Mutational scanning pinpoints distinct binding sites of key ATGL regulators in lipolysis.


Journal

Nature communications
ISSN: 2041-1723
Titre abrégé: Nat Commun
Pays: England
ID NLM: 101528555

Informations de publication

Date de publication:
21 Mar 2024
Historique:
received: 26 05 2023
accepted: 14 03 2024
medline: 22 3 2024
pubmed: 22 3 2024
entrez: 22 3 2024
Statut: epublish

Résumé

ATGL is a key enzyme in intracellular lipolysis and plays an important role in metabolic and cardiovascular diseases. ATGL is tightly regulated by a known set of protein-protein interaction partners with activating or inhibiting functions in the control of lipolysis. Here, we use deep mutational protein interaction perturbation scanning and generate comprehensive profiles of single amino acid variants that affect the interactions of ATGL with its regulatory partners: CGI-58, G0S2, PLIN1, PLIN5 and CIDEC. Twenty-three ATGL amino acid variants yield a specific interaction perturbation pattern when validated in co-immunoprecipitation experiments in mammalian cells. We identify and characterize eleven highly selective ATGL switch mutations which affect the interaction of one of the five partners without affecting the others. Switch mutations thus provide distinct interaction determinants for ATGL's key regulatory proteins at an amino acid resolution. When we test triglyceride hydrolase activity in vitro and lipolysis in cells, the activity patterns of the ATGL switch variants trace to their protein interaction profile. In the context of structural data, the integration of variant binding and activity profiles provides insights into the regulation of lipolysis and the impact of mutations in human disease.

Identifiants

pubmed: 38514628
doi: 10.1038/s41467-024-46937-x
pii: 10.1038/s41467-024-46937-x
doi:

Types de publication

Journal Article

Langues

eng

Sous-ensembles de citation

IM

Pagination

2516

Subventions

Organisme : Austrian Science Fund (Fonds zur Förderung der Wissenschaftlichen Forschung)
ID : 10.55776/DOC50
Organisme : Austrian Science Fund (Fonds zur Förderung der Wissenschaftlichen Forschung)
ID : P30162
Organisme : Karl-Franzens-Universität Graz (University of Graz)
ID : BioHealth

Informations de copyright

© 2024. The Author(s).

