the high heterogeneity of their sulfation patterns constituting the "sulfation code." Consequently, only a limited number of experimental protein-GAG structures have been elucidated. Hence, theoretical approaches are particularly promising in deciphering the code for understanding the structure-function relationship of these complex molecules. In this chapter, we focus on the particularities, challenges, and advances of computational methods such as molecular docking, molecular dynamics, and free-energy calculations when applied to GAG-containing systems. These computational approaches offer valuable insights into the enigmatic world of protein-GAG interactions, paving the way for their enhanced understanding and potential therapeutic applications.
    DOI:  https://doi.org/10.1007/164_2025_741
  16. Allergol Int. 2025 Mar 29. pii: S1323-8930(25)00003-6. [Epub ahead of print]
       BACKGROUND: Elderly asthma has distinct pathophysiologic and phenotypic characteristics compared with asthma in younger patients. However, a potential relationship between sex hormones and the severity of asthma remains unknown in the elderly population. The aim of the present study was to elucidate the relationship between the level of circulating free testosterone and severity of asthma among Japanese with elderly asthma.
    METHODS: The level of free testosterone was measured using sera from elderly patients with asthma aged ≥60 years (n = 192), and its association with the severity of asthma was examined after stratification by sex.
    RESULTS: Based on previous literature and our preliminary analysis showing that current oral corticosteroid (OCS) use might be a risk factor for a lower free testosterone level regardless of severity of asthma, analyzed patients were limited to those who were not currently using OCS (n = 164). Regarding elderly men who were not currently using OCS (n = 62), there was no significant association between free testosterone level and severity of asthma. However, in female counterparts (n = 102), a low free testosterone level was significantly associated with severe asthma even after adjustment for age (p for trend, 0.03).
    CONCLUSIONS: The present study showed a significant association between the serum free testosterone level and severity of asthma among elderly women who were not currently using OCS. Although the causal relationship is unclear, this finding may provide a clue to understand the sex difference in the mechanisms of severe asthma in elderly populations.
    Keywords:  Dehydroepiandrosterone sulfate; Elderly; Free testosterone; Oral corticosteroids; Severe asthma
    DOI:  https://doi.org/10.1016/j.alit.2024.12.008