Affinage

Showing STAR — STARD1 is a alias.

STAR

Guanylyl cyclase C · UniProt P25092

Length
1073 aa
Mass
123.4 kDa
Annotated
2026-06-10
100 papers in source corpus 21 papers cited in narrative 21 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 9/9 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

STAR (STARD1) mediates the rate-limiting step of steroidogenesis by transferring cholesterol from the outer to the inner mitochondrial membrane, where P450scc converts it to pregnenolone (PMID:11181954). It is a sterol-selective transfer protein: recombinant StAR moves cholesterol and beta-sitosterol—but not phosphatidylcholine—from liposomes to mitochondria, and a non-functional mutant fails to do so (PMID:9756854). StAR acts on the cytosolic face of the outer mitochondrial membrane, where interaction with protonated phospholipid head groups drives a pH-dependent molten-globule conformational transition; the partially unfolded C-terminal helix opens a hydrophobic cavity (residues ~83–140) that binds cholesterol (PMID:17433772, PMID:18331352, PMID:19138724). Import into the mitochondrial matrix terminates StAR activity and targets it for turnover by matrix proteases and the proteasome, so activity is intrinsically coupled to its residence at the outer membrane (PMID:11181954, PMID:17218054). StAR function is gated by phosphorylation: PKA phosphorylates S194/195 downstream of ACTH/cAMP signaling to maximize steroidogenic activity, while ERK1/2 phosphorylation and mitochondrial fusion control StAR retention at the outer membrane (PMID:10548884, PMID:25540920, PMID:27999527). Cholesterol delivery is organized within an outer-membrane complex involving the peripheral benzodiazepine receptor (PBR/TSPO), with PAP7 bridging PKA (RIα) to the PBR-StAR machinery (PMID:17433772, PMID:12530641). STAR transcription is controlled by SF-1/NR5A acting at the proximal and distal promoter elements and by cAMP-driven CREB/TORC signaling, and is directly repressed by HIF-1 under hypoxia, while the H19/let-7 axis tunes StAR post-transcriptionally (PMID:9029708, PMID:8923870, PMID:17210223, PMID:30400066, PMID:27813675). Beyond steroidogenesis, STARD1 governs mitochondrial cholesterol homeostasis in hepatocytes: it is induced by ER stress to drive mitochondrial cholesterol loading and GSH depletion in acetaminophen-induced liver failure (PMID:31029706), supplies cholesterol for primary bile acid synthesis via the alternative pathway and thereby promotes NASH-driven hepatocarcinogenesis (PMID:33515644), and also influences hepatic lipid metabolism and diacylglycerol accumulation (PMID:33670702, PMID:28153708). Genetic analysis of congenital lipoid adrenal hyperplasia mutations and the StAR-null mouse established its essential role in steroid hormone biosynthesis (PMID:11181954).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1996 High

    Established that StAR expression is concentrated in the most steroidogenic cells and is controlled at the transcriptional level by cAMP, defining the regulatory logic of acute steroidogenesis.

    Evidence In situ hybridization, nuclear run-on and reporter assays in primary human ovarian/granulosa-lutein cells

    PMID:8923870

    Open questions at the time
    • Did not identify the transcription factors mediating cAMP responsiveness
    • Did not address protein-level mechanism of cholesterol transfer
  2. 1997 High

    Identified SF-1 as the transcription factor required for StAR promoter activity, providing the molecular link between steroidogenic cell identity and StAR expression.

    Evidence Promoter-luciferase reporter assays, SF-1 cotransfection rescue in BeWo cells, cis-element deletion/mutation and binding assays

    PMID:9029708

    Open questions at the time
    • Did not resolve how cAMP signaling converges on the promoter
    • Other promoter regulators not characterized
  3. 1998 High

    Demonstrated directly that StAR is a sterol-selective transfer protein, answering whether StAR itself moves cholesterol rather than merely regulating another carrier.

    Evidence In vitro sterol transfer assay with recombinant StAR, liposomes, heat-treated mitochondria, with a non-functional mutant control and phosphatidylcholine selectivity test

    PMID:9756854

    Open questions at the time
    • Did not define the membrane on which StAR acts in cells
    • Structural basis of cholesterol selectivity not resolved
  4. 1999 High

    Showed PKA phosphorylation at S194/195 maximizes StAR activity and that mitochondrial import inactivates it, establishing that StAR acts before/at the outer membrane and is post-translationally gated.

    Evidence Truncation and site-directed mutagenesis with functional steroidogenesis and phosphorylation assays

    PMID:10548884

    Open questions at the time
    • Mechanism by which phosphorylation enhances activity not defined
    • Did not establish the structural change underlying activity
  5. 2001 High

    Synthesized genetic and biochemical evidence to fix the canonical model: StAR mediates the rate-limiting outer-to-inner membrane cholesterol transfer feeding P450scc, validated by congenital lipoid adrenal hyperplasia and the null mouse.