Références

Olzmann, J. A. & Carvalho, P. Dynamics and functions of lipid droplets. Nat. Rev. Mol. Cell Biol. 20, 137–155 (2019).
pubmed: 30523332 pmcid: 6746329 doi: 10.1038/s41580-018-0085-z
Grabner, G. F., Xie, H., Schweiger, M. & Zechner, R. Lipolysis: cellular mechanisms for lipid mobilization from fat stores. Nat. Metab. 3, 1445–1465 (2021).
pubmed: 34799702 doi: 10.1038/s42255-021-00493-6
Zimmermann, R. et al. Fat mobilization in adipose tissue is promoted by adipose triglyceride lipase. Science 306, 1383–1386 (2004).
pubmed: 15550674 doi: 10.1126/science.1100747
Haemmerle, G. et al. Defective lipolysis and altered energy metabolism in mice lacking adipose triglyceride lipase. Science 312, 734–737 (2006).
pubmed: 16675698 doi: 10.1126/science.1123965
Haemmerle, G. et al. ATGL-mediated fat catabolism regulates cardiac mitochondrial function via PPAR-α and PGC-1. Nat. Med. 17, 1076–1085 (2011).
pubmed: 21857651 pmcid: 3244833 doi: 10.1038/nm.2439
Schoiswohl, G. et al. Adipose triglyceride lipase plays a key role in the supply of the working muscle with fatty acids. J. Lipid Res. 51, 490–499 (2010).
pubmed: 19965578 pmcid: 2817579 doi: 10.1194/jlr.M001073
Schreiber, R. et al. Hypophagia and metabolic adaptations in mice with defective ATGL-mediated lipolysis cause resistance to HFD-induced obesity. Proc. Natl Acad. Sci. USA 112, 13850–13855 (2015).
pubmed: 26508640 pmcid: 4653216 doi: 10.1073/pnas.1516004112
Schweiger, M. et al. Pharmacological inhibition of adipose triglyceride lipase corrects high-fat diet-induced insulin resistance and hepatosteatosis in mice. Nat. Commun. 8, 14859 (2017).
pubmed: 28327588 pmcid: 5364409 doi: 10.1038/ncomms14859
Schoiswohl, G. et al. Impact of reduced ATGL-mediated adipocyte lipolysis on obesity-associated insulin resistance and inflammation in male mice. Endocrinology 156, 3610–3624 (2015).
pubmed: 26196542 pmcid: 4588821 doi: 10.1210/en.2015-1322
Grabner, G. F. et al. Small-molecule inhibitors targeting lipolysis in human adipocytes. J. Am. Chem. Soc. 144, 6237–6250 (2022).
pubmed: 35362954 pmcid: 9011347 doi: 10.1021/jacs.1c10836
Cerk, I. K., Wechselberger, L. & Oberer, M. Adipose triglyceride lipase regulation: an overview. Curr. protein Pept. Sci. 19, 221–233 (2018).
pubmed: 28925902 pmcid: 7613786
Granneman, J. G., Moore, H.-P. H., Krishnamoorthy, R. & Rathod, M. Perilipin controls lipolysis by regulating the interactions of AB-hydrolase containing 5 (Abhd5) and adipose triglyceride lipase (Atgl). J. Biol. Chem. 284, 34538–34544 (2009).
pubmed: 19850935 pmcid: 2787315 doi: 10.1074/jbc.M109.068478
Sahu-Osen, A. et al. CGI-58/ABHD5 is phosphorylated on Ser239 by protein kinase A: control of subcellular localization. J. Lipid Res. 56, 109–121 (2015).
pubmed: 25421061 pmcid: 4274058 doi: 10.1194/jlr.M055004
Recazens, E., Mouisel, E. & Langin, D. Hormone-sensitive lipase: sixty years later. Prog. Lipid Res. 82, 101084 (2021).
pubmed: 33387571 doi: 10.1016/j.plipres.2020.101084
Rondini, E. A. et al. Novel pharmacological probes reveal ABHD5 as a locus of lipolysis control in white and brown adipocytes. J. Pharmacol. Exp. Ther. 363, 367–376 (2017).
pubmed: 28928121 pmcid: 5698943 doi: 10.1124/jpet.117.243253
Lass, A. et al. Adipose triglyceride lipase-mediated lipolysis of cellular fat stores is activated by CGI-58 and defective in Chanarin-Dorfman Syndrome. Cell Metab. 3, 309–319 (2006).
pubmed: 16679289 doi: 10.1016/j.cmet.2006.03.005
