Affinage

HAS2

Hyaluronan synthase 2 · UniProt Q92819

Length
552 aa
Mass
63.6 kDa
Annotated
2026-06-10
100 papers in source corpus 32 papers cited in narrative 32 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

HAS2 is a plasma membrane hyaluronan synthase whose synthesis of high-molecular-mass hyaluronan (HA) drives cell migration, epithelial-mesenchymal transition, matrix homeostasis, and developmental morphogenesis (PMID:14729574, PMID:23108409, PMID:27094859). It assembles into functional homomeric complexes and heteromers with HAS1 and HAS3 through an N-terminal domain, with HAS1 co-expression dampening HAS2/HAS3-driven HA output (PMID:25795779), and exhibits a lower UDP-GlcNAc requirement than HAS1 such that HAS2 activity scales with UDP-sugar availability (PMID:23303191). Enzyme activity and trafficking are tightly controlled by post-translational modification: AMPK phosphorylation of Thr-110 is inhibitory and retains HAS2 in the ER, ubiquitination at K190 is required for HA synthesis and plasma membrane residence, and O-GlcNAcylation at Ser-221 is stimulatory, with these marks collectively governing ER-to-plasma-membrane transit (PMID:21228273, PMID:30394292). HAS2 protein is additionally turned over by autophagy through a dynamic interaction with ATG9A (PMID:32084457). The HAS2 gene is induced by TGFβ via Smad and p38 MAPK signaling (PMID:23108409), by NF-κB downstream of pro-inflammatory cytokines (PMID:20522558), and by transcription factors STAT3, SMAD4, ZEB1, FOXH1, and TCF7L2 (PMID:24847057, PMID:28086235, PMID:30282636, PMID:31489963, PMID:36358989), frequently acting in concert with its natural antisense transcript HAS2-AS1, which remodels chromatin at the HAS2 promoter (PMID:25183006); it is repressed by AMPK and SIRT1 (PMID:21228273, PMID:31932306) and by microRNAs miR-23 and miR-26b that target the HAS2 transcript directly (PMID:21778427, PMID:26887530). Functionally, HAS2-derived HA acts upstream of Rac1 to support lamellipodia and directed migration (PMID:14729574), promotes tumor invasion by suppressing TIMP-1 and sustaining FAK/PI3K/Akt signaling (PMID:22016393), drives EMT through a ZEB1-CD44 autocrine loop (PMID:28086235), and is essential in vivo for skeletal growth, chondrocyte maturation, synovial joint cavitation, aggrecan retention in cartilage, and cardiac endocardial cushion formation (PMID:19633173, PMID:27094859, PMID:21778427). Dysregulated HAS2 expression—via upstream duplication, 3'UTR shortening, or m6A-dependent mRNA stabilization—causes pathological HA accumulation linked to canine periodic fever, pulmonary hypertension, and breast cancer progression (PMID:21437276, PMID:35671866, PMID:37705505), while sustained high-molecular-mass HA production confers anti-cancer and anti-inflammatory benefits (PMID:37612507).

Mechanistic history

Synthesis pass · year-by-year structured walk · 20 steps
  1. 1999 Medium

    Established that HA production by HAS2 is not merely a structural byproduct but is itself sufficient to drive tumor cell proliferation and tumorigenicity.

    Evidence Stable HAS2 overexpression in HT1080 cells with colony formation and nude mouse xenograft assays

    PMID:10070975

    Open questions at the time
    • Does not resolve whether the effect requires extracellular HA or signaling through a specific HA receptor
    • Single cell line and single lab
  2. 2002 Medium

    Showed that the active process of HA synthesis by HAS2, rather than steady-state pericellular HA abundance, controls keratinocyte migration and lamellipodia formation.

    Evidence Bidirectional sense/antisense HAS2 transfection in keratinocytes with wounding assays and exogenous HA rescue

    PMID:12186949

    Open questions at the time
    • Molecular link between synthesis and cytoskeletal output not defined
    • Single lab
  3. 2004 High

    Placed Has2 genetically upstream of Rac1 in driving lamellipodia formation and directed cell migration in vivo, defining a signaling role for HA synthesis in morphogenesis.

    Evidence Morpholino knockdown and epistasis with CA/DN Rac1 constructs in zebrafish gastrulation

    PMID:14729574

    Open questions at the time
    • Direct biochemical mechanism linking HA to Rac1 activation unknown
    • Receptor mediating the signal not identified here
  4. 2009 High

    Defined the essential in vivo developmental requirement for HAS2-derived HA in skeletal growth, chondrocyte maturation, aggrecan deposition, and synovial joint formation.

    Evidence Prx1-Cre conditional Has2 knockout in limb bud mesoderm with histology and marker immunostaining

    PMID:19633173

    Open questions at the time
    • Does not separate intracellular from extracellular HA contributions
    • Cellular signaling downstream of HA loss not resolved
  5. 2011 High

    Identified AMPK phosphorylation of Thr-110 as a direct, isoform-specific inhibitory switch on HAS2 enzymatic activity.

    Evidence AMPK activators/inhibitors, KO cells, T110 mutagenesis, and in vitro kinase assay

    PMID:21228273

    Open questions at the time
    • Did not yet link Thr-110 phosphorylation to trafficking (resolved later)
    • Structural basis of inhibition unknown
  6. 2011 High

    Defined a HAS2→TIMP-1→FAK/PI3K/Akt pathway controlling tumor cell invasion, connecting HA synthesis to matrix proteolysis and adhesion signaling.

    Evidence Reciprocal HAS2 KD/OE with TIMP-1 siRNA and antibody rescue in MDA-MB-231 cells and invasion assays

    PMID:22016393

    Open questions at the time
    • Mechanism by which HAS2 suppresses TIMP-1 not defined
    • Receptor coupling HA to FAK signaling not identified
  7. 2010 Medium

    Established HAS2 as the specific isoform mediating cytokine-induced, NF-κB-dependent HA synthesis and monocyte adhesion in inflamed endothelium.

    Evidence HAS2 siRNA, NF-κB inhibition, and U937 monocyte adhesion assays

    PMID:20522558

    Open questions at the time
    • Direct NF-κB occupancy at the HAS2 locus not shown here
    • Single lab
  8. 2012 High

    Demonstrated that TGFβ drives EMT through HAS2 induction via Smad and p38 MAPK, with the EMT-promoting effect independent of extracellular HA or CD44.

