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. 2021 Feb 26;17(11):3671–3689. doi: 10.1080/15548627.2021.1886839

Table 1.

Proteins on human macrophage foam cell lipid droplets (LDs)

Protein Function Protein Function
    Cholesterol Efflux
CORO1A Phagolysosome assembly and maturation DSC1 Regulates apoAI-medidated cholesterol efflux
LAMTOR5 Lysosomal Ragulator complex/metabolic signaling PSAP Regulates cathepsin B/D activity, lipid trafficking and autophagy
PARK7 Positively regulates autophagy Lipid transport
VDAC2 Macroautophagy suppressor FABP4 Fatty acid transport protein
YWHAZ 14-3-3 protein zeta/delta FABP5 Fatty acid transport protein
YWHAG 14-3-3 protein gamma PPIA Regulates lipid droplet size
YWHAE 14-3-3 protein epsilon GOT2 Facilitates cellular uptake of long-chain free fatty acids
YWHAQ 14-3-3 protein theta Lysosomal function
DAP Negatively regulates autophagy CTSB,D,L,S Lysosomal cysteine proteases with roles in autophagy vescicle trafficking
HMGB1 Novel BECN1 interacting partner, promotes autophagy SLC25A5 Lysosomal solute carrier transporters implicated in autophagy/phagocytosis
    SLC25A6 Lysosomal solute carrier transporters implicated in autophagy/phagocytosis
TBCs Autophagy and protein aggregation MYL6 Myosin proteins linking autophagy receptors to autophagosomes
EEF1A1 Regulation of chaperone-mediated autophagy MYH6,7,9 Myosins
GRPEL1 Co-chaperone MYL12A,B Light chain myosin
HSP90AA1 Chaperone-mediated autophagy NPC2 Autophagy-lysosomal activity, mitophagy and lipid efflux from lysosomes
HSP90AB1 Molecular chaperone SCARB2 Lysosomal membrane protein, autophagy-lysosome pathway
HSP90B1 HSP90 paralogue, regulates innate and adaptive immunity Neutral lipases
HSPA8 Chaperone-mediated autophagy, autophagy and exocytosis CES1 Cholesterol ester hydrolase
HSPA9 Mitochondrial chaperone PNPLA2 Patatin-like phospholipase domain-containing protein 2
HSPB1 Regulation of autophagy LDAH Lipid droplet-associated hydrolase
HSPD1 Chaperonin FAF2 Inhibits PLPL2
HSPE1 Chaperonin Rab proteins  
HSPH1 Co-regulator of chaperone function RAB1A Regulation of autophagosome assembly
HYOU1 Hypoxia-regulated chaperone RAB2A Promotes autopahgy
P4HB ER chaperone RAB5C Member of the Ras-related protein family
PDIA3 ER chaperone RAB6A Regulates autophagy, binds to Atg8 family members
PDIA4 ER chaperone RAB6B Regulates autophagy, binds to Atg8 family members
TCP1 Chaperonin RAB7A Regulates lipophagy by mediating recruitement of lysosome to the LD
HSPA5 N-end rule proteolytic pathway, autophagy degradation RAB10 Required for LC3 recruitment ot autophagosomes
    RAB11A Regulates endosomal cholesterol movement to lipid droplets
H2A Core histone RAB11B Regulates endosomal cholesterol movement to lipid droplets
H2B Core histone RAB14 Phagolysosome assembly
H3 Core histone RAB18 Promotes LD growth and autophasosome formation
H4 Core histone RAB21 Promotes lipid droplet growth and autophasosome formation
    Stress granule regulation
PLIN2 Lipid droplet coat protein, intermediate-sized lipid droplets FUS Stress granule assembly factor
PLIN3 Lipid droplet coat protein, small lipid droplets PABPN1,C4 mRNA translation regulators
VIM Cytoskeletal protein Cellular Traficking
    COPB2 Golgi coatamer complex, delivery of PLIN2 to lipid droplets
ACADM Fatty acid oxidation CLTC Vesicle coat protein, intracellular trafficking and endocytosis
ACADVL Fatty acid oxidation ANXA2 Calcium dependent membrane protein involved in exocytosis
ACSL3 Fatty acid oxidation CHMP4B Part of the ESCRT machinery
ACSL4 Fatty acid oxidation Ubiquitination machinery
ALDH9A1 Aldehyde dehydrogenase AUP1 Ubiquitin binding, UBE2G2
ASAH1 Lysosomal ceramidase RPS27A Monoubiquitin
CS Citrate synthase UBB, UBC Polyubiquitin
CYB5R3 Desaturation and elongation of fatty acids, cholesterol synthesis UBA52 Monoubiquitin
DLD Lipoamide dehydrogenase VCP Aggresome assembly, autophagosome maturation
DLST Mitochondrial succinyltransferase Others  
GANAB Catalytic subunit of glucosidase II ALDOA Fructose-biphosphate aldolase
HSD17B11 Short-chain alcohol dehydrogenase ARHGDIA Rho GDP-dissociation inhibitor
LSS Lanosterol synthase ARHGDIB Rho GDP-dissociation inhibitor
MAN2B1 Alpha-mannosidase, lysososmal oligosaccharide degradation CDC42 GTPase
MDH2 Malate dehydrogenase GAPDH Converts D-glyceraldehyde 3-phosphate to 3-phospho-D-glyceroyl phosphate
METTL7A Methyltransferase GNB1 GTPase
PLA2G4C Phospholipase involved in lipid droplet biogenesis LDHA Converts pyruvate into lactate
RHOA,C Regulation of lipases RAC2 GTPase
SDHA Mitochondrial electron transport chain, Complex II RAP1A GTPase
SUCLG1 Hydrolyses succinyl-CoA TPD52L2 Promotes neutral lipid storage, contains a 14-3-3 binding motif
UQCRC1 Component of the ubiquinol-cytochrome c reductase complex VAPA Localized to membrane contact sites

LDs were purified from agLDL-loaded THP-1 macrophages by density gradient centrifugation and subjected to mass spectrometry. Names and cell function of identified proteins are shown.