Schweiger, M. et al. The C-terminal region of human adipose triglyceride lipase affects enzyme activity and lipid droplet binding. J. Biol. Chem. 283, 17211–17220 (2008).
pubmed: 18445597 doi: 10.1074/jbc.M710566200
Cornaciu, I. et al. The minimal domain of adipose triglyceride lipase (ATGL) ranges until leucine 254 and can be activated and inhibited by CGI-58 and G0S2, respectively. PLoS ONE 6, e26349 (2011).
pubmed: 22039468 pmcid: 3198459 doi: 10.1371/journal.pone.0026349
Tseng, Y. Y. et al. Structural and functional insights into ABHD5, a ligand-regulated lipase co-activator. Sci. Rep. 12, 2565 (2022).
pubmed: 35173175 pmcid: 8850477 doi: 10.1038/s41598-021-04179-7
Yang, X. et al. The G(0)/G(1) switch gene 2 regulates adipose lipolysis through association with adipose triglyceride lipase. Cell Metab. 11, 194–205 (2010).
pubmed: 20197052 pmcid: 3658843 doi: 10.1016/j.cmet.2010.02.003
Riegler-Berket, L. et al. Residues of the minimal sequence of G0S2 collectively contribute to ATGL inhibition while C-and N-terminal extensions promote binding to ATGL. Biochim. Biophys. Acta Mol. Cell Biol. Lipids 1867, 159105 (2022).
pubmed: 35026402 doi: 10.1016/j.bbalip.2021.159105
Puri, V. et al. Fat-specific protein 27, a novel lipid droplet protein that enhances triglyceride storage. J. Biol. Chem. 282, 34213–34218 (2007).
pubmed: 17884815 doi: 10.1074/jbc.M707404200
Grahn, T. H. M. et al. Fat-specific protein 27 (FSP27) interacts with adipose triglyceride lipase (ATGL) to regulate lipolysis and insulin sensitivity in human adipocytes. J. Biol. Chem. 289, 12029–12039 (2014).
pubmed: 24627478 pmcid: 4002109 doi: 10.1074/jbc.M113.539890
Yang, X., Heckmann, B. L., Zhang, X., Smas, C. M. & Liu, J. Distinct mechanisms regulate ATGL-mediated adipocyte lipolysis by lipid droplet coat proteins. Mol. Endocrinol. 27, 116–126 (2013).
pubmed: 23204327 doi: 10.1210/me.2012-1178
Najt, C. P., Devarajan, M. & Mashek, D. G. Perilipins at a glance. J. Cell Sci. 135, jcs259501 (2022).
Subramanian, V. et al. Perilipin A mediates the reversible binding of CGI-58 to lipid droplets in 3T3-L1 adipocytes. J. Biol. Chem. 279, 42062–42071 (2004).
pubmed: 15292255 doi: 10.1074/jbc.M407462200
Yamaguchi, T., Omatsu, N., Matsushita, S. & Osumi, T. CGI-58 interacts with perilipin and is localized to lipid droplets. Possible involvement of CGI-58 mislocalization in Chanarin-Dorfman syndrome. J. Biol. Chem. 279, 30490–30497 (2004).
pubmed: 15136565 doi: 10.1074/jbc.M403920200
Tansey, J. T. et al. Perilipin ablation results in a lean mouse with aberrant adipocyte lipolysis, enhanced leptin production, and resistance to diet-induced obesity. Proc. Natl Acad. Sci. USA 98, 6494–6499 (2001).
pubmed: 11371650 pmcid: 33496 doi: 10.1073/pnas.101042998
Miyoshi, H. et al. Perilipin overexpression in mice protects against diet-induced obesity. J. Lipid Res. 51, 975–982 (2010).
pubmed: 19797618 pmcid: 2853465 doi: 10.1194/jlr.M002352
Sawada, T. et al. Perilipin overexpression in white adipose tissue induces a brown fat-like phenotype. PLoS ONE 5, e14006 (2010).
pubmed: 21103377 pmcid: 2982838 doi: 10.1371/journal.pone.0014006
Wang, H. et al. Perilipin 5, a lipid droplet-associated protein, provides physical and metabolic linkage to mitochondria. J. Lipid Res. 52, 2159–2168 (2011).
pubmed: 21885430 pmcid: 3220284 doi: 10.1194/jlr.M017939
Pollak, N. M. et al. The interplay of protein kinase A and perilipin 5 regulates cardiac lipolysis. J. Biol. Chem. 290, 1295–1306 (2015).
pubmed: 25418045 doi: 10.1074/jbc.M114.604744
Granneman, J. G., Moore, H.-P. H., Mottillo, E. P., Zhu, Z. & Zhou, L. Interactions of perilipin-5 (Plin5) with adipose triglyceride lipase. J. Biol. Chem. 286, 5126–5135 (2011).
pubmed: 21148142 doi: 10.1074/jbc.M110.180711
Kien, B. et al. Lipid droplet-mitochondria coupling via perilipin 5 augments respiratory capacity but is dispensable for FA oxidation. J. Lipid Res. 63, 100172 (2022).
pubmed: 35065923 pmcid: 8953689 doi: 10.1016/j.jlr.2022.100172
Pollak, N. M. et al. Cardiac-specific overexpression of perilipin 5 provokes severe cardiac steatosis via the formation of a lipolytic barrier. J. Lipid Res. 54, 1092–1102 (2013).
pubmed: 23345410 pmcid: 3605985 doi: 10.1194/jlr.M034710
Kuramoto, K. et al. Perilipin 5, a lipid droplet-binding protein, protects heart from oxidative burden by sequestering fatty acid from excessive oxidation. J. Biol. Chem. 287, 23852–23863 (2012).
pubmed: 22532565 pmcid: 3390660 doi: 10.1074/jbc.M111.328708
Shendure, J. & Fields, S. Massively parallel. Genet. Genet. 203, 617–619 (2016).
Kunowska, N. & Stelzl, U. Decoding the cellular effects of genetic variation through interaction proteomics. Curr. Opin. Chem. Biol. 66, 102100 (2022).
pubmed: 34801969 doi: 10.1016/j.cbpa.2021.102100
Moesslacher, C. S., Kohlmayr, J. M. & Stelzl, U. Exploring absent protein function in yeast: assaying post translational modification and human genetic variation. Micro. Cell 8, 164–183 (2021).
doi: 10.15698/mic2021.08.756
Woodsmith, J. et al. Protein interaction perturbation profiling at amino-acid resolution. Nat. Methods 14, 1213–1221 (2017).
pubmed: 29039417 doi: 10.1038/nmeth.4464
Starita, L. M. et al. Massively parallel functional analysis of BRCA1 RING domain variants. Genetics 200, 413–422 (2015).
pubmed: 25823446 pmcid: 4492368 doi: 10.1534/genetics.115.175802
Diss, G. & Lehner, B. The genetic landscape of a physical interaction. eLife 7, e32472 (2018).
pubmed: 29638215 pmcid: 5896888 doi: 10.7554/eLife.32472
Hidalgo, F. et al. A saturation-mutagenesis analysis of the interplay between stability and activation in Ras. eLife 11, e76595 (2022).
pubmed: 35272765 pmcid: 8916776 doi: 10.7554/eLife.76595
Weng, C. Faure, A. J., Escobedo, A. & Lehner, B. The energetic and allosteric landscape for KRAS inhibition. Nature 626, 643–652 (2024).
Faure, A. J. et al. Mapping the energetic and allosteric landscapes of protein binding domains. Nature 604, 175–183 (2022).
pubmed: 35388192 doi: 10.1038/s41586-022-04586-4
Moesslacher, C. S. et al. Missense variant interaction scanning reveals a critical role of the FERM domain for tumor suppressor protein NF2 conformation and function. Life Sci. Alliance 6, e202302043 (2023).
pubmed: 37280085 pmcid: 10244618 doi: 10.26508/lsa.202302043
Worseck, J. M., Grossmann, A., Weimann, M., Hegele, A. & Stelzl, U. A stringent yeast two-hybrid matrix screening approach for protein-protein interaction discovery. Methods Mol. Biol. 812, 63–87 (2012).
pubmed: 22218854 doi: 10.1007/978-1-61779-455-1_4
Weimann, M. et al. A Y2H-seq approach defines the human protein methyltransferase interactome. Nat. Methods 10, 339–342 (2013).
pubmed: 23455924 doi: 10.1038/nmeth.2397
Wrenbeck, E. E. et al. Plasmid-based one-pot saturation mutagenesis. Nat. Methods 13, 928–930 (2016).
pubmed: 27723752 pmcid: 5666567 doi: 10.1038/nmeth.4029
Hegele, A. et al. Dynamic protein-protein interaction wiring of the human spliceosome. Mol. Cell 45, 567–580 (2012).
pubmed: 22365833 doi: 10.1016/j.molcel.2011.12.034
