Affinage

ME2

NAD-dependent malic enzyme, mitochondrial · UniProt P23368

Length
584 aa
Mass
65.4 kDa
Annotated
2026-06-10
50 papers in source corpus 16 papers cited in narrative 16 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

ME2 is a mitochondrial NAD(P)+-dependent malic enzyme that catalyzes the oxidative decarboxylation of malate to pyruvate, with distinct kinetics from its paralogs (high Km for malate, dual NAD/NADP cofactor use, allosteric activation by fumarate and inhibition by ATP), and functions across multiple tissues including pancreatic islets (PMID:19691144). By generating pyruvate and reducing equivalents, ME2 sustains mitochondrial respiration, ATP production via oxidative phosphorylation, and NADPH-dependent ROS clearance, such that its loss raises oxidative stress and triggers apoptosis (PMID:37079187). ME2 activity is tuned by a convergent set of activating post-translational modifications gated by nutrient state: SIRT5-mediated desuccinylation at K346 activates ME2 under glutamine deprivation to support proliferation (PMID:38007551), ACAT1-mediated acetylation at K156 activates it under low glucose to drive pyruvate/lactate output and chemoresistance (PMID:39951294), and PRMT1-mediated methylation activates it to promote hepatocellular carcinoma growth (PMID:39528487). Phosphorylation of the full-length cytoplasmic isoform (ME2fl) at S9 by AKT1 blocks its mitochondrial import and converts it into a scaffold that assembles glycolytic enzymes (PFKL, GAPDH, PKM2, LDHA), switching metabolism toward aerobic glycolysis (PMID:38263319). ME2-derived pyruvate also stabilizes β-catenin to promote migration and invasion (PMID:37110198). A homozygous frameshift variant producing truncated, unstable ME2 protein causes a human neurodevelopmental disorder, and the growth arrest of yeast lacking the ME2 ortholog is rescued by human ME2, establishing an essential mitochondrial role (PMID:39401966). The enzyme is allosterically druggable at a site distinct from the active site (embonic acid) and by active-site-directed inhibitors (NPD389) (PMID:26008970, PMID:24681895).

Mechanistic history

Synthesis pass · year-by-year structured walk · 8 steps
  1. 1985 Medium

    Before molecular characterization, the question was simply where ME2 resides in the genome; linkage mapping anchored the locus, providing the genetic foothold for later study.

    Evidence Family-based genetic linkage analysis placing ME2 on chromosome 6p near F13A

    PMID:2860058

    Open questions at the time
    • Chromosomal assignment only, no molecular or functional mechanism
    • No protein-level characterization
  2. 2009 Medium

    It was unclear how mitochondrial ME2 differs functionally from its paralogs; kinetic profiling established ME2 as a dual-cofactor, fumarate-activated, ATP-inhibited enzyme distinct from ME1 and ME3.

    Evidence Spectrophotometric enzyme activity and immunoblotting across human, rat, mouse islets and INS-1 cells

    PMID:19691144

    Open questions at the time
    • No structural basis for cofactor promiscuity or allosteric regulation
    • Physiological role in islets not resolved
  3. 2015 High

    Whether ME2 could be pharmacologically targeted and at what site was unknown; embonic acid was shown to allosterically inhibit ME2 at the fumarate site/dimer interface, distinct from the catalytic site, and induce p53-independent senescence.

    Evidence In vitro enzymatic inhibition, mutagenesis, thermal shift binding, shRNA knockdown and senescence assays in H1299 cells

    PMID:24681895 PMID:26008970

    Open questions at the time
    • Selectivity over ME1/ME3 not fully established
    • Off-target cellular effects of allosteric inhibition unclear
  4. 2023 High

    The first activating PTM regulation of ME2 was unknown; SIRT5 desuccinylation at K346 was identified as a nutrient-gated activation switch coupling glutamine availability to mitochondrial respiration.

    Evidence Reciprocal Co-IP, K346 mutagenesis, enzymatic and respiration assays, in vitro/in vivo cancer models

    PMID:38007551

    Open questions at the time
    • Succinyltransferase 'writer' not identified
    • Stoichiometry of K346 succinylation in vivo unknown
  5. 2023 Medium

    How ME2 enzymatic output feeds cellular signaling and survival was open; loss-of-function studies showed ME2 supplies pyruvate that stabilizes β-catenin to drive migration, and sustains OXPHOS/NADPH to suppress ROS and apoptosis.

    Evidence Knockdown/overexpression with pyruvate rescue, β-catenin assays; shRNA silencing and allosteric inhibitor with metabolite readouts and AML xenografts

    PMID:37079187 PMID:37110198

    Open questions at the time
    • Mechanism of pyruvate–β-catenin interaction not structurally defined
    • Tissue specificity of metabolic vulnerability unclear
  6. 2024 High

    Whether ME2 has a moonlighting non-catalytic function was unknown; AKT1 phosphorylation at S9 was shown to block mitochondrial import and repurpose cytoplasmic ME2fl as a glycolytic-enzyme scaffold, defining a metabolic switch toward aerobic glycolysis.

    Evidence Co-IP, S9 mutagenesis, subcellular fractionation, glycolytic flux assays, in vivo tumor models

    PMID:38263319

    Open questions at the time
    • Structural basis of the glycolytic scaffold assembly not resolved
    • Relative abundance of ME2fl scaffold vs mitochondrial pool not quantified
  7. 2024 Medium

    Whether arginine methylation regulates ME2 and disease relevance of ME2 loss were open; PRMT1 methylation was shown to activate ME2 in HCC, and a human frameshift variant plus yeast complementation established ME2 as essential for mitochondrial function and linked to neurodevelopmental disease.

    Evidence Co-IP and mutagenesis with respiration/proliferation assays (PRMT1); whole exome sequencing, truncated protein expression, yeast deletion/rescue

    PMID:39401966 PMID:39528487

    Open questions at the time
    • Exact methylated residue(s) not definitively mapped
    • Single patient; genotype-phenotype spectrum unknown
    • Mechanism connecting ME2 deficiency to neurodevelopment unclear
  8. 2025 High

    How ME2 activity is coupled to chemoresistance was unknown; ACAT1 acetylation at K156 was shown to activate ME2 under low glucose, driving lactate-dependent protein lactylation that supports homologous recombination repair.

    Evidence Co-IP, K156 mutagenesis, enzymatic/glucose-uptake assays, metabolomics, in vitro/in vivo chemoresistance models

    PMID:39951294

    Open questions at the time
    • Direct lactylation targets in HR repair not enumerated
    • Interplay between K156, K346 and other PTMs not resolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • An integrated model of how the multiple activating PTMs (succinylation, acetylation, methylation), AKT1 phosphorylation, and protein-stability control (CYP4F11) are coordinated under specific physiological conditions, and how ME2 dysfunction produces neurodevelopmental disease, remains unresolved.
  • No structural model of the regulated enzyme or its scaffold form
  • PTM hierarchy and crosstalk uncharacterized
  • Mechanistic link between ME2 loss and the neurodevelopmental phenotype not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016491 oxidoreductase activity 4 GO:0016829 lyase activity 1 GO:0060090 molecular adaptor activity 1
Localization
GO:0005739 mitochondrion 3 GO:0005829 cytosol 1
Pathway
R-HSA-1430728 Metabolism 3
Partners
ACAT1AKT1GAPDHLDHAPFKLPKM2PRMT1SIRT5

Evidence

Reading pass · 16 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2023 SIRT5 acts as a desuccinylase for ME2, removing succinylation at lysine 346. Glutamine deprivation enhances the SIRT5–ME2 interaction, promoting desuccinylation and activating ME2 enzymatic activity. Activated ME2 enhances mitochondrial respiration to support cell proliferation under glutamine deficiency. Co-immunoprecipitation, site-directed mutagenesis (K346), enzymatic activity assays, mitochondrial respiration measurement, in vitro and in vivo cancer models Cell death and differentiation High 38007551
2024 AKT1 phosphorylates the full-length cytoplasmic isoform of ME2 (ME2fl) at serine 9 in the mitochondrial localization signal peptide, preventing mitochondrial translocation. Cytoplasmic ME2fl functions as a scaffold assembling glycolytic enzymes PFKL, GAPDH, PKM2, and LDHA to promote glycolysis, thereby mediating a metabolic switch from mitochondrial TCA-cycle metabolism to aerobic glycolysis in tumor cells. Co-immunoprecipitation, site-directed mutagenesis (S9), subcellular fractionation, glycolytic flux assays, in vitro and in vivo tumor models, phosphorylation assays Nature communications High 38263319
2025 ACAT1 acetylates ME2 at lysine 156, potentiating ME2 enzymatic activity and facilitating pyruvate/lactate production from glutamine-derived malate. Decreased intracellular glucose (triggered by chemotherapy) drives this ACAT1-mediated ME2 acetylation, and ME2-derived lactate promotes protein lactylation involved in homologous recombination repair, contributing to chemoresistance. Co-immunoprecipitation, site-directed mutagenesis (K156), enzymatic activity assays, glucose uptake assays, in vitro and in vivo chemoresistance models, metabolomic analyses Advanced science High 39951294
2024 PRMT1 methylates ME2 (at arginine, with mutation at lysine 67 shown to mimic activated state), activating ME2 enzymatic activity and enhancing mitochondrial respiration to promote hepatocellular carcinoma cell growth and migration. ME2 was shown to directly interact with PRMT1 by co-immunoprecipitation. Co-immunoprecipitation, site-directed mutagenesis (R67K), enzymatic activity assays, mitochondrial respiration measurement, cell proliferation and migration assays Cell death & disease Medium 39528487
2015 The natural compound embonic acid (EA) allosterically inhibits mitochondrial NAD(P)+-dependent malic enzyme ME2 with an in vitro IC50 of ~1.4 µM. Mutagenesis and binding studies localized the EA binding site to the fumarate binding site or dimer interface, distinct from the catalytic active site. EA inhibition of ME2 in H1299 cancer cells induced cellular senescence via a p53-independent pathway. In vitro enzymatic inhibition assay, site-directed mutagenesis, thermal shift binding assay, shRNA knockdown, cellular senescence assay Oncotarget High 26008970
2014 NPD389 is an uncompetitive inhibitor of ME2 with respect to the substrate NAD+ and a mixed-type inhibitor with respect to L-malate, acting in a fast-binding mode. This was established by enzyme kinetics analysis of purified recombinant human ME2. High-throughput screening, enzyme kinetics analysis, thermal shift assay with purified recombinant human ME2 Acta pharmacologica Sinica Medium 24681895
2009 Mitochondrial ME2 has distinct kinetic properties from cytosolic ME1 and the other mitochondrial ME3: ME2 uses either NAD or NADP as cofactor, has a high Km for malate, and is allosterically activated by fumarate and inhibited by ATP. ME2 activity was directly detected and confirmed by immunoblotting in human, rat, and mouse pancreatic islets and INS-1 832/13 insulinoma cells. Spectrophotometric enzyme activity assay exploiting distinct kinetics, immunoblotting, tissue fractionation Archives of biochemistry and biophysics Medium 19691144
2023 ME2 promotes hepatocellular carcinoma cell migration and invasion by producing pyruvate, which directly binds to β-catenin and increases β-catenin protein levels. Pyruvate treatment rescued migration defects in ME2-depleted cells. ME2 knockdown and overexpression, cell migration/invasion assays, pyruvate rescue experiment, β-catenin binding and protein level measurements Metabolites Medium 37110198
2023 ME2 inhibition (by silencing or allosteric inhibitor disodium embonate) in AML cells decreases pyruvate and NADH, reducing ATP production via oxidative phosphorylation, and decreases NADPH leading to increased ROS and oxidative stress, ultimately inducing apoptosis. ME2 silencing also inhibited xenotransplanted human AML cell growth in vivo. shRNA silencing, allosteric inhibitor treatment, metabolite measurements (pyruvate, NADH, NADPH, ATP, ROS), xenograft mouse model Cellular oncology Medium 37079187
2021 ME2 inhibits mitochondrial ROS production and AMPK phosphorylation in glioblastoma cells, leading to SREBP-1 maturation and nuclear localization and enhancing the ACSS2 lipogenesis pathway. ME2 also promotes proneural-to-mesenchymal transition by upregulating mesenchymal markers (N-cadherin, vimentin, YKL40, MET) and downregulating OLIG2. ME2 overexpression and knockdown, ROS measurement, AMPK phosphorylation assay, SREBP-1 localization, gene expression analysis, cell migration/invasion assays Frontiers in oncology Low 34381734
2023 ME2 is a direct target of miR-214-3p, as established by dual luciferase reporter assay. Overexpression of ME2 ameliorated ferroptosis and cellular damage induced by miR-214-3p in cardiomyocytes, while ME2 depletion compromised the protective effects of miR-214-3p inhibitor against hypoxic injury. Dual luciferase reporter assay, ME2 overexpression and depletion in neonatal rat cardiomyocytes, hypoxia model, ferroptosis markers, mouse MI model Biochemical and biophysical research communications Medium 37087800
2024 A homozygous frameshift variant (c.1379_1380delTT, p.Phe460fs*22) in ME2 results in truncated, unstable ME2 protein and is associated with a neurodevelopmental disorder in a human patient. Deletion of the yeast ortholog of ME2 caused growth arrest rescued by overexpression of human ME2, supporting ME2's essential role in mitochondrial function. Whole exome sequencing, in vitro expression of truncated protein, yeast complementation (deletion + rescue by human ME2 overexpression) Clinical genetics Medium 39401966
1998 Murine E2-C (mE2-C), a cyclin-selective ubiquitin conjugating enzyme, is required for in vitro ubiquitination and degradation of cyclin B in human HeLa cell extracts. A dominant-negative form created by mutating the catalytic cysteine to serine blocked cyclin B ubiquitination and degradation. mE2-C expression varies with the cell cycle. Dominant-negative mutagenesis (catalytic cysteine to serine), in vitro ubiquitination assay in HeLa cell extracts, cell cycle expression analysis Oncogene Medium 9798675
2025 CYP4F11 prevents ubiquitin-proteasomal degradation of ME2, stabilizing ME2 protein levels. miR-195 suppresses CYP4F11, which leads to ME2 degradation and disrupts mitochondrial malate metabolism. Rescue experiments confirmed ME2 mediates CYP4F11's oncogenic effects in NSCLC. 3'-UTR luciferase reporter assay (miR-195/CYP4F11), CYP4F11 knockdown and overexpression, ME2 protein stability assay, rescue experiments, metabolomic analysis Frontiers of medicine Low 41359237
1985 Genetic linkage analysis placed the ME2 locus on the short arm of chromosome 6, closely linked to coagulation factor XIIIA (F13A) with a maximum lod score of 4.33 at recombination fraction 0.10. Map order established as GLO1–HLA–F13A–ME2. Family-based genetic linkage analysis (18 informative families, 54 children), lod score calculation Human genetics Medium 2860058
2024 In ME2-deficient pancreatic cancer cells (with co-deletion of SMAD4/ME2), ME3 is upregulated to compensate for ROS clearance. Targeted siRNA knockdown of ME3 in these ME2-deficient cells inhibited their capacity to clear ROS and, combined with doxorubicin-induced ROS increase, led to apoptosis — demonstrating collateral lethality exploiting ME2 loss. siRNA knockdown of ME3 in ME2-deficient PC cells, ROS measurement, apoptosis assay, targeted DNA nanostructure delivery in vivo Nano letters Low 39602246

Source papers

Stage 0 corpus · 50 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1985 Conjugal transfer from Streptococcus lactis ME2 of plasmids encoding phage resistance, nisin resistance and lactose-fermenting ability: evidence for a high-frequency conjugative plasmid responsible for abortive infection of virulent bacteriophage. Journal of general microbiology 88 3930657
1998 EWS/FLI1 up regulates mE2-C, a cyclin-selective ubiquitin conjugating enzyme involved in cyclin B destruction. Oncogene 82 9798675
2011 Me2-NHC based robust Ir catalyst for efficient water oxidation. Chemical communications (Cambridge, England) 73 21225028
1992 Molecular characterization of a second abortive phage resistance gene present in Lactococcus lactis subsp. lactis ME2. Journal of bacteriology 54 1429469
2023 SIRT5-mediated ME2 desuccinylation promotes cancer growth by enhancing mitochondrial respiration. Cell death and differentiation 52 38007551
2001 In vivo antitumor activity of bis(4,7-dimethyl-1,10-phenanthroline) sulfatooxovanadium(IV) (METVAN [VO(SO4)(Me2-Phen)2]). Clinical cancer research : an official journal of the American Association for Cancer Research 48 11448932
1988 Restriction and modification activities from Streptococcus lactis ME2 are encoded by a self-transmissible plasmid, pTN20, that forms cointegrates during mobilization of lactose-fermenting ability. Journal of bacteriology 44 2841286
2008 Combination of 2-methoxyestradiol (2-ME2) and eugenol for apoptosis induction synergistically in androgen independent prostate cancer cells. The Journal of steroid biochemistry and molecular biology 41 19084597
1994 Expression of basic-helix-loop-helix transcription factor ME2 during brain development and in the regions of neuronal plasticity in the adult brain. Brain research. Molecular brain research 40 7984047
1998 The effect of Me2+ cofactors at the initial stages of V(D)J recombination. The Journal of biological chemistry 38 9632694
2024 AKT1 phosphorylation of cytoplasmic ME2 induces a metabolic switch to glycolysis for tumorigenesis. Nature communications 35 38263319
2003 2-Methoxyestradiol exhibits a biphasic effect on VEGF-A in tumor cells and upregulation is mediated through ER-alpha: a possible signaling pathway associated with the impact of 2-ME2 on proliferative cells. Neoplasia (New York, N.Y.) 34 14670179
2015 Alkane metathesis with the tantalum methylidene [(≡SiO)Ta(═CH2)Me2]/[(≡SiO)2Ta(═CH2)Me] generated from well-defined surface organometallic complex [(≡SiO)Ta(V)Me4]. Journal of the American Chemical Society 32 25557135
1982 The structure of the EF-Tu . GDP . Me2+ complex. European journal of biochemistry 30 7200884
2021 Plant defense compound triggers mycotoxin synthesis by regulating H2B ub1 and H3K4 me2/3 deposition. The New phytologist 29 34480757
2015 Enhanced cytosolic NADP-ME2 activity in A. thaliana affects plant development, stress tolerance and specific diurnal and nocturnal cellular processes. Plant science : an international journal of experimental plant biology 29 26475199
2025 ACAT1-Mediated ME2 Acetylation Drives Chemoresistance in Ovarian Cancer by Linking Glutaminolysis to Lactate Production. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 27 39951294
1995 Differential expression and distinct DNA-binding specificity of ME1a and ME2 suggest a unique role during differentiation and neuronal plasticity. Brain research. Molecular brain research 27 7769987
2016 Sediment PAH source apportionment in the Liaohe River using the ME2 approach: A comparison to the PMF model. The Science of the total environment 26 26925728
2015 A small-molecule inhibitor suppresses the tumor-associated mitochondrial NAD(P)+-dependent malic enzyme (ME2) and induces cellular senescence. Oncotarget 24 26008970
2023 Inhibition of miR-214-3p attenuates ferroptosis in myocardial infarction via regulating ME2. Biochemical and biophysical research communications 22 37087800
2021 ME2 Promotes Proneural-Mesenchymal Transition and Lipogenesis in Glioblastoma. Frontiers in oncology 20 34381734
2017 Miz1 Controls Schwann Cell Proliferation via H3K36me2 Demethylase Kdm8 to Prevent Peripheral Nerve Demyelination. The Journal of neuroscience : the official journal of the Society for Neuroscience 20 29217679
2014 Discovery of a novel inhibitor of NAD(P)(+)-dependent malic enzyme (ME2) by high-throughput screening. Acta pharmacologica Sinica 18 24681895
2009 Mitochondrial malic enzyme (ME2) in pancreatic islets of the human, rat and mouse and clonal insulinoma cells. Archives of biochemistry and biophysics 18 19691144
1976 Me2+-(13 S) ATPase from Micrococcus sp. ATCC 398E. The effect of trypsin on the purified enzyme. Biochimica et biophysica acta 15 131581
2006 Innovative effect of illite on improved microbiological conversion of L-tyrosine to 3,4 dihydroxy phenyl L-alanine (L-DOPA) by Aspergillus oryzae ME2 under acidic reaction conditions. Current microbiology 13 17039388
2001 Apoptosis inducing novel anti-leukemic agent, bis(4,7-dimethyl-1,10 phenanthroline) sulfatooxovanadium(IV) [VO(SO4)(Me2-Phen)2] depolarizes mitochondrial membranes. Leukemia & lymphoma 13 11378580
1985 Linkage between the loci for mitochondrial malic enzyme (ME2) and coagulation factor XIIIA subunit (F13A). Human genetics 13 2860058
2015 Implementing constrained multi-time approach with bootstrap analysis in ME-2: An application to PM2.5 data from Florence (Italy). The Science of the total environment 12 26414851
2023 Targeting human mitochondrial NAD(P)+-dependent malic enzyme (ME2) impairs energy metabolism and redox state and exhibits antileukemic activity in acute myeloid leukemia. Cellular oncology (Dordrecht, Netherlands) 11 37079187
2024 NADP-malic Enzyme OsNADP-ME2 Modulates Plant Height Involving in Gibberellin Signaling in Rice. Rice (New York, N.Y.) 7 39152344
2016 The level and distribution pattern of HP1β in the embryonic brain correspond to those of H3K9me1/me2 but not of H3K9me3. Histochemistry and cell biology 7 26794325
2014 Dynamic effects dictate the mechanism and selectivity of dehydration-rearrangement reactions of protonated alcohols [Me2 (R)CCH(OH2 )Me](+) (R=Me, Et, iPr) in the gas phase. Chemistry (Weinheim an der Bergstrasse, Germany) 7 25179304
2023 Cucumber malate decarboxylase, CsNADP-ME2, functions in the balance of carbon and amino acid metabolism in fruit. Horticulture research 6 38077499
2022 Study on the Effect of Key Genes ME2 and adhE during Luzhou-Flavor Baijiu Brewing. Foods (Basel, Switzerland) 6 35267332
1982 Induction of 2H+/Me2+ exchange in rat-liver mitochondria. European journal of biochemistry 5 6293824
2024 ME2 Deficiency Is Associated With Recessive Neurodevelopmental Disorder. Clinical genetics 4 39401966
2019 Genome-wide identification of histone methylation (H3K9me2) and acetylation (H4K12ac) marks in two ecotypes of switchgrass (Panicum virgatum L.). BMC genomics 4 31438854
2015 Metabolic vulnerability in melanoma: a ME2 (me too) story. The Journal of investigative dermatology 4 25666673
2003 Xstir polymorphism and absence of sex linkage in Xenopus laevis ME2 gene. Folia biologica 4 12859020
2024 PRMT1-mediated methylation of ME2 promotes hepatocellular carcinoma growth by inhibiting ubiquitination. Cell death & disease 3 39528487
2023 ME2 Promotes Hepatocellular Carcinoma Cell Migration through Pyruvate. Metabolites 3 37110198
2018 Preparation and Properties of Nanoparticles, tRNA-Bivalent Metal Cation (Me2+) Complexes, and Prospects of Their Practical Use. Doklady. Biochemistry and biophysics 3 29779113
2024 A Targeted Octahedral DNA Nanostructure Co-delivers siME3 and Doxorubicin to Enhance Collateral Lethality in ME2-Deficient Pancreatic Cancer. Nano letters 1 39602246
2013 (N,N-diethyldithiocarbamato)[tris(3,5-dimethylpyrazol-1-yl)hydroborato]copper(II): a new copper(II) dithiocarbamate compound with the classic Tp(Me2) scorpionate. Acta crystallographica. Section C, Crystal structure communications 1 24005503
2025 CYP4F11 promotes lung cancer progression through the miR-195/ME2 pathway. Frontiers of medicine 0 41359237
2015 Long-term remission of therapy-related acute myeloid leukemia with a new t(11;18)(q23;q21.2) translocation and KMT2A-ME2 (MLL-ME2) fusion gene. Cancer genetics 0 26556690
2014 Competitive reaction pathways for the gas-phase reactivity of [Me2 AlNH2 ]3. Chemphyschem : a European journal of chemical physics and physical chemistry 0 24976567
1990 A population study of leukocyte enzymes (GOT2, ME2 and PGM3) in Galicia (NW Spain). Human heredity 0 2138119

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