    Evidence Congenital lipoid adrenal hyperplasia mutation analysis, StAR-null mouse phenotype, COS-1 complementation, recombinant protein on isolated mitochondria

    PMID:11181954

    Open questions at the time
    • Molecular machinery transferring cholesterol across the intermembrane space not fully defined
    • Outer-membrane partner proteins not yet identified
  6. 2002 Medium

    Placed StAR within an outer-membrane signaling-transport complex by showing PAP7 bridges PKA (RIα) to PBR, coupling hormone signaling to cholesterol import.

    Evidence Antisense knockdown, co-IP/interaction studies, overexpression and steroid assays in MA-10 Leydig cells

    PMID:12530641

    Open questions at the time
    • Direct StAR-PBR physical contact not biochemically resolved
    • Single lab, limited mechanistic resolution
  7. 2007 Medium

    Defined the conformational mechanism of StAR action—a pH/phospholipid-induced molten-globule transition at the outer membrane required for cholesterol transfer, with a proposed StAR-PBR cholesterol handoff.

    Evidence Biophysical spectroscopy, molecular dynamics, structure-function studies in synthetic and natural membranes (two complementary studies)

    PMID:17207924 PMID:17433772

    Open questions at the time
    • Not reconstituted as a complete biochemical pathway
    • Direct cholesterol transfer from PBR to inner membrane not demonstrated
  8. 2007 Medium

    Resolved StAR turnover dynamics, showing the cytoplasmic preprotein partitions between outer-membrane activity and proteasomal/matrix-protease degradation, mechanistically explaining why import inactivates StAR.

    Evidence Proteasome inhibitor treatment, immuno-EM, pulse-chase turnover, mitochondrial fractionation

    PMID:17218054

    Open questions at the time
    • E3 ligase / degradation signal not identified
    • Quantitative flux between active and degraded pools not defined
  9. 2007 Medium

    Connected cAMP signaling to StAR transcription beyond SF-1 by showing the CREB coactivator TORC is activated by PKA-driven dephosphorylation to promote StAR expression.

    Evidence Pharmacological inhibition, reporter assays and phosphorylation analysis in Y1 adrenocortical cells

    PMID:17210223

    Open questions at the time
    • TORC binding to the StAR promoter not directly mapped
    • Single lab pharmacological approach
  10. 2008 Medium

    Localized the cholesterol-binding region of the START domain to residues 83–140 and revealed differential binding behavior versus STARD3, refining the structural basis of sterol recognition.

    Evidence Photoaffinity labeling with [3H]azocholestanol, peptide mapping, trypsin protection assays

    PMID:18331352

    Open questions at the time
    • Atomic-resolution cholesterol-bound structure not solved
    • Functional consequence of differential trypsin protection unclear
  11. 2009 Low

    Provided solution-state structural evidence for a two-state gating model in which C-terminal helix unfolding permits cholesterol entry followed by refolding.

    Evidence Solution-state NMR (1H-15N-HSQC), homology modeling, structure-based thermodynamics

    PMID:19138724

    Open questions at the time
    • Full structure not solved and no mutagenesis validation in this study
    • Single lab
  12. 2014 Medium

    Extended phosphoregulation of StAR by showing ERK1/2 phosphorylation and mitochondrial fusion control StAR retention on the outer membrane and thereby its activity.

    Evidence Phosphorylation assays, kinase inhibitor experiments, mitochondrial fractionation and steroidogenesis assays

    PMID:25540920

    Open questions at the time
    • ERK phosphosite(s) not precisely mapped here
    • Mechanism linking fusion to retention unresolved
  13. 2017 Medium

    Identified post-transcriptional control of StAR by the H19/let-7 axis, adding a non-coding RNA layer to StAR regulation.

    Evidence H19 and let-7 overexpression with reporter assays in murine and human cell lines

    PMID:27813675

    Open questions at the time
    • Physiological contexts where this axis dominates not defined
    • Single lab
  14. 2017 Medium

    Revealed a hepatic metabolic role: StAR overexpression reduces hepatic lipid accumulation and insulin resistance via FXR activation and reduced DAG/PKCε signaling, extending StAR function beyond steroidogenesis.

    Evidence Adenoviral StAR overexpression in HFD mice and FFA-overloaded hepatocytes with FXR inhibition rescue

    PMID:28153708

    Open questions at the time
    • Mechanism linking StAR cholesterol transfer to FXR ligand generation not fully defined
    • Gain-of-function only
  15. 2019 High

    Established STARD1 as a driver of hepatotoxicity, where ER-stress-induced STARD1 loads mitochondrial cholesterol and depletes GSH upstream of SAB/JNK in acetaminophen-induced liver failure.

    Evidence Liver-specific conditional KO of Stard1, Sab and Jnk1/2 with epistasis, humanized liver mouse model, mitochondrial assays

    PMID:31029706

    Open questions at the time
    • Direct mechanism of mitochondrial GSH depletion by cholesterol not fully resolved
    • How ER stress induces STARD1 transcription not defined here
  16. 2019 High

    Demonstrated direct HIF-1 binding and repression of the STAR promoter, providing a hypoxia-responsive brake on cholesterol transport and testosterone synthesis distinct from effects on other steroidogenic enzymes.

    Evidence ChIP, EMSA supershift, reporter assays with binding-site mutation, in vivo and in vitro hypoxia in Leydig/TM3 cells

    PMID:30400066

    Open questions at the time
    • Interplay between HIF-1 repression and SF-1/CREB activation not integrated
    • Physiological hypoxia thresholds not defined
  17. 2021 High

    Linked STARD1-dependent cholesterol transfer to primary bile acid synthesis via the alternative mitochondrial pathway and to NASH-driven hepatocellular carcinoma through bile-acid effects on tumor-initiating cells.

    Evidence Hepatocyte-specific Stard1 deletion and overexpression in NASH-HCC mouse models, bile acid mass spectrometry, TIC assays

    PMID:33515644

    Open questions at the time
    • Receptor mediating bile-acid effects on stemness not defined
    • Translation to human HCC not established
  18. 2021 Medium

    Confirmed STAR requirement for steroidogenesis in a clean knockout and uncovered a cholesterol-transport-independent role in lipid droplet diacylglycerol accumulation.

    Evidence CRISPR/Cas9 STAR knockout in MA-10 cells with steroidogenesis assays, EM, lipid-droplet LC-MS lipidomics and transcriptomics

    PMID:33670702

    Open questions at the time
    • Mechanism of StAR influence on DAG metabolism unknown
    • Whether DAG role is direct or secondary unresolved
  19. 2021 Medium

    Identified acid ceramidase as an inverse regulator of STARD1 expression in Niemann-Pick C disease, connecting STARD1-driven mitochondrial cholesterol loading to lysosomal storage pathology via LRH-1.

    Evidence U18666A treatment in Stard1 mice, ACDase transfection in NPC patient fibroblasts, mitochondrial cholesterol/GSH and function assays

    PMID:34175669

    Open questions at the time
    • Direct transcriptional mechanism via LRH-1 not fully mapped
    • Single lab across mixed model systems

Open questions

Synthesis pass · forward-looking unresolved questions
  • How StAR physically delivers cholesterol across the intermembrane space and the atomic structure of the cholesterol-loaded, membrane-engaged state remain unresolved, as does the molecular basis of its non-steroidogenic lipid metabolic roles.
  • No full atomic structure of the membrane-engaged holo state
  • Mechanism of intermembrane cholesterol handoff not reconstituted
  • Mechanistic basis of DAG/lipid-droplet roles unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008289 lipid binding 4 GO:0140104 molecular carrier activity 2
Localization
GO:0005739 mitochondrion 3 GO:0005811 lipid droplet 1
Pathway
R-HSA-162582 Signal Transduction 4 R-HSA-1430728 Metabolism 3 R-HSA-74160 Gene expression (Transcription) 3
Partners
PAP7PRKAR1ATSPO