Taipale, M. et al. Chaperones as thermodynamic sensors of drug-target interactions reveal kinase inhibitor specificities in living cells. Nat. Biotechnol. 31, 630–637 (2013).
pubmed: 23811600 pmcid: 3774174 doi: 10.1038/nbt.2620
Schweiger, M. et al. Measurement of lipolysis. Methods Enzymol. 538, 171–193 (2014).
pubmed: 24529439 pmcid: 4018506 doi: 10.1016/B978-0-12-800280-3.00010-4
Itabe, H., Yamaguchi, T., Nimura, S. & Sasabe, N. Perilipins: a diversity of intracellular lipid droplet proteins. Lipids Health Dis. 16, 83 (2017).
pubmed: 28454542 pmcid: 5410086 doi: 10.1186/s12944-017-0473-y
Smirnova, E. et al. ATGL has a key role in lipid droplet/adiposome degradation in mammalian cells. EMBO Rep. 7, 106–113 (2006).
pubmed: 16239926 doi: 10.1038/sj.embor.7400559
Ellong, E. N. et al. Interaction between the triglyceride lipase ATGL and the Arf1 activator GBF1. PLoS ONE 6, e21889 (2011).
pubmed: 21789191 pmcid: 3138737 doi: 10.1371/journal.pone.0021889
Tavian, D. et al. Contribution of novel ATGL missense mutations to the clinical phenotype of NLSD-M: a strikingly low amount of lipase activity may preserve cardiac function. Hum. Mol. Genet. 21, 5318–5328 (2012).
pubmed: 22990388 pmcid: 3510752 doi: 10.1093/hmg/dds388
Fischer, J. et al. The gene encoding adipose triglyceride lipase (PNPLA2) is mutated in neutral lipid storage disease with myopathy. Nat. Genet. 39, 28–30 (2007).
pubmed: 17187067 doi: 10.1038/ng1951
Landrum, M. J. et al. ClinVar: improving access to variant interpretations and supporting evidence. Nucleic Acids Res. 46, D1062–D1067 (2018).
pubmed: 29165669 doi: 10.1093/nar/gkx1153
Karczewski, K. J. et al. The mutational constraint spectrum quantified from variation in 141,456 humans. Nature 581, 434–443 (2020).
pubmed: 32461654 pmcid: 7334197 doi: 10.1038/s41586-020-2308-7
Kulminskaya, N. & Oberer, M. Protein-protein interactions regulate the activity of Adipose Triglyceride Lipase in intracellular lipolysis. Biochimie 169, 62–68 (2020).
pubmed: 31404588 doi: 10.1016/j.biochi.2019.08.004
Jumper, J. et al. Highly accurate protein structure prediction with AlphaFold. Nature 596, 583–589 (2021).
pubmed: 34265844 pmcid: 8371605 doi: 10.1038/s41586-021-03819-2
Saha, P. K., Kojima, H., Martinez-Botas, J., Sunehag, A. L. & Chan, L. Metabolic adaptations in the absence of perilipin: increased beta-oxidation and decreased hepatic glucose production associated with peripheral insulin resistance but normal glucose tolerance in perilipin-null mice. J. Biol. Chem. 279, 35150–35158 (2004).
pubmed: 15197189 doi: 10.1074/jbc.M405499200
Kulminskaya, N. et al. Unmasking crucial residues in adipose triglyceride lipase (ATGL) for co-activation with comparative gene identification-58 (CGI-58). J. Lipid Res. 69, 100491 (2024).
doi: 10.1016/j.jlr.2023.100491
Xie, X. et al. Identification of a novel phosphorylation site in adipose triglyceride lipase as a regulator of lipid droplet localization. Am. J. Physiol. Endocrinol. Metab. 306, E1449–E1459 (2014).
pubmed: 24801391 pmcid: 4059987 doi: 10.1152/ajpendo.00663.2013
Ahmadian, M. et al. Desnutrin/ATGL is regulated by AMPK and is required for a brown adipose phenotype. Cell Metab. 13, 739–748 (2011).
pubmed: 21641555 pmcid: 3148136 doi: 10.1016/j.cmet.2011.05.002
Schweiger, M., Lass, A., Zimmermann, R., Eichmann, T. O. & Zechner, R. Neutral lipid storage disease: genetic disorders caused by mutations in adipose triglyceride lipase/PNPLA2 or CGI-58/ABHD5. Am. J. Physiol. Endocrinol. Metab. 297, E289–E296 (2009).
pubmed: 19401457 doi: 10.1152/ajpendo.00099.2009
Boeszoermenyi, A. et al. Structure of a CGI-58 motif provides the molecular basis of lipid droplet anchoring. J. Biol. Chem. 290, 26361–26372 (2015).
pubmed: 26350461 pmcid: 4646293 doi: 10.1074/jbc.M115.682203
Zhang, X., Heckmann, B. L., Campbell, L. E. & Liu, J. G0S2: a small giant controller of lipolysis and adipose-liver fatty acid flux. Biochimica Biophysica Acta Mol. Cell Biol. Lipids 1862, 1146–1154 (2017).
doi: 10.1016/j.bbalip.2017.06.007
Schweiger, M. et al. G0/G1 switch gene-2 regulates human adipocyte lipolysis by affecting activity and localization of adipose triglyceride lipase. J. Lipid Res. 53, 2307–2317 (2012).
pubmed: 22891293 pmcid: 3466000 doi: 10.1194/jlr.M027409
Cerk, I. K. et al. A peptide derived from G0/G1 switch gene 2 acts as noncompetitive inhibitor of adipose triglyceride lipase. J. Biol. Chem. 289, 32559–32570 (2014).
pubmed: 25258314 pmcid: 4239610 doi: 10.1074/jbc.M114.602599
Hubbard, S. R., Mohammadi, M. & Schlessinger, J. Autoregulatory mechanisms in protein-tyrosine kinases. J. Biol. Chem. 273, 11987–11990 (1998).
pubmed: 9575136 doi: 10.1074/jbc.273.20.11987
Huse, M. & Kuriyan, J. The conformational plasticity of protein kinases. Cell 109, 275–282 (2002).
pubmed: 12015977 doi: 10.1016/S0092-8674(02)00741-9
Sahni, N. et al. Widespread macromolecular interaction perturbations in human genetic disorders. Cell 161, 647–660 (2015).
pubmed: 25910212 pmcid: 4441215 doi: 10.1016/j.cell.2015.04.013
Yadav, A., Vidal, M. & Luck, K. Precision medicine - networks to the rescue. Curr. Opin. Biotechnol. 63, 177–189 (2020).
pubmed: 32199228 pmcid: 7308189 doi: 10.1016/j.copbio.2020.02.005
Woodsmith, J. & Stelzl, U. Understanding disease variants through the lens of protein interactions. Cell Syst. 5, 544–546 (2017).
pubmed: 29284128 doi: 10.1016/j.cels.2017.12.009
Gietz, R. D. & Schiestl, R. H. High-efficiency yeast transformation using the LiAc/SS carrier DNA/PEG method. Nat. Protoc. 2, 31–34 (2007).
pubmed: 17401334 doi: 10.1038/nprot.2007.13
Liu, H. & Naismith, J. H. An efficient one-step site-directed deletion, insertion, single and multiple-site plasmid mutagenesis protocol. BMC Biotechnol. 8, 91 (2008).
pubmed: 19055817 pmcid: 2629768 doi: 10.1186/1472-6750-8-91
Yadav, L. et al. Systematic analysis of human protein phosphatase interactions and dynamics. Cell Syst. 4, 430–444.e5 (2017).
pubmed: 28330616 doi: 10.1016/j.cels.2017.02.011

Auteurs

Johanna M Kohlmayr (JM)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.

Gernot F Grabner (GF)

Institute of Molecular Biosciences, Biochemistry, University of Graz, Graz, Austria.
Gottfried Schatz Research Center, Molecular Biology and Biochemistry, Medical University of Graz, Graz, Austria.

Anna Nusser (A)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.

Anna Höll (A)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.

Verina Manojlović (V)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.

Bettina Halwachs (B)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.
Field of Excellence BioHealth - University of Graz, Graz, Austria.

Sarah Masser (S)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.
BioTechMed-Graz, Graz, Austria.

Evelyne Jany-Luig (E)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.

Hanna Engelke (H)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria.
Field of Excellence BioHealth - University of Graz, