LDHB

UniProt ID: P07195
Organism: Homo sapiens
Review Status: INITIALIZED
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Gene Description

LDHB encodes the B (also called H, "heart") subunit of L-lactate dehydrogenase (LDH; EC 1.1.1.27), a cytosolic NAD-dependent oxidoreductase. It catalyses the reversible, stereospecific interconversion of pyruvate + NADH + H+ and L-lactate + NAD+. Catalytically active LDH is a tetramer assembled in variable ratios from LDHA (M) and LDHB (H) subunits, producing the five classic isoenzymes (M4 through H4). LDHB-rich isoenzymes (H4) are kinetically tuned to favour the oxidation of L-lactate back to pyruvate and predominate in oxidative tissues such as heart and kidney that consume lactate as a metabolic fuel, whereas LDHA-rich isoenzymes favour the reduction of pyruvate to lactate during fermentative glycolysis. LDHB also participates in cytosolic NAD+/NADH redox balance. Loss-of-function LDHB variants cause lactate dehydrogenase B (H-subunit) deficiency, a generally benign condition of interest chiefly to laboratory medicine.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0019244 pyruvate fermentation to lactate
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Phylogenetic (IBA) annotation to the pyruvate-to-lactate fermentation process for the LDH family node. LDHB is a bona fide L-lactate dehydrogenase and catalyses this reversible reaction, so the annotation is biologically correct at the family level, but its named direction (pyruvate -> lactate, fermentation) is the direction favoured by the LDHA (M) subunit; the LDHB-rich H4 isoenzyme is kinetically tuned to the reverse (lactate -> pyruvate) direction in oxidative tissues.
Reason: The reaction is correctly assigned to the family, but for LDHB specifically the fermentation (lactate-producing) direction is not the physiologically dominant role; the lactate-oxidising direction is more characteristic. Keep as non-core; the core metabolic role is better captured by lactate/pyruvate metabolic process.
Supporting Evidence:
PMID:27618187
it has been suggested that LDHA preferentially reduces pyruvate to lactate, while LDHB supports conversion of lactate to pyruvate in cells that utilize lactate as a nutrient source for oxidative metabolism or gluconeogenesis
GO:0004459 L-lactate dehydrogenase (NAD+) activity
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic (IBA) assignment of the core catalytic molecular function, L-lactate dehydrogenase (NAD+) activity (EC 1.1.1.27). This is the defining activity of LDHB and is corroborated by direct enzymatic, structural, and orthology evidence.
Reason: This is the core molecular function of LDHB, strongly supported across experimental (PMID:11276087, PMID:27618187), phylogenetic, and electronic evidence.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
PMID:11276087
Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with concomitant
GO:0005739 mitochondrion
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Phylogenetic (IBA) annotation placing LDHB activity in the mitochondrion. A minority of experimental work (PMID:27618187) reports LDHB association with the inner mitochondrial membrane and mitochondrial LDH activity, but the canonical, dominant localization of LDH is cytosolic.
Reason: There is experimental support for a mitochondrial pool of LDHB, but this is a non-canonical, still-debated localization for a classically cytosolic glycolytic enzyme. Retain as non-core rather than a core location.
Supporting Evidence:
PMID:27618187
gold-labeled LDHB localizes to the inner mitochondrial membrane
GO:0003824 catalytic activity
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro2GO electronic mapping to the root catalytic activity term. Correct but far too general given the well-defined L-lactate dehydrogenase (NAD+) activity.
Reason: Uninformative parent of the specific enzymatic function; the precise MF term GO:0004459 is annotated with strong experimental evidence and should be preferred.
GO:0004459 L-lactate dehydrogenase (NAD+) activity
IEA
GO_REF:0000120
ACCEPT
Summary: Electronic (multi-method IEA) assignment of the core L-lactate dehydrogenase (NAD+) activity, consistent with the EC 1.1.1.27 / RHEA:23444 mapping and with experimental evidence.
Reason: Correct core molecular function, redundant with but consistent with the experimental and phylogenetic annotations of the same term.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
GO:0005737 cytoplasm
IEA
GO_REF:0000120
ACCEPT
Summary: Electronic assignment of the cytoplasm, consistent with UniProt subcellular location and with LDH being a soluble cytosolic glycolytic enzyme.
Reason: Correct, though the more specific cytosol (GO:0005829) is the preferred core location. Cytoplasm is a valid broader statement.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane
GO:0005743 mitochondrial inner membrane
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: Electronic annotation derived from the UniProt Subcellular Location keyword mapping (Mitochondrion inner membrane), which in turn rests on the experimental report PMID:27618187. This is a non-canonical localization for LDHB.
Reason: Supported experimentally (PMID:27618187) via the UniProt subcellular location, but a minority/debated localization; the core location is cytosolic. Keep as non-core.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane
GO:0016491 oxidoreductase activity
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro2GO electronic mapping to the general oxidoreductase activity term. Correct but too general.
Reason: Parent of the specific L-lactate dehydrogenase (NAD+) activity; uninformative given the well-supported specific MF term.
GO:0016616 oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro2GO electronic mapping to an intermediate oxidoreductase term describing the CH-OH-group / NAD(P)-acceptor chemistry. Accurate for LDH chemistry but less specific than L-lactate dehydrogenase (NAD+) activity.
Reason: Correct grandparent-level chemistry but redundant with the specific MF term GO:0004459, which should be preferred.
GO:0005515 protein binding
IPI
PMID:21044950
Genome-wide YFP fluorescence complementation screen identifi...
MARK AS OVER ANNOTATED
Summary: IntAct two-hybrid/interaction datum with TERF1 (P54274). Bare protein binding conveys no specific function.
Reason: Uninformative generic protein-binding annotation from a high-throughput interaction screen; not a core function and no specific molecular function can be inferred. Retained per policy rather than removed.
GO:0005515 protein binding
IPI
PMID:21988832
Toward an understanding of the protein interaction network o...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LDHA (P00338), consistent with formation of mixed M/H LDH tetramers, but captured only as generic protein binding.
Reason: The underlying LDHA interaction is biologically real (heterotetramer assembly), but the bare protein binding term is uninformative; the tetramer relationship is better captured by identical protein binding / oxidoreductase complex.
GO:0005515 protein binding
IPI
PMID:24725412
Ribosomal protein s15 phosphorylation mediates LRRK2 neurode...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LRRK2 (Q5S007) from a Parkinson-disease study. Bare protein binding, no specific function implied for LDHB.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0005515 protein binding
IPI
PMID:25416956
A proteome-scale map of the human interactome network.
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LDHA isoform (P00338-3) from a proteome-scale interactome map. Bare protein binding.
Reason: Uninformative generic protein-binding annotation from a high-throughput interactome study; not a core function.
GO:0005515 protein binding
IPI
PMID:25910212
Widespread macromolecular interaction perturbations in human...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LDHA isoform (P00338-3) from a study of interaction perturbations in genetic disorders. Bare protein binding.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0005515 protein binding
IPI
PMID:28514442
Architecture of the human interactome defines protein commun...
MARK AS OVER ANNOTATED
Summary: IntAct interaction data (LDHA P00338, LDHC P07864) from an interactome community study. Consistent with LDH-family tetramer assembly but recorded as generic protein binding.
Reason: Uninformative generic protein-binding term; the biologically relevant LDH-family subunit interactions are better captured elsewhere.
GO:0005515 protein binding
IPI
PMID:31515488
Extensive disruption of protein interactions by genetic vari...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LDHA (P00338) from a study of variant-driven interaction disruption. Bare protein binding.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0005515 protein binding
IPI
PMID:32296183
A reference map of the human binary protein interactome.
MARK AS OVER ANNOTATED
Summary: IntAct interaction data (SDCBP O00560, LDHA isoform P00338-3) from a reference binary interactome map. Bare protein binding.
Reason: Uninformative generic protein-binding annotation from a high-throughput screen; not a core function.
GO:0005515 protein binding
IPI
PMID:32814053
Interactome Mapping Provides a Network of Neurodegenerative ...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with HTT (P42858) from a neurodegenerative-disease interactome study. Bare protein binding.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
MARK AS OVER ANNOTATED
Summary: IntAct interaction data (LDHA P00338, LDHC P07864) from a cell-specific interactome remodeling study. LDH-family subunit interactions recorded as generic protein binding.
Reason: Uninformative generic protein-binding term; the relevant subunit interactions are better represented by the tetramer/complex annotations.
GO:0005515 protein binding
IPI
PMID:35271311
OpenCell: Endogenous tagging for the cartography of human ce...
MARK AS OVER ANNOTATED
Summary: IntAct interaction data (LDHA P00338, LDHC P07864) from the OpenCell endogenous-tagging interactome. Bare protein binding.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0005515 protein binding
IPI
PMID:40205054
Multimodal cell maps as a foundation for structural and func...
MARK AS OVER ANNOTATED
Summary: IntAct interaction datum with LDHC (P07864) from a multimodal cell-map study. Bare protein binding.
Reason: Uninformative generic protein-binding annotation; not a core function.
GO:0042802 identical protein binding
IPI
PMID:25416956
A proteome-scale map of the human interactome network.
KEEP AS NON CORE
Summary: IntAct self-interaction (P07195 with P07195), reflecting the homotetrameric assembly of the LDHB (H4) isoenzyme.
Reason: Biologically meaningful (LDHB homotetramerization) and more informative than bare protein binding, but the self-association is a structural feature supporting the complex rather than the core catalytic function. Keep as non-core.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0042802 identical protein binding
IPI
PMID:25502805
A massively parallel pipeline to clone DNA variants and exam...
KEEP AS NON CORE
Summary: IntAct self-interaction (P07195 with P07195) from a variant-cloning / molecular-phenotype pipeline, again reflecting LDHB homotetramerization.
Reason: Consistent with homotetramer assembly; informative structural annotation but not the core catalytic function.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0042802 identical protein binding
IPI
PMID:31515488
Extensive disruption of protein interactions by genetic vari...
KEEP AS NON CORE
Summary: IntAct self-interaction (P07195 with P07195), reflecting LDHB homotetramerization.
Reason: Consistent with homotetramer assembly; informative structural annotation but not the core catalytic function.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0042802 identical protein binding
IPI
PMID:32296183
A reference map of the human binary protein interactome.
KEEP AS NON CORE
Summary: IntAct self-interaction (P07195 with P07195) from the reference binary interactome, reflecting LDHB homotetramerization.
Reason: Consistent with homotetramer assembly; informative structural annotation but not the core catalytic function.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0004457 lactate dehydrogenase activity
IEA
GO_REF:0000107
ACCEPT
Summary: Ensembl orthology (IEA) assignment of lactate dehydrogenase activity, the immediate parent of the NAD+-specific L-lactate dehydrogenase activity.
Reason: Correct and consistent with the core function; a slightly broader phrasing of the same catalytic activity. Acceptable as-is.
Supporting Evidence:
PMID:11276087
Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with concomitant
GO:0005829 cytosol
IEA
GO_REF:0000107
ACCEPT
Summary: Ensembl orthology (IEA) placement of LDHB activity in the cytosol, the canonical location of this glycolytic enzyme, concordant with the HPA IDA and Reactome TAS annotations to cytosol.
Reason: Correct core subcellular location; strongly corroborated by IDA (HPA) and TAS (Reactome) evidence.
GO:0006089 lactate metabolic process
IEA
GO_REF:0000107
ACCEPT
Summary: Ensembl orthology (IEA) assignment to lactate metabolic process, capturing LDHB's role in the interconversion of lactate and pyruvate. Also independently supported by ComplexPortal IDA (PMID:15117937).
Reason: Accurately reflects the core biological process in which LDHB participates (lactate/pyruvate interconversion), and is the process best matching the LDHB-favoured lactate-oxidising direction.
Supporting Evidence:
PMID:27618187
LDHB supports conversion of lactate to pyruvate in cells that utilize lactate as a nutrient source for oxidative metabolism or gluconeogenesis
GO:0019674 NAD+ metabolic process
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Ensembl orthology (IEA) assignment to NAD+ metabolic process, reflecting that LDHB regenerates/consumes NAD+/NADH during the lactate-pyruvate reaction and thereby contributes to cytosolic redox balance.
Reason: Biologically correct (the reaction interconverts NAD+ and NADH), but NAD+ metabolism is a downstream consequence of the catalytic activity rather than the core annotated function. Keep as non-core.
Supporting Evidence:
PMID:15117937
central to NAD(+) regeneration
GO:0019900 kinase binding
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: Ensembl orthology (IEA) transfer of a kinase-binding annotation from a rat ortholog. There is no specific experimental evidence that human LDHB binds a kinase as a defined molecular function.
Reason: Orthology-transferred generic binding annotation with no clear functional relevance to LDHB's role; likely an over-annotation and not a core function.
GO:0042802 identical protein binding
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Ensembl orthology (IEA) transfer of identical protein binding, consistent with the homotetrameric assembly of LDHB.
Reason: Reflects LDHB homotetramerization; informative structural feature but not the core catalytic function.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0051287 NAD binding
IEA
GO_REF:0000107
ACCEPT
Summary: Ensembl orthology (IEA) assignment of NAD binding. LDHB is an NAD-dependent dehydrogenase with a Rossmann-fold NAD-binding domain and defined NAD(+)-binding residues, so this is a correct and informative molecular function.
Reason: Correct; NAD binding is a genuine, structurally documented molecular function of LDHB (NAD-binding Rossmann domain; NAD(+) binding site residues in UniProt).
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
GO:0005829 cytosol
IDA
GO_REF:0000052
ACCEPT
Summary: Direct immunofluorescence (HPA) localization of LDHB to the cytosol, the canonical location of this glycolytic enzyme.
Reason: Experimentally supported core subcellular location, concordant with electronic and TAS cytosol annotations.
GO:0004459 L-lactate dehydrogenase (NAD+) activity
EXP
PMID:11276087
Structural basis for altered activity of M- and H-isozyme fo...
ACCEPT
Summary: Experimental (structural/biophysical) demonstration of L-lactate dehydrogenase (NAD+) activity for human LDH-H, via crystallographic ternary complexes with NADH and substrate analog and kinetic characterization.
Reason: This is the defining, experimentally established core molecular function of LDHB, characterized structurally and kinetically for the H isoform.
Supporting Evidence:
PMID:11276087
Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with concomitant
PMID:11276087
the H form, found mainly in cardiac muscle
GO:0006089 lactate metabolic process
IDA
PMID:15117937
Identification and activity of a series of azole-based compo...
ACCEPT
Summary: ComplexPortal IDA annotation of the LDH complex to lactate metabolic process. LDH is central to lactate/pyruvate interconversion and NAD+ regeneration.
Reason: Correctly captures the core biological process; LDHB participates in lactate metabolism as the H subunit of the LDH complex.
Supporting Evidence:
PMID:15117937
central to NAD(+) regeneration
GO:1990204 oxidoreductase complex
IPI
PMID:15117937
Identification and activity of a series of azole-based compo...
ACCEPT
Summary: ComplexPortal assertion that LDHB is part of the L-lactate dehydrogenase (oxidoreductase) complex, i.e. the LDH tetramer. UniProt/ComplexPortal record the A/B (M/H) tetramer variants (CPX-6592/6594/6598/6599).
Reason: Correct; catalytically active LDH is a tetramer, and LDHB is a constituent subunit. This appropriately captures the complex-level context of the enzyme.
Supporting Evidence:
PMID:27618187
Functional LDH is a homo or hetero tetramer composed of LDHA and LDHB subunits
file:human/LDHB/LDHB-uniprot.txt
Homotetramer (PubMed:11276087)
GO:0004459 L-lactate dehydrogenase (NAD+) activity
IDA
PMID:27618187
Lactate metabolism is associated with mammalian mitochondria...
ACCEPT
Summary: Direct assay of LDHB (L-lactate dehydrogenase) activity in whole-cell and isolated mitochondrial lysates, confirming the core catalytic function.
Reason: Experimentally supported core molecular function of LDHB.
Supporting Evidence:
PMID:27618187
the mitochondria had higher LDHB activity, further suggesting that LDHB activity is concentrated in mitochondria
GO:0005515 protein binding
IPI
PMID:33406399
An Upstream Open Reading Frame in Phosphatase and Tensin Hom...
MARK AS OVER ANNOTATED
Summary: UniProt-curated IPI interaction with the PTEN-uORF micropeptide MP31 (C0HLV8). MP31 binds LDHB and inhibits mitochondrial LDH activity by competing for NAD+, acting as a regulatory circuit breaker of lactate-pyruvate conversion; the interaction is abolished by LDHB mutations D53A and R100A.
Reason: This is the most biologically meaningful of the protein-binding annotations (a specific, functionally characterized regulatory interaction), but the term itself is the uninformative generic protein binding. Retained (not removed) per policy; a more specific enzyme-inhibitor / MP31-binding characterization would be preferable.
Supporting Evidence:
PMID:33406399
limits lactate-pyruvate conversion in mitochondria by
PMID:33406399
competing with mitochondrial lactate dehydrogenase (mLDH) for
GO:0005743 mitochondrial inner membrane
IDA
PMID:27618187
Lactate metabolism is associated with mammalian mitochondria...
KEEP AS NON CORE
Summary: TEM immunogold localization of LDHB to the inner mitochondrial membrane in HeLa cells, with concordant mitochondrial LDH activity in isolated mitochondria. A specific but non-canonical localization for this classically cytosolic enzyme.
Reason: Experimentally supported (PMID:27618187), but a minority/debated localization; the dominant location of LDHB is cytosolic. Retain as non-core.
Supporting Evidence:
PMID:27618187
gold-labeled LDHB localizes to the inner mitochondrial membrane
GO:0045121 membrane raft
IDA
PMID:25204797
Flotillin-1 facilitates toll-like receptor 3 signaling in hu...
MARK AS OVER ANNOTATED
Summary: Localization to membrane raft reported in a flotillin-1/TLR3 endothelial signaling study that used SILAC proteomics of raft/caveolae fractions. LDHB is a soluble cytosolic enzyme and its recovery in raft proteomic fractions most likely reflects abundant-protein co-purification rather than a functional raft localization.
Reason: A soluble glycolytic enzyme is not expected to be a genuine membrane raft resident; this is likely a proteomic co-isolation artifact and an over-annotation.
Supporting Evidence:
PMID:25204797
Flotillin-1 and -2 are markers of membrane rafts
GO:0070062 extracellular exosome
HDA
PMID:23533145
In-depth proteomic analyses of exosomes isolated from expres...
MARK AS OVER ANNOTATED
Summary: High-throughput proteomic detection of LDHB in urinary/prostatic-secretion exosome preparations. Abundant cytosolic enzymes are commonly detected in exosome proteomes without evidence of a functional role there.
Reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization and not a core annotation.
GO:0016020 membrane
HDA
PMID:19946888
Defining the membrane proteome of NK cells.
MARK AS OVER ANNOTATED
Summary: High-throughput proteomic detection of LDHB in an NK-cell membrane proteome preparation. Non-specific membrane assignment for a soluble enzyme.
Reason: Generic membrane assignment for a cytosolic protein from a proteomic membrane-fraction study; likely co-isolation and an over-annotation.
GO:0070062 extracellular exosome
HDA
PMID:19056867
Large-scale proteomics and phosphoproteomics of urinary exos...
MARK AS OVER ANNOTATED
Summary: High-throughput proteomic detection of LDHB in urinary exosomes.
Reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization and not a core annotation.
GO:0070062 extracellular exosome
HDA
PMID:20458337
MHC class II-associated proteins in B-cell exosomes and pote...
MARK AS OVER ANNOTATED
Summary: High-throughput proteomic detection of LDHB in B-cell exosomes.
Reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization and not a core annotation.
GO:0005829 cytosol
TAS
Reactome:R-HSA-70510
ACCEPT
Summary: Reactome traceable-author-statement placing LDHB in the cytosol in the context of LDH tetramer oxidises LACT to PYR, the lactate-oxidising direction favoured by the LDHB-rich isoenzyme.
Reason: Correct core cytosolic location, consistent with IDA and IEA cytosol annotations and with the canonical glycolytic-enzyme localization.
GO:0005829 cytosol
TAS
Reactome:R-HSA-71849
ACCEPT
Summary: Reactome traceable-author-statement placing LDHB in the cytosol in the context of LDH tetramer reduces PYR to LACT.
Reason: Correct core cytosolic location, redundant with but consistent with the other cytosol annotations.
GO:0005737 cytoplasm
TAS
PMID:16130169
Proteomics of human umbilical vein endothelial cells applied...
ACCEPT
Summary: Traceable-author-statement (UniProt) cytoplasm localization derived from a HUVEC proteomics study. Consistent with the soluble cytoplasmic nature of LDHB.
Reason: Correct broader localization; the more specific cytosol is preferred as the core location.
Supporting Evidence:
file:human/LDHB/LDHB-uniprot.txt
SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane
GO:0004459 L-lactate dehydrogenase (NAD+) activity
TAS
PMID:8314553
Analysis of a genetic mutation in an electrophoretic variant...
ACCEPT
Summary: Traceable-author-statement of L-lactate dehydrogenase (NAD+) activity, from a study characterizing an electrophoretic LDH-B(H) subunit variant (K7E) with decreased heat stability and slightly reduced activity.
Reason: Correct core molecular function; the paper explicitly concerns the human LDH-B (H) subunit and its enzymatic activity.
Supporting Evidence:
PMID:8314553
An electrophoretic variant of the lactate dehydrogenase (LDH)-B(H) subunit was

Core Functions

Cytosolic NAD-dependent L-lactate dehydrogenase (H/B subunit) catalysing the reversible interconversion of pyruvate + NADH and L-lactate + NAD+; in LDHB-rich (H4) isoenzymes this favours oxidation of lactate to pyruvate in oxidative tissues.

Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • PMID:11276087
    Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with concomitant
  • file:human/LDHB/LDHB-uniprot.txt
    Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);

As a subunit of the tetrameric L-lactate dehydrogenase complex, LDHB assembles into homotetramers (H4) and heterotetramers with LDHA (M), forming the catalytically active oxidoreductase complex.

Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • PMID:27618187
    Functional LDH is a homo or hetero tetramer composed of LDHA and LDHB subunits
  • file:human/LDHB/LDHB-uniprot.txt
    Homotetramer (PubMed:11276087)

References

Gene Ontology annotation through association of InterPro records with GO terms
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Gene Ontology annotation based on curation of immunofluorescence data
Automatic transfer of experimentally verified manual GO annotation data to orthologs using Ensembl Compara
Combined Automated Annotation using Multiple IEA Methods
file:human/LDHB/LDHB-uniprot.txt
UniProtKB entry P07195 (LDHB_HUMAN)
Structural basis for altered activity of M- and H-isozyme forms of human lactate dehydrogenase.
  • Human LDH-H and LDH-M interconvert pyruvate and lactate with concomitant NADH/NAD+ interconversion; the H form is found mainly in cardiac muscle and is structurally indistinguishable at the active site from the M form.
Identification and activity of a series of azole-based compounds with lactate dehydrogenase-directed anti-malarial activity.
  • LDH is central to NAD+ regeneration during glycolysis coupled with homolactic fermentation.
Proteomics of human umbilical vein endothelial cells applied to etoposide-induced apoptosis.
Large-scale proteomics and phosphoproteomics of urinary exosomes.
Defining the membrane proteome of NK cells.
MHC class II-associated proteins in B-cell exosomes and potential functional implications for exosome biogenesis.
Genome-wide YFP fluorescence complementation screen identifies new regulators for telomere signaling in human cells.
Toward an understanding of the protein interaction network of the human liver.
In-depth proteomic analyses of exosomes isolated from expressed prostatic secretions in urine.
Ribosomal protein s15 phosphorylation mediates LRRK2 neurodegeneration in Parkinson's disease.
Flotillin-1 facilitates toll-like receptor 3 signaling in human endothelial cells.
  • Flotillins are membrane raft markers; LDHB was recovered in the raft/caveolae proteomic analysis of this study.
A proteome-scale map of the human interactome network.
A massively parallel pipeline to clone DNA variants and examine molecular phenotypes of human disease mutations.
Widespread macromolecular interaction perturbations in human genetic disorders.
Lactate metabolism is associated with mammalian mitochondria.
  • Functional LDH is a tetramer of LDHA and LDHB subunits; LDHB is the predominant heart (H) subunit and supports conversion of lactate to pyruvate in lactate-consuming oxidative cells. LDHB was localized by TEM immunogold to the inner mitochondrial membrane and mitochondrial lysates showed LDHB activity.
Architecture of the human interactome defines protein communities and disease networks.
Extensive disruption of protein interactions by genetic variants across the allele frequency spectrum in human populations.
A reference map of the human binary protein interactome.
Interactome Mapping Provides a Network of Neurodegenerative Disease Proteins and Uncovers Widespread Protein Aggregation in Affected Brains.
An Upstream Open Reading Frame in Phosphatase and Tensin Homolog Encodes a Circuit Breaker of Lactate Metabolism.
  • MP31, a micropeptide from the PTEN uORF, binds LDHB and limits lactate-pyruvate conversion in mitochondria by competing with mitochondrial LDH for NAD+.
Dual proteome-scale networks reveal cell-specific remodeling of the human interactome.
OpenCell: Endogenous tagging for the cartography of human cellular organization.
Multimodal cell maps as a foundation for structural and functional genomics.
Analysis of a genetic mutation in an electrophoretic variant of the human lactate dehydrogenase-B(H) subunit.
  • An electrophoretic LDH-B(H) subunit variant (K7E) with decreased heat stability and slightly lower LDH activity was characterized, confirming LDHB as the H subunit of human LDH.
Reactome:R-HSA-70510
LDH tetramer oxidises LACT to PYR
Reactome:R-HSA-71849
LDH tetramer reduces PYR to LACT

Suggested Questions for Experts

Q: Is the mitochondrial (inner-membrane) pool of LDHB a genuine, functionally significant localization in vivo, or is it largely restricted to specific cell types (e.g. fermenting cancer cells) and experimental conditions?

Q: To what extent does the MP31 (PTEN-uORF) circuit-breaker regulation of LDHB operate in normal physiology versus tumour metabolism?

Suggested Experiments

Experiment: Compartment-resolved activity assays (cytosol vs mitochondrial sub-fractions) across oxidative and glycolytic human tissues to quantify the physiological contribution of any mitochondrial LDHB pool.

Experiment: Structure-guided kinetic comparison of purified LDHB homotetramer versus defined M/H heterotetramers to quantify the directional (lactate to pyruvate) preference under physiological substrate/redox conditions.

📚 Additional Documentation

Notes

(LDHB-notes.md)

LDHB (L-lactate dehydrogenase B chain, P07195) — review notes

Core biology

LDHB encodes the B / H ("heart") subunit of L-lactate dehydrogenase (LDH; EC 1.1.1.27).
Active LDH is a tetramer assembled from LDHA (M) and LDHB (H) subunits, giving rise to
the classic five isoenzymes (M4, M3H1, M2H2, M1H3, H4). LDHB-rich isoenzymes (H4)
kinetically favour oxidation of L-lactate back to pyruvate, suiting oxidative tissues
(heart, kidney) that consume lactate as fuel.

  • Catalytic reaction (UniProt CATALYTIC ACTIVITY, RHEA:23444, EC 1.1.1.27):
    (S)-lactate + NAD(+) = pyruvate + NADH + H(+). Reversible; UniProt notes both
    left-to-right (RHEA:23445) and right-to-left (RHEA:23446) physiological directions.
  • UniProt FUNCTION [ECO:0000269|PubMed:27618187]: "Interconverts simultaneously and
    stereospecifically pyruvate and lactate with concomitant interconversion of NADH and
    NAD(+)."
  • SUBUNIT: Homotetramer (PubMed:11276087). Interacts with the PTEN-uORF micropeptide
    MP31, which inhibits mitochondrial LDH activity (PubMed:33406399).
  • Tissue specificity: "Predominantly expressed in aerobic tissues such as cardiac
    muscle" [ECO:0000305|PubMed:11276087]. HPA: tissue-enhanced heart muscle, kidney.

Kinetic / structural basis (PMID:11276087)

Read et al. crystallised human LDH-M and LDH-H ternary complexes with NADH + oxamate.
"Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with concomitant
interconversion of NADH and NAD(+)." The two isoforms are structurally
indistinguishable at the active site; kinetic differences arise from peripheral charged
surface residues. Confirms homotetramer, NAD+ binding, and MF = L-lactate dehydrogenase
(NAD+) activity. [full_text_available: false — abstract only]

Mitochondrial association (PMID:27618187, Chen et al. Nat Chem Biol 2016)

Full text available. Key verbatim findings:
- "Functional LDH is a homo or hetero tetramer composed of LDHA and LDHB subunits"
- "LDHB is the predominant isoform found in heart muscle and is often referred to as the
H subunit"
- "it has been suggested that LDHA preferentially reduces pyruvate to lactate, while
LDHB supports conversion of lactate to pyruvate in cells that utilize lactate as a
nutrient source for oxidative metabolism or gluconeogenesis"
- TEM immunogold: "gold-labeled LDHB localizes to the inner mitochondrial membrane"
- LDHB activity detected in whole-cell and isolated mitochondrial lysates.
This is the experimental basis for the IDA GO:0004459 and IDA GO:0005743 (mito inner
membrane) annotations. Note this is a specific, somewhat contested localization; the
dominant/canonical localization remains cytosolic (glycolytic enzyme).

MP31 interaction (PMID:33406399)

MP31 is a micropeptide from the PTEN uORF that "limits lactate-pyruvate conversion in
mitochondria by competing with mitochondrial lactate dehydrogenase (mLDH) for
nicotinamide adenine dinucleotide (NAD+)." UniProt: interaction abolished by LDHB
mutants D53A and R100A. Supports a real, functionally meaningful protein-protein
interaction (regulatory), curated by UniProt as GO:0005515 IPI with C0HLV8 (MP31).

Disease

LDHB deficiency (LDHBD, MIM:614128): "A condition with no deleterious effects on
health." Clinically mild/asymptomatic; laboratory-medicine interest (LDH isoenzyme
misdiagnosis). Many missense variants documented in UniProt (K7E, R107W inactive, etc.).

Annotation assessment summary

  • MF core: GO:0004459 L-lactate dehydrogenase (NAD+) activity — strongly supported
    (EXP PMID:11276087, IDA PMID:27618187, IBA, IEA, TAS). ACCEPT.
  • GO:0004457 lactate dehydrogenase activity — parent term, ACCEPT (broader is fine).
  • BP: GO:0006089 lactate metabolic process, GO:0042867 pyruvate catabolic /
    GO:0019244 pyruvate fermentation to lactate, GO:0019674 NAD+ metabolic process —
    all consistent. IBA pyruvate fermentation to lactate reflects the PANTHER family node;
    for LDHB (H4) the physiologically favoured direction is lactate->pyruvate, so
    "fermentation to lactate" is somewhat directionally biased toward LDHA, but the
    reaction is the same and reversible — keep, mark non-core where over-specific.
  • CC: cytosol GO:0005829 (IDA HPA, IEA, TAS Reactome) — core canonical location.
    cytoplasm GO:0005737 — ACCEPT (broader). mitochondrion / mito inner membrane
    (IBA + IDA PMID:27618187) — real but non-canonical; keep as non-core. exosome /
    membrane / membrane raft — mass-spec / proteomics HDA/IDA; over-annotations for a
    soluble glycolytic enzyme, mark over-annotated.
  • protein binding (GO:0005515) IPIs — bare, uninformative; MARK_AS_OVER_ANNOTATED
    (do NOT REMOVE per policy). The MP31 one (PMID:33406399) is the most biologically
    meaningful.
  • identical protein binding GO:0042802 — reflects homotetramerization; keep as
    non-core (informative-ish, supports tetramer).
  • kinase binding / NAD binding (Ensembl IEA GO_REF:0000107) — NAD binding is correct
    (NAD-dependent enzyme); kinase binding is an orthology-transferred over-annotation,
    mark over-annotated.

📄 View Raw YAML

id: P07195
gene_symbol: LDHB
product_type: PROTEIN
status: INITIALIZED
taxon:
  id: NCBITaxon:9606
  label: Homo sapiens
description: LDHB encodes the B (also called H, "heart") subunit of L-lactate dehydrogenase
  (LDH; EC 1.1.1.27), a cytosolic NAD-dependent oxidoreductase. It catalyses the reversible,
  stereospecific interconversion of pyruvate + NADH + H+ and L-lactate + NAD+. Catalytically
  active LDH is a tetramer assembled in variable ratios from LDHA (M) and LDHB (H) subunits,
  producing the five classic isoenzymes (M4 through H4). LDHB-rich isoenzymes (H4) are
  kinetically tuned to favour the oxidation of L-lactate back to pyruvate and predominate
  in oxidative tissues such as heart and kidney that consume lactate as a metabolic fuel,
  whereas LDHA-rich isoenzymes favour the reduction of pyruvate to lactate during
  fermentative glycolysis. LDHB also participates in cytosolic NAD+/NADH redox balance.
  Loss-of-function LDHB variants cause lactate dehydrogenase B (H-subunit) deficiency, a
  generally benign condition of interest chiefly to laboratory medicine.
existing_annotations:
- term:
    id: GO:0019244
    label: pyruvate fermentation to lactate
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: Phylogenetic (IBA) annotation to the pyruvate-to-lactate fermentation process
      for the LDH family node. LDHB is a bona fide L-lactate dehydrogenase and catalyses
      this reversible reaction, so the annotation is biologically correct at the family
      level, but its named direction (pyruvate -> lactate, fermentation) is the direction
      favoured by the LDHA (M) subunit; the LDHB-rich H4 isoenzyme is kinetically tuned to
      the reverse (lactate -> pyruvate) direction in oxidative tissues.
    action: KEEP_AS_NON_CORE
    reason: The reaction is correctly assigned to the family, but for LDHB specifically the
      fermentation (lactate-producing) direction is not the physiologically dominant role;
      the lactate-oxidising direction is more characteristic. Keep as non-core; the core
      metabolic role is better captured by lactate/pyruvate metabolic process.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: it has been suggested that LDHA preferentially reduces pyruvate to
        lactate, while LDHB supports conversion of lactate to pyruvate in cells that utilize
        lactate as a nutrient source for oxidative metabolism or gluconeogenesis
- term:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: enables
  review:
    summary: Phylogenetic (IBA) assignment of the core catalytic molecular function,
      L-lactate dehydrogenase (NAD+) activity (EC 1.1.1.27). This is the defining activity
      of LDHB and is corroborated by direct enzymatic, structural, and orthology evidence.
    action: ACCEPT
    reason: This is the core molecular function of LDHB, strongly supported across
      experimental (PMID:11276087, PMID:27618187), phylogenetic, and electronic evidence.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
    - reference_id: PMID:11276087
      supporting_text: Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with
        concomitant
- term:
    id: GO:0005739
    label: mitochondrion
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: is_active_in
  review:
    summary: Phylogenetic (IBA) annotation placing LDHB activity in the mitochondrion. A
      minority of experimental work (PMID:27618187) reports LDHB association with the inner
      mitochondrial membrane and mitochondrial LDH activity, but the canonical, dominant
      localization of LDH is cytosolic.
    action: KEEP_AS_NON_CORE
    reason: There is experimental support for a mitochondrial pool of LDHB, but this is a
      non-canonical, still-debated localization for a classically cytosolic glycolytic
      enzyme. Retain as non-core rather than a core location.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: gold-labeled LDHB localizes to the inner mitochondrial membrane
- term:
    id: GO:0003824
    label: catalytic activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: InterPro2GO electronic mapping to the root catalytic activity term. Correct
      but far too general given the well-defined L-lactate dehydrogenase (NAD+) activity.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative parent of the specific enzymatic function; the precise MF term
      GO:0004459 is annotated with strong experimental evidence and should be preferred.
- term:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  qualifier: enables
  review:
    summary: Electronic (multi-method IEA) assignment of the core L-lactate dehydrogenase
      (NAD+) activity, consistent with the EC 1.1.1.27 / RHEA:23444 mapping and with
      experimental evidence.
    action: ACCEPT
    reason: Correct core molecular function, redundant with but consistent with the
      experimental and phylogenetic annotations of the same term.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  qualifier: located_in
  review:
    summary: Electronic assignment of the cytoplasm, consistent with UniProt subcellular
      location and with LDH being a soluble cytosolic glycolytic enzyme.
    action: ACCEPT
    reason: Correct, though the more specific cytosol (GO:0005829) is the preferred core
      location. Cytoplasm is a valid broader statement.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: 'SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane'
- term:
    id: GO:0005743
    label: mitochondrial inner membrane
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: Electronic annotation derived from the UniProt Subcellular Location keyword
      mapping (Mitochondrion inner membrane), which in turn rests on the experimental
      report PMID:27618187. This is a non-canonical localization for LDHB.
    action: KEEP_AS_NON_CORE
    reason: Supported experimentally (PMID:27618187) via the UniProt subcellular location,
      but a minority/debated localization; the core location is cytosolic. Keep as
      non-core.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: 'SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane'
- term:
    id: GO:0016491
    label: oxidoreductase activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: InterPro2GO electronic mapping to the general oxidoreductase activity term.
      Correct but too general.
    action: MARK_AS_OVER_ANNOTATED
    reason: Parent of the specific L-lactate dehydrogenase (NAD+) activity; uninformative
      given the well-supported specific MF term.
- term:
    id: GO:0016616
    label: oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP
      as acceptor
  evidence_type: IEA
  original_reference_id: GO_REF:0000002
  qualifier: enables
  review:
    summary: InterPro2GO electronic mapping to an intermediate oxidoreductase term
      describing the CH-OH-group / NAD(P)-acceptor chemistry. Accurate for LDH chemistry
      but less specific than L-lactate dehydrogenase (NAD+) activity.
    action: MARK_AS_OVER_ANNOTATED
    reason: Correct grandparent-level chemistry but redundant with the specific MF term
      GO:0004459, which should be preferred.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21044950
  qualifier: enables
  review:
    summary: IntAct two-hybrid/interaction datum with TERF1 (P54274). Bare protein binding
      conveys no specific function.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation from a high-throughput
      interaction screen; not a core function and no specific molecular function can be
      inferred. Retained per policy rather than removed.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21988832
  qualifier: enables
  review:
    summary: IntAct interaction datum with LDHA (P00338), consistent with formation of
      mixed M/H LDH tetramers, but captured only as generic protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: The underlying LDHA interaction is biologically real (heterotetramer assembly),
      but the bare protein binding term is uninformative; the tetramer relationship is
      better captured by identical protein binding / oxidoreductase complex.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:24725412
  qualifier: enables
  review:
    summary: IntAct interaction datum with LRRK2 (Q5S007) from a Parkinson-disease study.
      Bare protein binding, no specific function implied for LDHB.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25416956
  qualifier: enables
  review:
    summary: IntAct interaction datum with LDHA isoform (P00338-3) from a proteome-scale
      interactome map. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation from a high-throughput
      interactome study; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25910212
  qualifier: enables
  review:
    summary: IntAct interaction datum with LDHA isoform (P00338-3) from a study of
      interaction perturbations in genetic disorders. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:28514442
  qualifier: enables
  review:
    summary: IntAct interaction data (LDHA P00338, LDHC P07864) from an interactome
      community study. Consistent with LDH-family tetramer assembly but recorded as
      generic protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding term; the biologically relevant LDH-family
      subunit interactions are better captured elsewhere.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:31515488
  qualifier: enables
  review:
    summary: IntAct interaction datum with LDHA (P00338) from a study of variant-driven
      interaction disruption. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:32296183
  qualifier: enables
  review:
    summary: IntAct interaction data (SDCBP O00560, LDHA isoform P00338-3) from a reference
      binary interactome map. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation from a high-throughput screen;
      not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:32814053
  qualifier: enables
  review:
    summary: IntAct interaction datum with HTT (P42858) from a neurodegenerative-disease
      interactome study. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:33961781
  qualifier: enables
  review:
    summary: IntAct interaction data (LDHA P00338, LDHC P07864) from a cell-specific
      interactome remodeling study. LDH-family subunit interactions recorded as generic
      protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding term; the relevant subunit interactions
      are better represented by the tetramer/complex annotations.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:35271311
  qualifier: enables
  review:
    summary: IntAct interaction data (LDHA P00338, LDHC P07864) from the OpenCell
      endogenous-tagging interactome. Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:40205054
  qualifier: enables
  review:
    summary: IntAct interaction datum with LDHC (P07864) from a multimodal cell-map study.
      Bare protein binding.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative generic protein-binding annotation; not a core function.
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:25416956
  qualifier: enables
  review:
    summary: IntAct self-interaction (P07195 with P07195), reflecting the homotetrameric
      assembly of the LDHB (H4) isoenzyme.
    action: KEEP_AS_NON_CORE
    reason: Biologically meaningful (LDHB homotetramerization) and more informative than
      bare protein binding, but the self-association is a structural feature supporting the
      complex rather than the core catalytic function. Keep as non-core.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:25502805
  qualifier: enables
  review:
    summary: IntAct self-interaction (P07195 with P07195) from a variant-cloning /
      molecular-phenotype pipeline, again reflecting LDHB homotetramerization.
    action: KEEP_AS_NON_CORE
    reason: Consistent with homotetramer assembly; informative structural annotation but
      not the core catalytic function.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:31515488
  qualifier: enables
  review:
    summary: IntAct self-interaction (P07195 with P07195), reflecting LDHB
      homotetramerization.
    action: KEEP_AS_NON_CORE
    reason: Consistent with homotetramer assembly; informative structural annotation but
      not the core catalytic function.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:32296183
  qualifier: enables
  review:
    summary: IntAct self-interaction (P07195 with P07195) from the reference binary
      interactome, reflecting LDHB homotetramerization.
    action: KEEP_AS_NON_CORE
    reason: Consistent with homotetramer assembly; informative structural annotation but
      not the core catalytic function.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0004457
    label: lactate dehydrogenase activity
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: enables
  review:
    summary: Ensembl orthology (IEA) assignment of lactate dehydrogenase activity, the
      immediate parent of the NAD+-specific L-lactate dehydrogenase activity.
    action: ACCEPT
    reason: Correct and consistent with the core function; a slightly broader phrasing of
      the same catalytic activity. Acceptable as-is.
    supported_by:
    - reference_id: PMID:11276087
      supporting_text: Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with
        concomitant
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: is_active_in
  review:
    summary: Ensembl orthology (IEA) placement of LDHB activity in the cytosol, the
      canonical location of this glycolytic enzyme, concordant with the HPA IDA and
      Reactome TAS annotations to cytosol.
    action: ACCEPT
    reason: Correct core subcellular location; strongly corroborated by IDA (HPA) and TAS
      (Reactome) evidence.
- term:
    id: GO:0006089
    label: lactate metabolic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: involved_in
  review:
    summary: Ensembl orthology (IEA) assignment to lactate metabolic process, capturing
      LDHB's role in the interconversion of lactate and pyruvate. Also independently
      supported by ComplexPortal IDA (PMID:15117937).
    action: ACCEPT
    reason: Accurately reflects the core biological process in which LDHB participates
      (lactate/pyruvate interconversion), and is the process best matching the LDHB-favoured
      lactate-oxidising direction.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: LDHB supports conversion of lactate to pyruvate in cells that utilize
        lactate as a nutrient source for oxidative metabolism or gluconeogenesis
- term:
    id: GO:0019674
    label: NAD+ metabolic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: involved_in
  review:
    summary: Ensembl orthology (IEA) assignment to NAD+ metabolic process, reflecting that
      LDHB regenerates/consumes NAD+/NADH during the lactate-pyruvate reaction and thereby
      contributes to cytosolic redox balance.
    action: KEEP_AS_NON_CORE
    reason: Biologically correct (the reaction interconverts NAD+ and NADH), but NAD+
      metabolism is a downstream consequence of the catalytic activity rather than the core
      annotated function. Keep as non-core.
    supported_by:
    - reference_id: PMID:15117937
      supporting_text: central to NAD(+) regeneration
- term:
    id: GO:0019900
    label: kinase binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: enables
  review:
    summary: Ensembl orthology (IEA) transfer of a kinase-binding annotation from a rat
      ortholog. There is no specific experimental evidence that human LDHB binds a kinase
      as a defined molecular function.
    action: MARK_AS_OVER_ANNOTATED
    reason: Orthology-transferred generic binding annotation with no clear functional
      relevance to LDHB's role; likely an over-annotation and not a core function.
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: enables
  review:
    summary: Ensembl orthology (IEA) transfer of identical protein binding, consistent with
      the homotetrameric assembly of LDHB.
    action: KEEP_AS_NON_CORE
    reason: Reflects LDHB homotetramerization; informative structural feature but not the
      core catalytic function.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0051287
    label: NAD binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  qualifier: enables
  review:
    summary: Ensembl orthology (IEA) assignment of NAD binding. LDHB is an NAD-dependent
      dehydrogenase with a Rossmann-fold NAD-binding domain and defined NAD(+)-binding
      residues, so this is a correct and informative molecular function.
    action: ACCEPT
    reason: Correct; NAD binding is a genuine, structurally documented molecular function
      of LDHB (NAD-binding Rossmann domain; NAD(+) binding site residues in UniProt).
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: GO_REF:0000052
  qualifier: located_in
  review:
    summary: Direct immunofluorescence (HPA) localization of LDHB to the cytosol, the
      canonical location of this glycolytic enzyme.
    action: ACCEPT
    reason: Experimentally supported core subcellular location, concordant with electronic
      and TAS cytosol annotations.
- term:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  evidence_type: EXP
  original_reference_id: PMID:11276087
  qualifier: enables
  review:
    summary: Experimental (structural/biophysical) demonstration of L-lactate dehydrogenase
      (NAD+) activity for human LDH-H, via crystallographic ternary complexes with NADH and
      substrate analog and kinetic characterization.
    action: ACCEPT
    reason: This is the defining, experimentally established core molecular function of
      LDHB, characterized structurally and kinetically for the H isoform.
    supported_by:
    - reference_id: PMID:11276087
      supporting_text: Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with
        concomitant
    - reference_id: PMID:11276087
      supporting_text: the H form, found mainly in cardiac muscle
- term:
    id: GO:0006089
    label: lactate metabolic process
  evidence_type: IDA
  original_reference_id: PMID:15117937
  qualifier: involved_in
  review:
    summary: ComplexPortal IDA annotation of the LDH complex to lactate metabolic process.
      LDH is central to lactate/pyruvate interconversion and NAD+ regeneration.
    action: ACCEPT
    reason: Correctly captures the core biological process; LDHB participates in lactate
      metabolism as the H subunit of the LDH complex.
    supported_by:
    - reference_id: PMID:15117937
      supporting_text: central to NAD(+) regeneration
- term:
    id: GO:1990204
    label: oxidoreductase complex
  evidence_type: IPI
  original_reference_id: PMID:15117937
  qualifier: part_of
  review:
    summary: ComplexPortal assertion that LDHB is part of the L-lactate dehydrogenase
      (oxidoreductase) complex, i.e. the LDH tetramer. UniProt/ComplexPortal record the
      A/B (M/H) tetramer variants (CPX-6592/6594/6598/6599).
    action: ACCEPT
    reason: Correct; catalytically active LDH is a tetramer, and LDHB is a constituent
      subunit. This appropriately captures the complex-level context of the enzyme.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: Functional LDH is a homo or hetero tetramer composed of LDHA and
        LDHB subunits
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: Homotetramer (PubMed:11276087)
- term:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  evidence_type: IDA
  original_reference_id: PMID:27618187
  qualifier: enables
  review:
    summary: Direct assay of LDHB (L-lactate dehydrogenase) activity in whole-cell and
      isolated mitochondrial lysates, confirming the core catalytic function.
    action: ACCEPT
    reason: Experimentally supported core molecular function of LDHB.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: the mitochondria had higher LDHB activity, further suggesting that
        LDHB activity is concentrated in mitochondria
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:33406399
  qualifier: enables
  review:
    summary: UniProt-curated IPI interaction with the PTEN-uORF micropeptide MP31 (C0HLV8).
      MP31 binds LDHB and inhibits mitochondrial LDH activity by competing for NAD+, acting
      as a regulatory circuit breaker of lactate-pyruvate conversion; the interaction is
      abolished by LDHB mutations D53A and R100A.
    action: MARK_AS_OVER_ANNOTATED
    reason: This is the most biologically meaningful of the protein-binding annotations
      (a specific, functionally characterized regulatory interaction), but the term itself
      is the uninformative generic protein binding. Retained (not removed) per policy; a
      more specific enzyme-inhibitor / MP31-binding characterization would be preferable.
    supported_by:
    - reference_id: PMID:33406399
      supporting_text: limits lactate-pyruvate conversion in mitochondria by
    - reference_id: PMID:33406399
      supporting_text: competing with mitochondrial lactate dehydrogenase (mLDH) for
- term:
    id: GO:0005743
    label: mitochondrial inner membrane
  evidence_type: IDA
  original_reference_id: PMID:27618187
  qualifier: located_in
  review:
    summary: TEM immunogold localization of LDHB to the inner mitochondrial membrane in
      HeLa cells, with concordant mitochondrial LDH activity in isolated mitochondria. A
      specific but non-canonical localization for this classically cytosolic enzyme.
    action: KEEP_AS_NON_CORE
    reason: Experimentally supported (PMID:27618187), but a minority/debated localization;
      the dominant location of LDHB is cytosolic. Retain as non-core.
    supported_by:
    - reference_id: PMID:27618187
      supporting_text: gold-labeled LDHB localizes to the inner mitochondrial membrane
- term:
    id: GO:0045121
    label: membrane raft
  evidence_type: IDA
  original_reference_id: PMID:25204797
  qualifier: located_in
  review:
    summary: Localization to membrane raft reported in a flotillin-1/TLR3 endothelial
      signaling study that used SILAC proteomics of raft/caveolae fractions. LDHB is a
      soluble cytosolic enzyme and its recovery in raft proteomic fractions most likely
      reflects abundant-protein co-purification rather than a functional raft localization.
    action: MARK_AS_OVER_ANNOTATED
    reason: A soluble glycolytic enzyme is not expected to be a genuine membrane raft
      resident; this is likely a proteomic co-isolation artifact and an over-annotation.
    supported_by:
    - reference_id: PMID:25204797
      supporting_text: Flotillin-1 and -2 are markers of membrane rafts
- term:
    id: GO:0070062
    label: extracellular exosome
  evidence_type: HDA
  original_reference_id: PMID:23533145
  qualifier: located_in
  review:
    summary: High-throughput proteomic detection of LDHB in urinary/prostatic-secretion
      exosome preparations. Abundant cytosolic enzymes are commonly detected in exosome
      proteomes without evidence of a functional role there.
    action: MARK_AS_OVER_ANNOTATED
    reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization
      and not a core annotation.
- term:
    id: GO:0016020
    label: membrane
  evidence_type: HDA
  original_reference_id: PMID:19946888
  qualifier: located_in
  review:
    summary: High-throughput proteomic detection of LDHB in an NK-cell membrane proteome
      preparation. Non-specific membrane assignment for a soluble enzyme.
    action: MARK_AS_OVER_ANNOTATED
    reason: Generic membrane assignment for a cytosolic protein from a proteomic
      membrane-fraction study; likely co-isolation and an over-annotation.
- term:
    id: GO:0070062
    label: extracellular exosome
  evidence_type: HDA
  original_reference_id: PMID:19056867
  qualifier: located_in
  review:
    summary: High-throughput proteomic detection of LDHB in urinary exosomes.
    action: MARK_AS_OVER_ANNOTATED
    reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization
      and not a core annotation.
- term:
    id: GO:0070062
    label: extracellular exosome
  evidence_type: HDA
  original_reference_id: PMID:20458337
  qualifier: located_in
  review:
    summary: High-throughput proteomic detection of LDHB in B-cell exosomes.
    action: MARK_AS_OVER_ANNOTATED
    reason: Cytosolic-enzyme detection in exosome proteomics; not a functional localization
      and not a core annotation.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-70510
  qualifier: located_in
  review:
    summary: Reactome traceable-author-statement placing LDHB in the cytosol in the context
      of LDH tetramer oxidises LACT to PYR, the lactate-oxidising direction favoured by the
      LDHB-rich isoenzyme.
    action: ACCEPT
    reason: Correct core cytosolic location, consistent with IDA and IEA cytosol
      annotations and with the canonical glycolytic-enzyme localization.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: TAS
  original_reference_id: Reactome:R-HSA-71849
  qualifier: located_in
  review:
    summary: Reactome traceable-author-statement placing LDHB in the cytosol in the context
      of LDH tetramer reduces PYR to LACT.
    action: ACCEPT
    reason: Correct core cytosolic location, redundant with but consistent with the other
      cytosol annotations.
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: TAS
  original_reference_id: PMID:16130169
  qualifier: located_in
  review:
    summary: Traceable-author-statement (UniProt) cytoplasm localization derived from a
      HUVEC proteomics study. Consistent with the soluble cytoplasmic nature of LDHB.
    action: ACCEPT
    reason: Correct broader localization; the more specific cytosol is preferred as the
      core location.
    supported_by:
    - reference_id: file:human/LDHB/LDHB-uniprot.txt
      supporting_text: 'SUBCELLULAR LOCATION: Cytoplasm. Mitochondrion inner membrane'
- term:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  evidence_type: TAS
  original_reference_id: PMID:8314553
  qualifier: enables
  review:
    summary: Traceable-author-statement of L-lactate dehydrogenase (NAD+) activity, from a
      study characterizing an electrophoretic LDH-B(H) subunit variant (K7E) with decreased
      heat stability and slightly reduced activity.
    action: ACCEPT
    reason: Correct core molecular function; the paper explicitly concerns the human LDH-B
      (H) subunit and its enzymatic activity.
    supported_by:
    - reference_id: PMID:8314553
      supporting_text: An electrophoretic variant of the lactate dehydrogenase (LDH)-B(H)
        subunit was
core_functions:
- description: Cytosolic NAD-dependent L-lactate dehydrogenase (H/B subunit) catalysing the
    reversible interconversion of pyruvate + NADH and L-lactate + NAD+; in LDHB-rich (H4)
    isoenzymes this favours oxidation of lactate to pyruvate in oxidative tissues.
  molecular_function:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  directly_involved_in:
  - id: GO:0006089
    label: lactate metabolic process
  locations:
  - id: GO:0005829
    label: cytosol
  supported_by:
  - reference_id: PMID:11276087
    supporting_text: Lactate dehydrogenase (LDH) interconverts pyruvate and lactate with
      concomitant
  - reference_id: file:human/LDHB/LDHB-uniprot.txt
    supporting_text: Reaction=(S)-lactate + NAD(+) = pyruvate + NADH + H(+);
- description: As a subunit of the tetrameric L-lactate dehydrogenase complex, LDHB
    assembles into homotetramers (H4) and heterotetramers with LDHA (M), forming the
    catalytically active oxidoreductase complex.
  molecular_function:
    id: GO:0004459
    label: L-lactate dehydrogenase (NAD+) activity
  directly_involved_in:
  - id: GO:0006089
    label: lactate metabolic process
  in_complex:
    id: GO:1990204
    label: oxidoreductase complex
  locations:
  - id: GO:0005829
    label: cytosol
  supported_by:
  - reference_id: PMID:27618187
    supporting_text: Functional LDH is a homo or hetero tetramer composed of LDHA and LDHB
      subunits
  - reference_id: file:human/LDHB/LDHB-uniprot.txt
    supporting_text: Homotetramer (PubMed:11276087)
suggested_questions:
- question: Is the mitochondrial (inner-membrane) pool of LDHB a genuine, functionally
    significant localization in vivo, or is it largely restricted to specific cell types
    (e.g. fermenting cancer cells) and experimental conditions?
- question: To what extent does the MP31 (PTEN-uORF) circuit-breaker regulation of LDHB
    operate in normal physiology versus tumour metabolism?
suggested_experiments:
- description: Compartment-resolved activity assays (cytosol vs mitochondrial
    sub-fractions) across oxidative and glycolytic human tissues to quantify the
    physiological contribution of any mitochondrial LDHB pool.
- description: Structure-guided kinetic comparison of purified LDHB homotetramer versus
    defined M/H heterotetramers to quantify the directional (lactate to pyruvate)
    preference under physiological substrate/redox conditions.
references:
- id: GO_REF:0000002
  title: Gene Ontology annotation through association of InterPro records with GO terms
  findings: []
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
  findings: []
- id: GO_REF:0000044
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location
    vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
  findings: []
- id: GO_REF:0000052
  title: Gene Ontology annotation based on curation of immunofluorescence data
  findings: []
- id: GO_REF:0000107
  title: Automatic transfer of experimentally verified manual GO annotation data to
    orthologs using Ensembl Compara
  findings: []
- id: GO_REF:0000120
  title: Combined Automated Annotation using Multiple IEA Methods
  findings: []
- id: file:human/LDHB/LDHB-uniprot.txt
  title: UniProtKB entry P07195 (LDHB_HUMAN)
  findings: []
- id: PMID:11276087
  title: Structural basis for altered activity of M- and H-isozyme forms of human
    lactate dehydrogenase.
  findings:
  - statement: Human LDH-H and LDH-M interconvert pyruvate and lactate with concomitant
      NADH/NAD+ interconversion; the H form is found mainly in cardiac muscle and is
      structurally indistinguishable at the active site from the M form.
    reference_section_type: ABSTRACT
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: Abstract-only cache; establishes the core L-lactate dehydrogenase (NAD+)
      activity and the H (heart) subunit identity of LDHB.
- id: PMID:15117937
  title: Identification and activity of a series of azole-based compounds with lactate
    dehydrogenase-directed anti-malarial activity.
  findings:
  - statement: LDH is central to NAD+ regeneration during glycolysis coupled with
      homolactic fermentation.
    reference_section_type: ABSTRACT
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: Primarily about Plasmodium LDH inhibitors; used by ComplexPortal as the
      reference for the human LDH complex annotations. Supports LDH's central role in NAD+
      regeneration.
- id: PMID:16130169
  title: Proteomics of human umbilical vein endothelial cells applied to etoposide-induced
    apoptosis.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: HUVEC proteomics; source of a TAS cytoplasm localization. Contextual only.
- id: PMID:19056867
  title: Large-scale proteomics and phosphoproteomics of urinary exosomes.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Exosome proteomics; abundant-cytosolic-protein detection, not a functional
      localization.
- id: PMID:19946888
  title: Defining the membrane proteome of NK cells.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Membrane-fraction proteomics; generic membrane detection for a soluble
      enzyme.
- id: PMID:20458337
  title: MHC class II-associated proteins in B-cell exosomes and potential functional
    implications for exosome biogenesis.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Exosome proteomics; abundant-cytosolic-protein detection.
- id: PMID:21044950
  title: Genome-wide YFP fluorescence complementation screen identifies new regulators
    for telomere signaling in human cells.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: High-throughput interaction screen (TERF1); source of a generic protein
      binding datum.
- id: PMID:21988832
  title: Toward an understanding of the protein interaction network of the human liver.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Interactome study (LDHA); generic protein binding datum consistent with
      LDH tetramer assembly.
- id: PMID:23533145
  title: In-depth proteomic analyses of exosomes isolated from expressed prostatic
    secretions in urine.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Exosome proteomics; abundant-cytosolic-protein detection.
- id: PMID:24725412
  title: Ribosomal protein s15 phosphorylation mediates LRRK2 neurodegeneration in
    Parkinson's disease.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: LRRK2 interactome context; generic protein binding datum.
- id: PMID:25204797
  title: Flotillin-1 facilitates toll-like receptor 3 signaling in human endothelial cells.
  findings:
  - statement: Flotillins are membrane raft markers; LDHB was recovered in the raft/caveolae
      proteomic analysis of this study.
    reference_section_type: ABSTRACT
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Membrane-raft proteomics; the raft localization of a soluble glycolytic
      enzyme is most likely a co-isolation artifact.
- id: PMID:25416956
  title: A proteome-scale map of the human interactome network.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Interactome map; source of protein binding (LDHA) and identical protein
      binding (self) data.
- id: PMID:25502805
  title: A massively parallel pipeline to clone DNA variants and examine molecular
    phenotypes of human disease mutations.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Variant-cloning interaction pipeline; source of an identical protein
      binding (self) datum consistent with homotetramerization.
- id: PMID:25910212
  title: Widespread macromolecular interaction perturbations in human genetic disorders.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Interaction-perturbation study (LDHA); generic protein binding datum.
- id: PMID:27618187
  title: Lactate metabolism is associated with mammalian mitochondria.
  findings:
  - statement: Functional LDH is a tetramer of LDHA and LDHB subunits; LDHB is the
      predominant heart (H) subunit and supports conversion of lactate to pyruvate in
      lactate-consuming oxidative cells. LDHB was localized by TEM immunogold to the inner
      mitochondrial membrane and mitochondrial lysates showed LDHB activity.
    reference_section_type: RESULTS
  reference_review:
    relevance: HIGH
    correctness: VERIFIED
    review_notes: Full text available; direct experimental support for LDHB catalytic
      activity, tetramer composition, H-subunit identity, and a non-canonical
      mitochondrial localization.
- id: PMID:28514442
  title: Architecture of the human interactome defines protein communities and disease
    networks.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Interactome community study (LDHA/LDHC); generic protein binding data.
- id: PMID:31515488
  title: Extensive disruption of protein interactions by genetic variants across the
    allele frequency spectrum in human populations.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Variant interaction study (LDHA); source of protein binding and identical
      protein binding data.
- id: PMID:32296183
  title: A reference map of the human binary protein interactome.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Binary interactome map; source of protein binding (SDCBP/LDHA) and
      identical protein binding (self) data.
- id: PMID:32814053
  title: Interactome Mapping Provides a Network of Neurodegenerative Disease Proteins
    and Uncovers Widespread Protein Aggregation in Affected Brains.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Neurodegeneration interactome (HTT); generic protein binding datum.
- id: PMID:33406399
  title: An Upstream Open Reading Frame in Phosphatase and Tensin Homolog Encodes
    a Circuit Breaker of Lactate Metabolism.
  findings:
  - statement: MP31, a micropeptide from the PTEN uORF, binds LDHB and limits
      lactate-pyruvate conversion in mitochondria by competing with mitochondrial LDH for
      NAD+.
    reference_section_type: ABSTRACT
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: Abstract-only cache; UniProt curates this as the LDHB-MP31 (C0HLV8)
      interaction abolished by LDHB D53A/R100A. Functionally meaningful regulatory
      interaction, though annotated only as generic protein binding.
- id: PMID:33961781
  title: Dual proteome-scale networks reveal cell-specific remodeling of the human
    interactome.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Cell-specific interactome remodeling (LDHA/LDHC); generic protein binding
      data.
- id: PMID:35271311
  title: 'OpenCell: Endogenous tagging for the cartography of human cellular organization.'
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: OpenCell interactome (LDHA/LDHC); generic protein binding data.
- id: PMID:40205054
  title: Multimodal cell maps as a foundation for structural and functional genomics.
  findings: []
  reference_review:
    relevance: LOW
    correctness: VERIFIED
    review_notes: Multimodal cell-map interactome (LDHC); generic protein binding datum.
- id: PMID:8314553
  title: Analysis of a genetic mutation in an electrophoretic variant of the human
    lactate dehydrogenase-B(H) subunit.
  findings:
  - statement: An electrophoretic LDH-B(H) subunit variant (K7E) with decreased heat
      stability and slightly lower LDH activity was characterized, confirming LDHB as the
      H subunit of human LDH.
    reference_section_type: ABSTRACT
  reference_review:
    relevance: MEDIUM
    correctness: VERIFIED
    review_notes: Abstract-only cache; source of the TAS L-lactate dehydrogenase (NAD+)
      activity annotation and confirms LDHB = H subunit.
- id: Reactome:R-HSA-70510
  title: LDH tetramer oxidises LACT to PYR
  findings: []
- id: Reactome:R-HSA-71849
  title: LDH tetramer reduces PYR to LACT
  findings: []