Evidence

Reading pass · 21 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2001 StAR protein mediates the rate-limiting step in steroidogenesis by transferring cholesterol from the outer to the inner mitochondrial membrane, where the cholesterol side-chain cleavage enzyme (P450scc) converts it to pregnenolone. StAR acts exclusively on the outer mitochondrial membrane, and its import into mitochondria is not essential for activity but rather represents a means of inactivating it. Genetic analysis of congenital lipoid adrenal hyperplasia mutations, StAR null mouse phenotype, functional complementation in COS-1 cells, recombinant protein studies on isolated mitochondria Annual review of physiology High 11181954
1998 Recombinant StAR protein (lacking the N-terminal 62 aa mitochondrial targeting sequence) stimulates transfer of cholesterol and beta-sitosterol from liposomes to heat-treated mitochondria in a dose-, time-, and temperature-dependent manner, demonstrating sterol transfer activity. A non-functional StAR mutant did not promote sterol transfer. Unlike SCP2, StAR did not stimulate phosphatidylcholine transfer, indicating sterol selectivity. In vitro sterol transfer assay using recombinant proteins, liposomes, and heat-treated mitochondria; site-directed mutagenesis of non-functional StAR The Journal of biological chemistry High 9756854
1999 StAR is a target for serine phosphorylation by protein kinase A (PKA) at S194/195 (mouse/human), and this phosphorylation is required for maximizing StAR steroidogenic activity. Import into mitochondria inactivates StAR; the C-terminus contains the functionally important domains for steroidogenic activity. Truncation mutations, site-directed mutagenesis, functional assays in steroidogenic cells, phosphorylation studies Recent progress in hormone research High 10548884
1997 Steroidogenic factor 1 (SF-1) is required for StAR gene transcription. The human StAR promoter contains two cis elements (distal and proximal) governing basal and cAMP-regulated expression; the distal element is a high-affinity SF-1 binding site. Cotransfection of SF-1 allows StAR promoter function in BeWo cells that normally lack activity; deletion or mutation of either element substantially reduces SF-1-supported activity. Promoter-luciferase reporter assays, cotransfection with SF-1 expression plasmid, deletion and site-directed mutation of cis elements, gel shift / binding assays Steroids High 9029708
1996 StAR mRNA and protein are expressed in the most steroidogenic compartments of the human ovary (theca of preovulatory follicles, luteinized granulosa and thecal cells). cAMP analog 8-Br-cAMP increases StAR mRNA by increasing StAR gene transcription (not mRNA stability). Phorbol myristate acetate (protein kinase C activator) antagonizes the cAMP stimulatory effect on StAR expression. In situ hybridization, Northern blot, nuclear run-on transcription assays, mRNA stability assays, promoter-luciferase transfection in granulosa-lutein cells The Journal of clinical endocrinology and metabolism High 8923870
2007 StAR's interaction with protonated phospholipid head groups on the outer mitochondrial membrane induces a molten globule conformational change required for its cholesterol transfer activity. StAR requires cholesterol binding and acts on the outer membrane. A model is proposed wherein StAR removes cholesterol from the cholesterol-binding domain of the peripheral benzodiazepine receptor (PBR) and delivers it to the inner mitochondrial membrane. Biophysical spectroscopy, molecular dynamics simulations, structure-function studies in synthetic and natural membranes Biochimica et biophysica acta Medium 17433772
2007 StAR acts on the outer mitochondrial membrane, requires cholesterol binding, and requires a pH-dependent molten globule structural change for function. Functional interaction between StAR and PBR (peripheral benzodiazepine receptor) is indicated. Structure-function analysis, biophysical studies, outer membrane activity assays Molecular and cellular endocrinology Medium 17207924
2002 PBR (peripheral benzodiazepine receptor), StAR, and PKA cooperate in hormone-induced mitochondrial cholesterol transport. PAP7, a protein that interacts with both PBR and the PKA regulatory subunit RIα, is present in mitochondria of adrenal and gonadal cells; overexpression of PAP7 increases hormone-induced steroid production, and inhibition of PAP7 reduces it, indicating PAP7 bridges PKA to PBR-StAR complex. Antisense oligonucleotide knockdown, co-immunoprecipitation/protein interaction studies, overexpression, steroid production assays in MA-10 Leydig cells Endocrine research Medium 12530641
2014 ERK1/2 phosphorylates StAR, and this phosphorylation regulates StAR retention on the outer mitochondrial membrane (OMM) and StAR activity. Mitochondrial fusion also plays a role in regulating StAR retention on the OMM, thereby modulating its steroidogenic activity. Phosphorylation assays, kinase inhibitor experiments, mitochondrial fractionation, functional steroidogenesis assays Molecular and cellular endocrinology Medium 25540920
2007 Once synthesized on free polyribosomes, StAR preprotein either associates with the outer mitochondrial membrane to mediate cholesterol transfer, or is degraded by the proteasome. Upon mitochondrial import, StAR is subject to rapid turnover: one pool is degraded by matrix proteases shortly after import; a second pool undergoes slower degradation after translocation to the matrix face of the inner membrane. Proteasome inhibitors (MG132, clasto-lactacystin beta-lactone but not epoxomicin) can inhibit turnover of both cytoplasmic preprotein and intra-mitochondrial StAR. Proteasome inhibitor treatment, immuno-electron microscopy, pulse-chase protein turnover assays, mitochondrial fractionation Molecular and cellular endocrinology Medium 17218054
2008 Photoaffinity labeling with [(3)H]azocholestanol predominantly labeled a 6.2 kDa fragment (amino acids 83-140) of STARD1-START, which contains residues proposed to interact with cholesterol in a hydrophobic cavity. Cholesterol preferentially interacts with one side wall of this cavity. By contrast, cholesterol had no protective effect against trypsin degradation of STARD1-START (unlike STARD3), suggesting differential cholesterol-binding properties between the two START domains. Photoaffinity labeling with radiolabeled azocholestanol, trypsin protection assays, peptide mapping The FEBS journal Medium 18331352
2008 NMR studies of StAR in apo- and holo-states at physiological pH show well-dispersed resonances with key spectral differences between states, consistent with a two-state model in which the C-terminal alpha-helix undergoes partial unfolding (molten globule transition) to allow cholesterol binding, followed by stabilization and refolding upon cholesterol binding. This structural gating mechanism is proposed for cholesterol access. Solution-state NMR ((1)H-(15)N-HSQC), homology modeling, structure-based thermodynamics Molecular and cellular endocrinology Low 19138724
2007 TORC (transducer of regulated CREB activity), a CREB coactivator, regulates StAR gene expression: dephosphorylated TORC is active and promotes StAR transcription. PKA phosphorylates CREB and simultaneously inhibits TORC kinases, leading to TORC dephosphorylation and activation. Staurosporine (kinase inhibitor) increases dephospho-TORC and induces StAR expression in Y1 adrenocortical cells. Pharmacological inhibition (staurosporine, KG501), reporter gene assays, phosphorylation analysis, expression studies in Y1 adrenocortical cells Molecular and cellular endocrinology Medium 17210223
2019 Endoplasmic reticulum (ER) stress induces STARD1 upregulation in hepatocytes, which mediates mitochondrial cholesterol accumulation, sustained mitochondrial GSH depletion, and mitochondrial dysfunction leading to acetaminophen-induced acute liver failure. Liver-specific STARD1 deletion (Stard1ΔHep) protected mice from APAP/VPA-induced ALF despite increased mitochondrial GSH and phosphorylated JNK. STARD1 acts upstream of SAB (SH3BP5) and JNK1/2 phosphorylation in this hepatotoxicity pathway. Liver-specific conditional knockout mice (Stard1ΔHep, SabΔHep, Jnk1+2ΔHep), pharmacological ER stress inhibition (TUDCA), histology, mitochondrial function assays, humanized liver mouse model (FRGN) Gastroenterology High 31029706
2021 STARD1 promotes generation of primary bile acids (β-muricholic acid, cholic acid and their tauroconjugates) through the alternative mitochondrial pathway in hepatocytes. STARD1 overexpression in NASH mouse models increases liver tumor multiplicity, while hepatocyte-specific STARD1 deletion reduces tumor burden. The STARD1-generated bile acids act on tumor-initiated stem-like cells (TICs) to stimulate stemness, pluripotency and inflammation, linking STARD1 to HCC pathogenesis. Hepatocyte-specific STARD1 deletion (Stard1ΔHep) and overexpression in NASH-HCC mouse models, bile acid profiling by mass spectrometry, TIC and primary hepatocyte incubation assays Journal of hepatology High 33515644
2019 HIF-1 (hypoxia-inducible factor 1) directly binds to the StAR/STAR promoter at three specific binding sites (-2082/-2078, -2064/-2060, -1910/-1906) and represses STAR transcription, leading to reduced cholesterol transport and decreased testosterone synthesis in Leydig cells. This was shown to specifically affect cholesterol transport (blocked by pregnenolone rescue but not cAMP), while other steroidogenic enzymes (3b-HSD, 17b-HSD, P450scc) were not significantly affected. ChIP, EMSA supershift, dual-luciferase reporter assay, site-directed mutation of HIF-1 binding sites, hypoxia exposure in vivo (mice) and in vitro (rat primary Leydig cells, TM3 cells) Journal of molecular endocrinology High 30400066
2017 StAR is a novel target of the microRNA let-7, which inhibits StAR at the post-transcriptional level. The long noncoding RNA H19 stimulates StAR expression by sponging/antagonizing let-7, thereby relieving let-7-mediated repression of StAR. Overexpression of H19 and let-7, reporter assays, murine and human cell line experiments Endocrinology Medium 27813675
2021 STAR knockout in MA-10 mouse Leydig cells abolishes progesterone formation in response to dibutyryl-cAMP and TSPO drug ligands (but not to the membrane-permeable 22(R)-hydroxycholesterol). STAR KO cells show significantly altered lipid droplet density and composition, with marked increases in diacylglycerol (DAG, particularly DAG 38:1), cholesteryl ester, and phosphatidylcholine in lipid droplets, suggesting constitutive STAR has a role in DAG accumulation in lipid droplets beyond cholesterol transport. CRISPR/Cas9 STAR knockout in MA-10 cells, steroid production assays, electron microscopy, liquid chromatography-mass spectrometry of lipid droplet content, transcriptomic analysis International journal of molecular sciences Medium 33670702
2021 Acid ceramidase (ACDase) inversely regulates STARD1 expression: reduced ACDase in NPC disease correlates with increased STARD1 and mitochondrial cholesterol. Transfection of ACDase in NPC patient fibroblasts decreased STARD1 expression and mitochondrial cholesterol accumulation, resulting in increased mitochondrial GSH, improved mitochondrial function, and decreased oxidative stress. The STARD1 upregulation in NPC is dissociated from ER stress and linked to LRH-1 levels. U18666A treatment in Stard1f/f and Stard1ΔHep mice, ACDase transfection in NPC patient fibroblasts, mitochondrial cholesterol measurement, GSH assay, mitochondrial functional assays Redox biology Medium 34175669
2017 StAR overexpression in a high-fat diet NAFLD mouse model reduced hepatic lipid accumulation and attenuated insulin resistance through activation of the farnesoid X receptor (FXR) and reduction of intracellular diacylglycerol levels with consequent decreased PKCε phosphorylation. FXR inactivation reversed these beneficial effects of StAR overexpression. Recombinant adenovirus-mediated StAR overexpression in HFD mice and FFA-overloaded hepatocytes, lipid measurement, insulin signaling assays, FXR inhibition experiments Biochimica et biophysica acta. Molecular basis of disease Medium 28153708
2016 ACTH activates StAR through the cAMP-PKA signaling pathway. PKA-dependent phosphorylation of StAR at S194/195 (mouse/human) is required for StAR function. The current model places StAR translation and phosphorylation at the outer mitochondrial membrane as the site of StAR action. Review/synthesis of mutagenesis and phosphorylation studies; the mechanistic conclusions are grounded in cited primary experimental work on PKA phosphorylation and site-directed mutants Frontiers in neuroscience Medium 27999527

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2012 STAR: ultrafast universal RNA-seq aligner. Bioinformatics (Oxford, England) 38488 23104886
2015 Circular RNA: A new star of noncoding RNAs. Cancer letters 1436 26052092
2015 Mapping RNA-seq Reads with STAR. Current protocols in bioinformatics 959 26334920
2001 StAR protein and the regulation of steroid hormone biosynthesis. Annual review of physiology 694 11181954
2007 The pancreatic stellate cell: a star on the rise in pancreatic diseases. The Journal of clinical investigation 582 17200706
2019 CircRNA: a rising star in gastric cancer. Cellular and molecular life sciences : CMLS 326 31659415
2021 Superoxide dismutase nanozymes: an emerging star for anti-oxidation. Journal of materials chemistry. B 295 34161407
2013 MiR-200, a new star miRNA in human cancer. Cancer letters 289 24262661
2021 Exosomes, a New Star for Targeted Delivery. Frontiers in cell and developmental biology 280 34692704
2020 Extracellular vesicles: A bright star of nanomedicine. Biomaterials 275 33189359
2007 Steroidogenic acute regulatory protein (StAR), a novel mitochondrial cholesterol transporter. Biochimica et biophysica acta 241 17433772
2005 Target RNA motif and target mRNAs of the Quaking STAR protein. Nature structural & molecular biology 239 16041388
1996 Role of the steroidogenic acute regulatory protein (StAR) in steroidogenesis. Biochemical pharmacology 224 8573184
2000 The role of the StAR protein in steroidogenesis: challenges for the future. The Journal of endocrinology 219 10694364
2023 Extracellular vesicles: a rising star for therapeutics and drug delivery. Journal of nanobiotechnology 216 37475025
2009 Transcriptional regulation of steroidogenic genes: STARD1, CYP11A1 and HSD3B. Experimental biology and medicine (Maywood, N.J.) 215 19491374
2015 PVT1: a rising star among oncogenic long noncoding RNAs. BioMed research international 192 25883951
1998 Steroidogenic acute regulatory protein (StAR) is a sterol transfer protein. The Journal of biological chemistry 180 9756854
2016 Optimizing RNA-Seq Mapping with STAR. Methods in molecular biology (Clifton, N.J.) 171 27115637
2000 Steroidogenic acute regulatory protein (StAR) and the intramitochondrial translocation of cholesterol. Biochimica et biophysica acta 155 11111087
2001 Tracking the role of a star in the sky of the new millennium. Molecular endocrinology (Baltimore, Md.) 148 11463850
2021 Chimeric STAR receptors using TCR machinery mediate robust responses against solid tumors. Science translational medicine 135 33762437
1996 Expression of steroidogenic acute regulatory protein (StAR) in the human ovary. The Journal of clinical endocrinology and metabolism 129 8923870
2014 BTG2: a rising star of tumor suppressors (review). International journal of oncology 121 25405282
2023 Circular RNA: A promising new star of vaccine. Journal of translational internal medicine 113 38130633
2009 Hepatic stellate cell: a star cell in the liver. The international journal of biochemistry & cell biology 113 19433304
2019 Endoplasmic Reticulum Stress-Induced Upregulation of STARD1 Promotes Acetaminophen-Induced Acute Liver Failure. Gastroenterology 112 31029706
2020 Sensitization in transplantation: Assessment of risk (STAR) 2019 Working Group Meeting Report. American journal of transplantation : official journal of the American Society of Transplantation and the American Society of Transplant Surgeons 99 32342639
2012 STAR RNA-binding protein Quaking suppresses cancer via stabilization of specific miRNA. Genes & development 98 22751500
1999 The steroidogenic acute regulatory protein (StAR): a window into the complexities of intracellular cholesterol trafficking. Recent progress in hormone research 98 10548884
2018 A Rising Star in Pancreatic Diseases: Pancreatic Stellate Cells. Frontiers in physiology 97 29967585
1999 The STAR protein QKI-6 is a translational repressor. Proceedings of the National Academy of Sciences of the United States of America 97 10535969
2000 Conservation of steroidogenic acute regulatory (StAR) protein structure and expression in vertebrates. Molecular and cellular endocrinology 94 11064158
2020 HDAC11: a rising star in epigenetics. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 92 32841898
2015 Argonaute 2: A Novel Rising Star in Cancer Research. Journal of Cancer 91 26284139
1997 Regulation of expression of the steroidogenic acute regulatory protein (StAR) gene: a central role for steroidogenic factor 1. Steroids 90 9029708
2022 NLRP3 inflammasome: The rising star in cardiovascular diseases. Frontiers in cardiovascular medicine 83 36204568
2018 Circular RNA: new star, new hope in cancer. BMC cancer 82 30126353
2013 A star is born: new insights into the mechanism of astrogenesis. Cellular and molecular life sciences : CMLS 81 23907612
2019 Circular RNAs: a rising star in respiratory diseases. Respiratory research 76 30611252
2004 Neurosteroids: the StAR protein in the brain. Journal of neuroendocrinology 76 15344917
1999 Transcriptional regulation of the StAR gene. Molecular and cellular endocrinology 76 10411331
2021 STARD1 promotes NASH-driven HCC by sustaining the generation of bile acids through the alternative mitochondrial pathway. Journal of hepatology 74 33515644
2020 CircRNA Is a Rising Star in Researches of Ocular Diseases. Frontiers in cell and developmental biology 72 33015046
1997 A StAR search: implications in controlling steroidgenesis. Biology of reproduction 72 9116128
2001 Leydig cell aging: steroidogenic acute regulatory protein (StAR) and cholesterol side-chain cleavage enzyme. Journal of andrology 71 11191081
2008 Astrocyte, the star avatar: redefined. Journal of biosciences 69 19005240
2012 Star-PAP control of BIK expression and apoptosis is regulated by nuclear PIPKIα and PKCδ signaling. Molecular cell 68 22244330
2012 Gonadal transactivation of STARD1, CYP11A1 and HSD3B. Frontiers in bioscience (Landmark edition) 66 22201776
2021 Role of Constitutive STAR in Leydig Cells. International journal of molecular sciences 65 33670702
2021 NK Cell Therapy: A Rising Star in Cancer Treatment. Cancers 64 34439285
2016 Structural basis of RNA recognition and dimerization by the STAR proteins T-STAR and Sam68. Nature communications 60 26758068
2004 The role of potential splicing factors including RBMY, RBMX, hnRNPG-T and STAR proteins in spermatogenesis. International journal of andrology 60 15595951
2020 LncRNA SNHG5: A new budding star in human cancers. Gene 58 32360843
2014 The role of mitochondrial fusion and StAR phosphorylation in the regulation of StAR activity and steroidogenesis. Molecular and cellular endocrinology 58 25540920
2002 PBR, StAR, and PKA: partners in cholesterol transport in steroidogenic cells. Endocrine research 56 12530641
2016 Circular RNA: a new star in neurological diseases. The International journal of neuroscience 53 27619342
2019 LncRNA SNHG15: A new budding star in human cancers. Cell proliferation 51 31774607
2013 Expression and roles of steroidogenic acute regulatory (StAR) protein in 'non-classical', extra-adrenal and extra-gonadal cells and tissues. Molecular and cellular endocrinology 50 23415713
2017 The Steroidogenic Acute Regulatory Protein (StAR) Is Regulated by the H19/let-7 Axis. Endocrinology 49 27813675
2022 Extrachromosomal circular DNA (eccDNA): an emerging star in cancer. Biomarker research 48 35883211
2023 β-cell neogenesis: A rising star to rescue diabetes mellitus. Journal of advanced research 44 37839502
2008 Cholesterol interaction with the related steroidogenic acute regulatory lipid-transfer (START) domains of StAR (STARD1) and MLN64 (STARD3). The FEBS journal 43 18331352
2022 CircRNA: a rising star in plant biology. Journal of genetics and genomics = Yi chuan xue bao 42 35644325
2007 Mechanism of StAR's regulation of mitochondrial cholesterol import. Molecular and cellular endocrinology 42 17207924
2012 Hyperbranched polymeric "star vectors" for effective DNA or siRNA delivery. Accounts of chemical research 41 22353143
1999 Steroidogenic acute regulatory (StAR) protein: what's new? BioEssays : news and reviews in molecular, cellular and developmental biology 39 10462417
2022 CircRNA: An emerging star in the progression of glioma. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 36 35623170
2017 Exosomes: A Rising Star in Falling Hearts. Frontiers in physiology 36 28751864
2013 Intact cluster and chordate-like expression of ParaHox genes in a sea star. BMC biology 36 23803323
2024 Nanozyme as a rising star for metabolic disease management. Journal of nanobiotechnology 35 38711066
2020 Evaluation of STAR and Kallisto on Single Cell RNA-Seq Data Alignment. G3 (Bethesda, Md.) 35 32220951
2014 Perylene-cored star-shaped polycations for fluorescent gene vectors and bioimaging. ACS applied materials & interfaces 35 25159606
2024 Lactate: a rising star in tumors and inflammation. Frontiers in immunology 33 39660139
2020 Sestrin 2, a potential star of antioxidant stress in cardiovascular diseases. Free radical biology & medicine 33 33310138
2023 circRNA: a promising all-around star in the future. Epigenomics 32 37578015
2007 Dephosphorylation of TORC initiates expression of the StAR gene. Molecular and cellular endocrinology 32 17210223
2022 A novel adoptive synthetic TCR and antigen receptor (STAR) T-Cell therapy for B-Cell acute lymphoblastic leukemia. American journal of hematology 31 35491511
2022 Dental Pulp Fibroblast: A Star Cell. Journal of endodontics 31 35577145
2020 Reassessing Sarcopenia in Hypertension: STAR and ACE Inhibitors Excel. International journal of clinical practice 31 33108697
2018 Checkpoint-inhibition in ovarian cancer: rising star or just a dream? Journal of gynecologic oncology 31 30207101
2015 A-Raf: A new star of the family of raf kinases. Critical reviews in biochemistry and molecular biology 29 26508523
2007 Turnover of StAR protein: roles for the proteasome and mitochondrial proteases. Molecular and cellular endocrinology 29 17218054
2021 Acid ceramidase improves mitochondrial function and oxidative stress in Niemann-Pick type C disease by repressing STARD1 expression and mitochondrial cholesterol accumulation. Redox biology 28 34175669
2019 HIF 1 inhibits STAR transcription and testosterone synthesis in murine Leydig cells. Journal of molecular endocrinology 28 30400066
2010 The star family member QKI and cell signaling. Advances in experimental medicine and biology 28 21189683
2023 Startle: A star homoplasy approach for CRISPR-Cas9 lineage tracing. Cell systems 27 38128483
2024 Extracellular Vesicles: A New Star for Gene Drug Delivery. International journal of nanomedicine 26 38465204
2009 Differential regulation of the STARD1 subfamily of START lipid trafficking proteins in human macrophages. FEBS letters 26 19272380
2021 Exosomal circRNAs: A new star in cancer. Life sciences 25 33454367
2018 Cholesterol signaling in single cells: lessons from STAR and sm-FISH. Journal of molecular endocrinology 25 29691317
2008 Toward the NMR structure of StAR. Molecular and cellular endocrinology 25 19138724
2018 STAR Chimeric Post for rapid detection of circular RNA and fusion transcripts. Bioinformatics (Oxford, England) 24 29474638
2018 Triptolide: A new star for treating human malignancies. Journal of cancer research and therapeutics 23 29970675
2016 ACTH Action on StAR Biology. Frontiers in neuroscience 23 27999527
2009 Sam68: a new STAR in the male fertility firmament. Journal of andrology 23 19875495
2020 STARD1 and NPC1 expression as pathological markers associated with astrogliosis in post-mortem brains from patients with Alzheimer's disease and Down syndrome. Aging 22 31902793
2018 StAR protein and steroidogenic enzyme expressions in the rat Harderian gland. Comptes rendus biologies 22 29534958
2017 Steroidogenic acute regulatory protein (StAR) overexpression attenuates HFD-induced hepatic steatosis and insulin resistance. Biochimica et biophysica acta. Molecular basis of disease 22 28153708
2010 STAR trek: An introduction to STAR family proteins and review of quaking (QKI). Advances in experimental medicine and biology 22 21189682

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