    Evidence HAS2 siRNA, pathway inhibitors, hyaluronidase, and CD44 blockade in NMuMG cells with EMT marker and migration readouts

    PMID:23108409

    Open questions at the time
    • Intracellular mechanism independent of secreted HA not mechanistically explained
    • Single epithelial model
  9. 2013 Medium

    Characterized the distinct enzymatic kinetics of HAS2, showing it requires lower UDP-GlcNAc than HAS1 and scales activity with UDP-sugar availability.

    Evidence HAS1-3 transfection in COS-1 cells with UDP-sugar HPLC and HA ELISA under substrate manipulation

    PMID:23303191

    Open questions at the time
    • No structural basis for substrate affinity differences
    • In vitro overexpression context
  10. 2014 High

    Identified multiple direct transcriptional inputs to HAS2 (STAT3 and an O-GlcNAcylation/NF-κB/HAS2-AS1 chromatin axis), revealing layered metabolic and signaling control of HAS2 expression.

    Evidence ChIP for STAT3 and NF-κB at HAS2/HAS2-AS1 promoters, OGT inhibition, and HAS2-AS1 siRNA in keratinocytes and aortic smooth muscle cells

    PMID:24847057 PMID:25183006

    Open questions at the time
    • Interplay between the various transcription factors not integrated
    • HAS2-AS1 chromatin mechanism partially defined
  11. 2015 High

    Showed HAS2 assembles into homo- and heteromeric complexes with HAS1 and HAS3 via its N-terminal domain, establishing isoform cross-regulation at the protein level.

    Evidence FRET, PLA, co-IP, and deletion mapping in live cells with co-transfection functional assays

    PMID:25795779

    Open questions at the time
    • Stoichiometry and structural architecture of complexes unknown
    • Physiological regulation of complex formation unclear
  12. 2016 High

    Demonstrated that HAS2-produced HA is strictly required to build the pericellular matrix and retain aggrecan in chondrocytes.

    Evidence CRISPR/Cas9 Has2 knockout with adenoviral rescue and pericellular matrix exclusion assays in chondrosarcoma cells

    PMID:27094859

    Open questions at the time
    • Does not address signaling consequences of matrix loss
    • Cell line model
  13. 2018 High

    Integrated three post-translational modifications (ubiquitination at K190, phosphorylation at T110, O-GlcNAcylation at S221) into a unified model coupling HAS2 enzymatic activity, stability, and ER-to-plasma-membrane trafficking.

    Evidence Site-directed mutagenesis, Dendra2/EGFP-HAS2 imaging, surface biotinylation, and Rab10 siRNA

    PMID:30394292

    Open questions at the time
    • Enzymes catalyzing K190 ubiquitination and S221 O-GlcNAcylation not all identified
    • Crosstalk dynamics between modifications not resolved
  14. 2018 Medium

    Revealed a ZEB1–HAS2–CD44 positive feedback loop driving EMT and metastasis, showing HA can reinforce its own production transcriptionally.

    Evidence ZEB1 ChIP at HAS2 promoter, siRNA, and HA-conditioned medium experiments in breast cancer cells

    PMID:28086235

    Open questions at the time
    • Feedback loop kinetics not quantified
    • Single lab
  15. 2018 Medium

    Extended HAS2 transcriptional control to developmental and antisense-dependent contexts, including FOXH1-mediated activation in mesendoderm differentiation and a TGFβ/Has2as requirement for EMT.

    Evidence siRNA, ChIP for FOXH1/H3K27me3, and differentiation/migration assays in human ESCs and mammary epithelial cells

    PMID:30194979 PMID:30282636

    Open questions at the time
    • Mechanism by which Has2as acts in trans versus cis not fully reconciled
    • Single lab per context
  16. 2019 Medium

    Demonstrated that HAS2 exerts anti-procatabolic effects in chondrocytes through an intracellular mechanism independent of secreted HA, and identified SMAD4 as a direct transcriptional activator coupled to a CD44-Caspase3 axis.

    Evidence Inducible adenoviral HAS2 overexpression with cell-autonomous readouts; SMAD4 ChIP and KD in granulosa cells

    PMID:31270213 PMID:31489963

    Open questions at the time
    • The intracellular HAS2 effector mechanism is not molecularly defined
    • Different cell systems not cross-validated
  17. 2020 Medium

    Identified two distinct turnover/repression routes for HAS2: SIRT1-mediated transcriptional repression via NF-κB/HAS2-AS1, and autophagic degradation of HAS2 protein through ATG9A.

    Evidence SIRT1 activators with NF-κB translocation analysis; live-cell/super-resolution imaging of HAS2-ATG9A with in vivo chloroquine treatment

    PMID:31932306 PMID:32084457

    Open questions at the time
    • Autophagy receptor selecting HAS2 not identified
    • Whether ATG9A interaction is direct unresolved
  18. 2022 High

    Showed that post-transcriptional HAS2 dysregulation through 3'UTR shortening causes HA hyper-synthesis, bioenergetic dysfunction, and pulmonary hypertension in vivo.

    Evidence NUDT21 knockdown, 3'UTR reporters, transgenic HAS2 overexpression and conditional knockout, Seahorse flux, and hemodynamics

    PMID:35671866

    Open questions at the time
    • Mechanism linking HA synthesis to mitochondrial dysfunction not fully defined
    • Relevance to human PH genetics not established
  19. 2023 High

    Established m6A-dependent stabilization of HAS2 mRNA (via VIRMA/IGF2BP3) as a driver of breast cancer progression, and showed sustained high-molecular-mass HA confers cancer resistance and lifespan extension.

    Evidence VIRMA shRNA, VIRMA-IGF2BP3 co-IP, mRNA stability assays; transgenic nmrHas2 mice with tumor/lifespan/inflammation readouts and exogenous HMM-HA rescue

    PMID:37612507 PMID:37705505

    Open questions at the time
    • Context-dependence of HA being pro- versus anti-tumorigenic not reconciled
    • Receptors mediating protective HMM-HA effects not defined here
  20. 2025 Medium

    Revealed a protective, sex-specific role for fibroblast-derived Has2 in limiting acute heart failure after myocardial infarction, independent of fibrosis.

    Evidence Fibroblast-specific Has2 conditional knockout with echocardiography and histopathology in a myocardial infarction model

    PMID:41250926

    Open questions at the time
    • Mechanism of cardioprotection and basis of sex specificity unknown
    • Single lab

Open questions

Synthesis pass · forward-looking unresolved questions
  • How HAS2-derived HA produces signaling outcomes intracellularly versus extracellularly, and the structural basis for HAS2 catalysis, complex assembly, and PTM-controlled trafficking, remain unresolved.
  • No high-resolution structure of HAS2 in the captured literature
  • Molecular identity of the intracellular HA-independent effector mechanism unknown
  • Direct receptor coupling HA to Rac1, FAK, and ZEB1 signaling not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016740 transferase activity 3
Localization
GO:0005794 Golgi apparatus 2 GO:0005886 plasma membrane 2 GO:0005783 endoplasmic reticulum 1
Pathway
R-HSA-1266738 Developmental Biology 3 R-HSA-1643685 Disease 3 R-HSA-1474244 Extracellular matrix organization 2
Partners
ATG9AHAS1HAS3
Complex memberships
HAS2 homomerHAS2-HAS1 heteromerHAS2-HAS3 heteromer

Evidence

Reading pass · 32 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 HAS2 overexpression in human HT1080 cells directly increases hyaluronan production and promotes anchorage-independent growth and tumorigenicity in nude mice, demonstrating that HA production by tumor cells per se drives cell proliferation in tissues. Stable transfection of HAS2, colony formation in semisolid medium, xenograft tumor growth assay Cancer Research Medium 10070975
2002 HAS2-driven hyaluronan synthesis (rather than abundance of pericellular hyaluronan per se) controls keratinocyte migration and lamellipodia formation; HAS2-antisense cells show delayed S-phase entry, smaller lamellipodia, and increased vinculin-containing adhesion plaques. Stable sense/antisense HAS2 transfection in keratinocytes, in vitro wounding assay, exogenous HA rescue experiments, Streptomyces hyaluronidase treatment Journal of Cell Science Medium 12186949
2004 Zebrafish Has2 is required upstream of Rac1 for lamellipodia formation and dorsal migration of lateral cells during gastrulation; epistasis analyses with constitutively active and dominant-negative Rac1 place Has2 upstream of Rac1 activation, and the effect is cell-autonomous. Antisense morpholino knockdown, epistasis with CA/DN Rac1 constructs, ectopic has2 expression, cell migration analysis in zebrafish embryos Development High 14729574
2004 Vasodilatory prostaglandins (prostacyclin analogue iloprost, PGE2) upregulate HAS2 mRNA and HA synthesis in human arterial smooth muscle cells via EP2 and IP receptors and cAMP signaling; COX-2 activity is required for basal HAS2 expression; HAS2-specific siRNA knockdown abolishes iloprost-stimulated HA secretion and promotes cell spreading. RT-PCR for HAS2, RNA interference (RNAi) targeting HAS2, pharmacological receptor agonists/antagonists, cAMP analogues and forskolin treatment Circulation Research Medium 14752026
2009 Conditional inactivation of Has2 in limb bud mesoderm severely shortens skeletal elements, disrupts growth plate organization, reduces aggrecan deposition, decreases hypertrophic chondrocyte number, prevents secondary ossification center formation, and abolishes synovial joint cavity formation, demonstrating essential roles for HA in skeletal growth, patterning, chondrocyte maturation, and joint formation. Conditional knockout using floxed Has2 allele and Prx1-Cre transgene; histology, immunostaining for aggrecan and hypertrophic markers Development High 19633173
2010 Proinflammatory cytokines (IL-1β, TNF-α, TNF-β) induce HA synthesis and monocyte adhesion in human endothelial cells specifically through HAS2 via the NF-κB signaling pathway; HAS2-specific siRNA knockdown abolishes cytokine-induced HA synthesis and monocyte adhesion. siRNA knockdown of HAS2, NF-κB pathway inhibition, U937 monocyte adhesion assay, RT-PCR for HAS2 Journal of Biological Chemistry Medium 20522558
2011 AMPK phosphorylates Thr-110 of human HAS2, directly inhibiting its enzymatic activity and reducing HA synthesis; the other two HAS isoenzymes (HAS1 and HAS3) are not modified by AMPK. This inhibition reduces AoSMC proliferation, migration, and immune cell recruitment. AMPK activators (AICAR, metformin), AMPK-specific inhibitor and knockout cell lines, site-specific mutagenesis of HAS2 Thr-110, in vitro kinase assay, migration and proliferation assays Journal of Biological Chemistry High 21228273
2011 HAS2 knockdown in a bone-metastatic MDA-MB-231 clone completely suppresses invasion through induction of TIMP-1 and dephosphorylation of focal adhesion kinase; HAS2 also supports EGF-mediated FAK/PI3K/Akt signaling. Rescue by HAS2 re-expression, TIMP-1 siRNA, or TIMP-1-blocking antibodies confirms the pathway. siRNA knockdown and overexpression of HAS2, basement membrane invasion assay, Western blot for FAK phosphorylation, PI3K/Akt pathway inhibitors, TIMP-1 siRNA rescue Journal of Biological Chemistry High 22016393
2011 miR-23 directly targets Has2 (and Icat, Tmem2) to restrict endocardial cushion formation in zebrafish; Has2 upregulation is responsible for excessive endocardial cushion cell differentiation in dicer mutants; miR-23 also inhibits TGF-β-induced endothelial-to-mesenchymal transition in mouse endothelial cells by suppressing Has2. Zebrafish dicer mutants, miR-23 gain/loss-of-function, in silico target prediction combined with in vivo testing, endocardial cushion phenotype analysis Circulation Research High 21778427
2011 A novel unstable duplication upstream of HAS2 in Shar-Pei dogs increases HAS2 expression, leading to HA accumulation in skin (thick folded skin phenotype) and a periodic fever syndrome; higher copy number of the 16.1 kb duplication is associated with both increased HAS2 expression and fever syndrome. Genome-wide SNP analysis, targeted resequencing, copy number analysis, expression quantification of HAS2 PLoS Genetics Medium 21437276
2012 TGFβ potently stimulates HA synthesis via upregulation of HAS2 in NMuMG mammary epithelial cells through kinase-active type I TGFβ receptor, Smad signaling, and p38 MAPK activation; HAS2 knockdown inhibits TGFβ-induced EMT (~50% reduction), suppresses EMT markers (fibronectin, Snail1, Zeb1), and completely abolishes TGFβ-induced cell migration; extracellular HA or CD44 blocking are not required. siRNA knockdown of HAS2, Smad pathway inhibition, p38 MAPK inhibition, immunostaining for EMT markers, real-time PCR, cell migration assay, Streptomyces hyaluronidase treatment, CD44-blocking antibodies Oncogene High 23108409
2013 HAS2 requires lower cellular UDP-GlcNAc concentration than HAS1 to synthesize hyaluronan; HAS2 activity increases with UDP-sugar availability; transfected HAS2 consumes enough UDP-sugars to reduce their cellular content. These differences define distinct kinetic properties among the three HAS isoenzymes. Transfection of HAS1-3 into COS-1 cells, glucosamine supplementation and glucose deprivation, HPLC measurement of UDP-sugars, ELISA for secreted HA Journal of Biological Chemistry Medium 23303191
2014 O-GlcNAcylation (induced by glucosamine or PUGNAC) specifically increases HAS2 mRNA among the three HAS isoenzymes; the natural antisense transcript HAS2-AS1 is absolutely required for this O-GlcNAcylation-induced HAS2 transcription; O-GlcNAcylation recruits NF-κB subunit p65 to the HAS2-AS1 promoter, and HAS2-AS1 then regulates HAS2 transcription in cis by altering chromatin structure (O-GlcNAcylation and acetylation) around the HAS2 proximal promoter. Glucosamine and PUGNAC treatment, OGT inhibition, siRNA knockdown of HAS2-AS1, NF-κB ChIP, chromatin acetylation analysis, RT-PCR for HAS2 and HAS2-AS1 in human aortic smooth muscle cells Journal of Biological Chemistry Medium 25183006
2014 STAT3 phosphorylated at Tyr705 (via JAK2 and ERK1/2 activation downstream of P2Y14 receptor stimulation by UDP-glucose) directly binds to the HAS2 promoter and induces HAS2 transcription in keratinocytes; chromatin immunoprecipitation confirmed increased Tyr705-STAT3 promoter binding at the time of HAS2 induction. Chromatin immunoprecipitation (ChIP) for STAT3 at HAS2 promoter, JAK2/STAT3/Gi inhibitors, UDP-glucose treatment, RT-PCR for HAS2, keratinocyte migration assay Journal of Biological Chemistry High 24847057
2015 HAS2 forms functionally relevant homomeric and heteromeric complexes with HAS1 and HAS3; complexes exist in both Golgi and plasma membrane; interaction is mediated mainly via the N-terminal 86-amino acid domain; HAS1 co-transfection reduces HAS2/HAS3-driven HA synthesis, indicating functional cooperation. HAS2 immunoprecipitates contain functional HAS2 homomers and heteromers with HAS3. FRET microscopy, flow cytometric FRET quantification, acceptor photobleaching, proximity ligation assay with endogenous HAS antibodies, C-terminal deletion constructs, co-transfection functional assays Journal of Biological Chemistry High 25795779
2016 CRISPR/Cas9 knockout of HAS2 in rat chondrosarcoma chondrocytes abolishes the pericellular HA matrix and completely prevents aggrecan retention; restoration of HAS2 by adenoviral transduction rescues pericellular matrix and aggrecan binding, demonstrating that HA produced by HAS2 is essential for aggrecan retention. CRISPR/Cas9 gene editing of Has2, adenoviral rescue with Has2, pericellular matrix exclusion assay, exogenous aggrecan addition, pellet culture neocartilage model Matrix Biology High 27094859
2018 Post-translational modifications control HAS2 trafficking and activity: ubiquitination (K190R mutation) blocks HA synthesis and reduces enzyme degradation while increasing plasma membrane residence; phosphorylation site (T110A) retains HAS2 in ER, blocks PM trafficking, and abolishes HA synthesis; O-GlcNAcylation (S221A) reduces HA synthesis; S221 phosphomimetics (S221D/E) block synthesis and accelerate decay, indicating alternative regulation by O-GlcNAc versus phosphorylation. Site-directed mutagenesis of K190, T110, S221; Dendra2-/EGFP-HAS2 fusions; confocal microscopy; TIRF microscopy; cell-surface biotinylation; photo-conversion of Dendra2; Rab10 siRNA; glucosamine treatment Matrix Biology High 30394292
2018 ZEB1 directly activates HAS2 expression, and HAS2-derived HA in turn elevates ZEB1 via CD44s, forming a positive autocrine feedback loop that promotes EMT and breast cancer metastasis. siRNA knockdown, ChIP for ZEB1 at HAS2 promoter, HA-conditioned medium experiments, correlation analysis in cancer cell lines Oncotarget Medium 28086235
2018 TGFβ induces Has2, Has2as (antisense), and Hmga2 expression via Smad and non-Smad pathways in mouse mammary epithelial cells; Has2as abrogation suppresses TGFβ-induced EMT markers (Snai1, Hmga2, Fn1) and mesenchymal phenotype; CD44, but not Hmmr, is required for TGFβ-mediated EMT phenotype; Akt and Erk1/2 activation is required for Has2as/Has2 induction and cell motility. siRNA knockdown of Has2as, CD44, Hmmr; Smad pathway inhibition; Akt/ERK inhibitors; EMT marker RT-PCR; migration assay Matrix Biology Medium 30194979
2018 Activin/Smad2 and Wnt/β-catenin signals cooperate via FOXH1 on open chromatin (following EZH2-PRC2 eviction) to activate HAS2 expression during mesendoderm differentiation of human ESCs; HAS2 knockdown greatly attenuates mesendoderm differentiation. siRNA knockdown of HAS2, ChIP for H3K27me3 and FOXH1, promoter occupancy analysis, Activin/Wnt treatment, differentiation assays Journal of Biological Chemistry Medium 30282636
2019 HAS2 overexpression in chondrocytes inhibits MMP3, MMP13, TSG6, and other procatabolic markers and enhances aggrecan retention; however, this inhibitory effect occurs only in HAS2-transduced cells (not in adjacent non-transduced cells), indicating an intracellular mechanism independent of extracellular HA. Inducible adenoviral HAS2 overexpression, ELISA for HA, RT-PCR and Western blot for MMPs and procatabolic markers, IL-1β/LPS/TNFα/HA oligosaccharide stimulation, metabolic flux analysis Journal of Biological Chemistry Medium 31270213
2019 SMAD4 directly binds to the HAS2 promoter to induce HAS2 expression and HA secretion in porcine granulosa cells; the downstream CD44-Caspase3 axis is activated by SMAD4 through this HAS2-HA system; miR-26b attenuates HAS2 expression via SMAD4-dependent and -independent mechanisms. ChIP for SMAD4 at HAS2 promoter, siRNA knockdown of SMAD4, HAS2 overexpression/knockdown, luciferase reporter assay, apoptosis/proliferation assays Journal of Cellular Physiology Medium 31489963
2020 SIRT1 activation reduces HAS2 expression and HA accumulation in aortic smooth muscle cells by preventing nuclear translocation of NF-κB (p65), which reduces HAS2-AS1 levels, and HAS2-AS1 epigenetically controls HAS2 mRNA expression; SIRT1 also reduces RHAMM and TSG6 expression to inhibit HA-mediated monocyte adhesion and cell migration. SIRT1 activators (SRT1720, resveratrol), NF-κB nuclear translocation analysis, HAS2-AS1 quantification, monocyte adhesion assay, cell migration assay Journal of Biological Chemistry Medium 31932306
2020 HAS2 is degraded in vascular endothelial cells via autophagy (evoked by nutrient deprivation, mTOR inhibition, or proteoglycan fragments endorepellin/endostatin); live-cell and super-resolution microscopy reveal dynamic interaction between HAS2 and ATG9A during autophagic degradation; inhibiting autophagic flux with chloroquine increases HAS2 levels in heart and aorta in vivo; autophagic induction suppresses HA production and inhibits angiogenic sprouting. Live-cell confocal and super-resolution microscopy, nutrient deprivation, mTOR inhibitors, chloroquine treatment in vivo, endorepellin treatment, angiogenic sprouting assay Matrix Biology Medium 32084457
2022 3'UTR shortening of HAS2 (caused by depletion of NUDT21, a master regulator of alternative polyadenylation) leads to HAS2 hyper-expression in pulmonary artery smooth muscle cells, driving HA hyper-synthesis, bioenergetic dysfunction (impaired mitochondrial oxidative capacity and glycolytic shift), and pro-remodeling phenotypes; transgenic mice mimicking HAS2 hyper-synthesis in smooth muscle cells develop spontaneous pulmonary hypertension; targeted HAS2 deletion prevents experimental PH. NUDT21 knockdown, 3'UTR cloning and reporter assay, transgenic HAS2 overexpression in smooth muscle cells, conditional HAS2 knockout, Seahorse metabolic flux analysis, pulmonary hemodynamic measurements Matrix Biology High 35671866
2023 KIAA1429/VIRMA (a component of the m6A methyltransferase complex) binds to m6A-reader IGF2BP3, leading to stabilization of m6A-modified HAS2 mRNA in the cytosol of breast cancer cells; VIRMA knockdown inhibits breast cancer cell proliferation, migration, and invasion. shRNA knockdown of KIAA1429/VIRMA, co-immunoprecipitation of VIRMA with IGF2BP3, m6A RNA-binding analysis, mRNA stability assays, proliferation/migration/invasion assays EMBO Reports Medium 37705505
2023 Transgenic mice overexpressing naked mole-rat Has2 (nmrHas2) show increased high-molecular-mass hyaluronan in several tissues, reduced spontaneous and induced cancer incidence, extended lifespan, and attenuated multi-tissue inflammation; these beneficial effects are conferred by HMM-HA and are not specific to the nmrHas2 gene sequence, as they can be recapitulated by exogenous HMM-HA. Transgenic mouse overexpressing nmrHas2, tumor incidence studies, lifespan analysis, transcriptome profiling, immune cell functional assays, gut barrier function assays, exogenous HMM-HA administration Nature High 37612507
2011 Nephronectin acts as an upstream regulator of Bmp4-Has2 signaling in zebrafish AV canal differentiation; inhibition of has2 in npnt morphants rescues the endocardial (but not myocardial) expansion, placing Has2 downstream of Bmp4 and upstream of endocardial AV cell fate specification. Morpholino knockdown of npnt and has2 in zebrafish, epistasis analysis, in situ hybridization for has2 and AV markers, BMP inhibition Development Medium 21937601
2016 miR-26b directly binds the 3'UTR of HAS2 mRNA (confirmed by luciferase reporter assay) and negatively regulates HAS2 expression and HA synthesis in porcine granulosa cells; reduced HAS2 via miR-26b promotes granulosa cell apoptosis through the HAS2-HA-CD44-Caspase3 pathway. Luciferase reporter assay for miR-26b binding to HAS2 3'UTR, miR-26b mimic/inhibitor transfection, HAS2 KD/OE, HA ELISA, CD44/Caspase3 expression, apoptosis assays Scientific Reports Medium 26887530
2018 Extracellular ATP activates HAS2 expression in keratinocytes via P2Y2 receptor → Ca2+/calmodulin-dependent protein kinase II, PKC, MAPK, and CREB-dependent pathways; AMP and adenosine (ATP degradation products) inhibit HAS2 expression, providing a feedback mechanism to shut off HA production. Pharmacological inhibitors of PKC, CaMKII, MAPK, CREB, JAK2; Gi-coupled receptor inhibitor; purinergic receptor agonists; RT-PCR for HAS2; HA ELISA; cell migration assay Biochemical Journal Medium 29626161
2025 Fibroblast-derived Has2 (deleted by fibroblast-specific Cre) limits acute heart failure following myocardial infarction in male mice; Has2-deficient male mice show exacerbated heart failure (lower cardiac output, stroke volume) at 1 week post-MI without changes in fibrosis, cardiomyocyte size, or capillary density; sex-specific effect not evident in females. Fibroblast-specific Has2 conditional knockout, non-reperfused myocardial infarction model, echocardiography, histopathology, collagen and HA measurements Physiological Reports Medium 41250926
2022 FGF9 promotes HAS2 expression in palatal mesenchyme via the Wnt/β-catenin/TCF7L2 pathway; TCF7L2 binds the HAS2 promoter and activates its transcription; Fgf9 knockout reduces TCF7L2 and HAS2, and TCF-dependent agonist-induced HA expression is blocked in Fgf9-null palate due to TCF7L2 loss. Fgf9 conditional knockout (Ddx4-Cre), immunohistochemistry, Western blot, ELISA for HA, JASPAR binding site prediction, rescue with TCF-dependent agonist BML-284 Biomolecules Medium 36358989

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1999 Overproduction of hyaluronan by expression of the hyaluronan synthase Has2 enhances anchorage-independent growth and tumorigenicity. Cancer research 244 10070975
2005 The over-expression of HAS2, Hyal-2 and CD44 is implicated in the invasiveness of breast cancer. Experimental cell research 186 16125700
2003 EGF upregulates, whereas TGF-beta downregulates, the hyaluronan synthases Has2 and Has3 in organotypic keratinocyte cultures: correlations with epidermal proliferation and differentiation. The Journal of investigative dermatology 139 12787132
2012 Efficient TGFβ-induced epithelial-mesenchymal transition depends on hyaluronan synthase HAS2. Oncogene 127 23108409
2004 Has2 is required upstream of Rac1 to govern dorsal migration of lateral cells during zebrafish gastrulation. Development (Cambridge, England) 120 14729574
2009 Conditional inactivation of Has2 reveals a crucial role for hyaluronan in skeletal growth, patterning, chondrocyte maturation and joint formation in the developing limb. Development (Cambridge, England) 119 19633173
2014 Natural antisense transcript for hyaluronan synthase 2 (HAS2-AS1) induces transcription of HAS2 via protein O-GlcNAcylation. The Journal of biological chemistry 109 25183006
2010 Proinflammatory cytokines induce hyaluronan synthesis and monocyte adhesion in human endothelial cells through hyaluronan synthase 2 (HAS2) and the nuclear factor-kappaB (NF-kappaB) pathway. The Journal of biological chemistry 108 20522558
2011 A novel unstable duplication upstream of HAS2 predisposes to a breed-defining skin phenotype and a periodic fever syndrome in Chinese Shar-Pei dogs. PLoS genetics 107 21437276
2011 MicroRNA-23 restricts cardiac valve formation by inhibiting Has2 and extracellular hyaluronic acid production. Circulation research 103 21778427
2011 Hyaluronan synthesis is inhibited by adenosine monophosphate-activated protein kinase through the regulation of HAS2 activity in human aortic smooth muscle cells. The Journal of biological chemistry 102 21228273
2011 Hyaluronan synthase 2 (HAS2) promotes breast cancer cell invasion by suppression of tissue metalloproteinase inhibitor 1 (TIMP-1). The Journal of biological chemistry 98 22016393
2013 Hyaluronan synthase 1 (HAS1) requires higher cellular UDP-GlcNAc concentration than HAS2 and HAS3. The Journal of biological chemistry 97 23303191
2023 Increased hyaluronan by naked mole-rat Has2 improves healthspan in mice. Nature 93 37612507
2011 The human hyaluronan synthase 2 (HAS2) gene and its natural antisense RNA exhibit coordinated expression in the renal proximal tubular epithelial cell. The Journal of biological chemistry 82 21357421
2017 Long noncoding RNA HAS2-AS1 mediates hypoxia-induced invasiveness of oral squamous cell carcinoma. Molecular carcinogenesis 77 28485478
2018 Tumor-suppressive functions of 4-MU on breast cancer cells of different ER status: Regulation of hyaluronan/HAS2/CD44 and specific matrix effectors. Matrix biology : journal of the International Society for Matrix Biology 70 29673760
2004 Induction of hyaluronic acid synthase 2 (HAS2) in human vascular smooth muscle cells by vasodilatory prostaglandins. Circulation research 68 14752026
2018 Interconnected feedback loops among ESRP1, HAS2, and CD44 regulate epithelial-mesenchymal plasticity in cancer. APL bioengineering 65 31069317
2012 Hyaluronic acid, HAS1, and HAS2 are significantly upregulated during muscle hypertrophy. American journal of physiology. Cell physiology 63 22785117
2017 A novel ZEB1/HAS2 positive feedback loop promotes EMT in breast cancer. Oncotarget 62 28086235
2020 Hsa_circRNA_102002 facilitates metastasis of papillary thyroid cancer through regulating miR-488-3p/HAS2 axis. Cancer gene therapy 60 32862195
2002 Changed lamellipodial extension, adhesion plaques and migration in epidermal keratinocytes containing constitutively expressed sense and antisense hyaluronan synthase 2 (Has2) genes. Journal of cell science 58 12186949
2014 HAS2 and CD44 in breast tumorigenesis. Advances in cancer research 57 25081531
2009 Targeted suppression of Has2 mRNA in mouse cumulus cell-oocyte complexes by adenovirus-mediated short-hairpin RNA expression. Molecular reproduction and development 55 18951380
2007 Putative role of hyaluronan and its related genes, HAS2 and RHAMM, in human early preimplantation embryogenesis and embryonic stem cell characterization. Stem cells (Dayton, Ohio) 55 17872502
2011 Nephronectin regulates atrioventricular canal differentiation via Bmp4-Has2 signaling in zebrafish. Development (Cambridge, England) 52 21937601
2016 Conserved miR-26b enhances ovarian granulosa cell apoptosis through HAS2-HA-CD44-Caspase-3 pathway by targeting HAS2. Scientific reports 51 26887530
2014 Extensive CD44-dependent hyaluronan coats on human bone marrow-derived mesenchymal stem cells produced by hyaluronan synthases HAS1, HAS2 and HAS3. The international journal of biochemistry & cell biology 50 24406795
2018 Has2 natural antisense RNA and Hmga2 promote Has2 expression during TGFβ-induced EMT in breast cancer. Matrix biology : journal of the International Society for Matrix Biology 45 30194979
2020 Sirtuin 1 reduces hyaluronan synthase 2 expression by inhibiting nuclear translocation of NF-κB and expression of the long-noncoding RNA HAS2-AS1. The Journal of biological chemistry 44 31932306
2014 Extracellular UDP-glucose activates P2Y14 Receptor and Induces Signal Transducer and Activator of Transcription 3 (STAT3) Tyr705 phosphorylation and binding to hyaluronan synthase 2 (HAS2) promoter, stimulating hyaluronan synthesis of keratinocytes. The Journal of biological chemistry 42 24847057
2019 HAS2-AS1 is a novel LH/hCG target gene regulating HAS2 expression and enhancing cumulus cells migration. Journal of ovarian research 40 30819231
2019 CREB1 induced lncRNA HAS2-AS1 promotes epithelial ovarian cancer proliferation and invasion via the miR-466/RUNX2 axis. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 40 31082772
2018 Effects of mutations in the post-translational modification sites on the trafficking of hyaluronan synthase 2 (HAS2). Matrix biology : journal of the International Society for Matrix Biology 37 30394292
2015 Fluorescence resonance energy transfer (FRET) and proximity ligation assays reveal functionally relevant homo- and heteromeric complexes among hyaluronan synthases HAS1, HAS2, and HAS3. The Journal of biological chemistry 36 25795779
2014 The ABA-deficiency suppressor locus HAS2 encodes the PPR protein LOI1/MEF11 involved in mitochondrial RNA editing. Molecular plant 35 25708384
2018 Extracellular ATP activates hyaluronan synthase 2 (HAS2) in epidermal keratinocytes via P2Y2, Ca2+ signaling, and MAPK pathways. The Biochemical journal 32 29626161
2020 The transcription factor USF1 promotes glioma cell invasion and migration by activating lncRNA HAS2-AS1. Bioscience reports 31 32776110
2016 CRISPR/Cas9 knockout of HAS2 in rat chondrosarcoma chondrocytes demonstrates the requirement of hyaluronan for aggrecan retention. Matrix biology : journal of the International Society for Matrix Biology 31 27094859
2004 Light and metabolic regulation of HAS1, HAS1.1 and HAS2, three asparagine synthetase genes in Helianthus annuus. Plant physiology and biochemistry : PPB 30 15246064
2016 GREM1, EGFR, and HAS2; the oocyte competence markers for improved buffalo embryo production in vitro. Theriogenology 29 27448692
2020 Autophagic degradation of HAS2 in endothelial cells: A novel mechanism to regulate angiogenesis. Matrix biology : journal of the International Society for Matrix Biology 28 32084457
2010 Elevated glucose induces congenital heart defects by altering the expression of tbx5, tbx20, and has2 in developing zebrafish embryos. Birth defects research. Part A, Clinical and molecular teratology 28 20306498
2019 Long noncoding RNA HAS2-AS1 promotes tumor progression in glioblastoma via functioning as a competing endogenous RNA. Journal of cellular biochemistry 26 31385362
2023 KIAA1429/VIRMA promotes breast cancer progression by m6 A-dependent cytosolic HAS2 stabilization. EMBO reports 24 37705505
2007 LIF inhibits osteoblast differentiation at least in part by regulation of HAS2 and its product hyaluronan. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 24 17451373
2016 The mediator complex subunit Med10 regulates heart valve formation in zebrafish by controlling Tbx2b-mediated Has2 expression and cardiac jelly formation. Biochemical and biophysical research communications 22 27343557
2014 Genomewide association study identifies HAS2 as a novel susceptibility gene for adult asthma in a Japanese population. Clinical and experimental allergy : journal of the British Society for Allergy and Clinical Immunology 22 25251750
2012 Melanoma cell-derived factors stimulate hyaluronan synthesis in dermal fibroblasts by upregulating HAS2 through PDGFR-PI3K-AKT and p38 signaling. Histochemistry and cell biology 22 22825838
2023 Human TMEM2 is not a catalytic hyaluronidase, but a regulator of hyaluronan metabolism via HYBID (KIAA1199/CEMIP) and HAS2 expression. The Journal of biological chemistry 21 37196767
2015 Undifferentiated myxoid lipoblastoma with PLAG1-HAS2 fusion in an infant; morphologically mimicking primitive myxoid mesenchymal tumor of infancy (PMMTI)--diagnostic importance of cytogenetic and molecular testing and literature review. Cancer genetics 21 26701195
2006 Has2 expression in heart forming regions is independent of BMP signaling. Gene expression patterns : GEP 21 16458617
2019 Hyaluronan synthase 2 (HAS2) overexpression diminishes the procatabolic activity of chondrocytes by a mechanism independent of extracellular hyaluronan. The Journal of biological chemistry 20 31270213
2011 Increased HAS2-driven hyaluronic acid synthesis in shar-pei dogs with hereditary cutaneous hyaluronosis (mucinosis). Veterinary dermatology 20 21718367
2011 Fragmented hyaluronan is an autocrine chemokinetic motility factor supported by the HAS2-HYAL2/CD44 system on the plasma membrane. International journal of oncology 20 21743962
2019 TGF-β/SMAD4 signaling pathway activates the HAS2-HA system to regulate granulosa cell state. Journal of cellular physiology 18 31489963
2009 Hyaluronan oligosaccharide treatment of chondrocytes stimulates expression of both HAS-2 and MMP-3, but by different signaling pathways. Osteoarthritis and cartilage 18 19874928
2021 LncRNA HAS2-AS1 Promotes Glioblastoma Proliferation by Sponging miR-137. Frontiers in oncology 16 34094916
2019 Silencing of HAS2-AS1 mediates PI3K/AKT signaling pathway to inhibit cell proliferation, migration, and invasion in glioma. Journal of cellular biochemistry 16 30790335
2025 Metastatic tumor growth in steatotic liver is promoted by HAS2-mediated fibrotic tumor microenvironment. The Journal of clinical investigation 14 39946200
2022 The natural antisense transcript HAS2-AS1 regulates breast cancer cells aggressiveness independently from hyaluronan metabolism. Matrix biology : journal of the International Society for Matrix Biology 14 35395387
2021 The Attenuated Secretion of Hyaluronan by UVA-Exposed Human Fibroblasts Is Associated with Up- and Downregulation of HYBID and HAS2 Expression via Activated and Inactivated Signaling of the p38/ATF2 and JAK2/STAT3 Cascades. International journal of molecular sciences 14 33669634
2021 The Transcription Factor C/EBPβ Promotes HFL-1 Cell Migration, Proliferation, and Inflammation by Activating lncRNA HAS2-AS1 in Hypoxia. Frontiers in cell and developmental biology 14 33777961
2020 Vitamin D regulation of HAS2, hyaluronan synthesis and metabolism in triple negative breast cancer cells. The Journal of steroid biochemistry and molecular biology 14 32360595
2020 Long noncoding RNA HAS2-AS1 accelerates non-small cell lung cancer chemotherapy resistance by targeting LSD1/EphB3 pathway. American journal of translational research 13 32269726
2024 Enhancing Endogenous Hyaluronic Acid in Osteoarthritic Joints with an Anti-Inflammatory Supramolecular Nanofiber Hydrogel Delivering HAS2 Lentivirus. Small (Weinheim an der Bergstrasse, Germany) 12 38593309
2021 GNA14 stimulation of KLF7 promotes malignant growth of endometrial cancer through upregulation of HAS2. BMC cancer 12 33892667
2019 Atherosclerosis-associated endothelial cell apoptosis by miRNA let7-b-mediated downregulation of HAS-2. Journal of cellular biochemistry 12 31736114
2018 Activin/Smad2 and Wnt/β-catenin up-regulate HAS2 and ALDH3A2 to facilitate mesendoderm differentiation of human embryonic stem cells. The Journal of biological chemistry 12 30282636
2017 Evaluating The Effect of Melatonin on HAS2, and PGR expression, as well as Cumulus Expansion, and Fertility Potential in Mice. Cell journal 12 29308626
2023 Downregulation of HAS‑2 regulates the chondrocyte cytoskeleton and induces cartilage degeneration by activating the RhoA/ROCK signaling pathway. International journal of molecular medicine 11 37232339
2022 FGF9 Promotes Expression of HAS2 in Palatal Elevation via the Wnt/β-Catenin/TCF7L2 Pathway. Biomolecules 11 36358989
2021 WNT5A Enhances LH-Mediated Expression of HAS2 in Granulosa Cells. Reproductive sciences (Thousand Oaks, Calif.) 11 34542891
2020 Inhibition of HAS2 and hyaluronic acid production by 1,25-Dihydroxyvitamin D3 in breast cancer. Oncotarget 11 32774770
2013 Inhibition of HAS2 induction enhances the radiosensitivity of cancer cells via persistent DNA damage. Biochemical and biophysical research communications 11 24333416
2022 3'UTR shortening of HAS2 promotes hyaluronan hyper-synthesis and bioenergetic dysfunction in pulmonary hypertension. Matrix biology : journal of the International Society for Matrix Biology 9 35671866
2024 Calycosin inhibits the proliferation and metastasis of renal cell carcinoma through the MAZ/HAS2 signaling pathway. Phytotherapy research : PTR 8 39120474
2022 Cell-Type-Specific Expression of Hyaluronan Synthases HAS2 and HAS3 Promotes Goblet Cell Hyperplasia in Allergic Airway Inflammation. American journal of respiratory cell and molecular biology 8 35679095
2022 The Upregulation of HAS2-AS1 Relates to the Granulosa Cell Dysfunction by Repressing TGF-β Signaling and Upregulating HAS2. Molecular and cellular biology 8 35938797
2021 Hyaluronan synthase 2 (HAS2) regulates cell phenotype and invadopodia formation in luminal-like breast cancer cells. Molecular and cellular biochemistry 8 33954907
2016 In Vitro Effects of HAS-2 Gene Silencing on the Proliferation and Apoptosis of the MCF-7 Human Breast Cancer Cell Line. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 8 27915342
2014 Spatially restricted hyaluronan production by Has2 drives epithelial tubulogenesis in vitro. American journal of physiology. Cell physiology 8 25163516
2013 Triiodothyronine (T3) inhibits hyaluronate synthesis in a human dermal equivalent by downregulation of HAS2. In vitro cellular & developmental biology. Animal 8 23397370
2022 MicroRNA-101 regulates oocyte maturation in vitro via targeting HAS2 in porcine cumulus cells. Theriogenology 7 35567989
2017 Targeted HAS2 Expression Lessens Airway Responsiveness in Chronic Murine Allergic Airway Disease. American journal of respiratory cell and molecular biology 7 28787175
2024 Upregulation of HAS2 promotes glioma cell proliferation and chemoresistance via c-myc. Cellular signalling 6 38734194
2023 MicroRNA-376b is involved in the pathogenesis of thyroid-associated ophthalmopathy by regulating HAS2. Endocrine 6 37231239
2022 HAS2-Ezrin-ER axis plays a role in acquired antiestrogen resistance of ER-positive breast cancer. Frontiers in pharmacology 6 36386154
2020 HAS2-AS1 Acts as a Molecular Sponge for miR-137 and Promotes the Invasion and Migration of Glioma Cells by Targeting EZH2. Cell cycle (Georgetown, Tex.) 6 33064966
2016 Impact of Prunus Cerasus on PGR and HAS2 in Cumulus Cells and Fertility Outcome. Advanced pharmaceutical bulletin 6 27123419
2024 Piceatannol enhances hyaluronic acid synthesis through SIRT1-Mediated HAS2 upregulation in human dermal fibroblasts. Biochemistry and biophysics reports 5 38910870
2022 Has2 Regulates the Development of Ovalbumin-Induced Airway Remodeling and Steroid Insensitivity in Mice. Frontiers in immunology 5 35069543
2017 Effect of MT3 on Retinal and Choroidal TGF-β2 and HAS2 Expressions in Form Deprivation Myopia of Guinea Pig. Journal of ophthalmology 5 29163988
2010 Over-expression of has2 in synovium-derived mesenchymal stem cells may prevent adhesions following surgery of the digital flexor tendons. Medical hypotheses 5 21084166
2024 HAS2 facilitates glioma cell malignancy and suppresses ferroptosis in an FZD7-dependent manner. Cancer science 4 38816349
2012 Genetic association and gene-gene interaction of HAS2, HABP1 and HYAL3 implicate hyaluronan metabolic genes in glaucomatous neurodegeneration. Disease markers 4 22960332
2025 Fibroblast Has2 limits acute heart failure following myocardial infarction in male mice. Physiological reports 3 41250926
2024 Tanshinone IIA Protects Ischemia/Reperfusion-Induced Cardiomyocyte Injury by Inhibiting the HAS2/FGF9 Axis. Cardiology research and practice 3 39568660
2019 Expression of PTX3, HAS2 AND TNFAIP6 genes in relation to real-time proliferation of porcine endometrial luminal epithelial cells in primary cultivation model. Journal of biological regulators and homeostatic agents 2 31189490

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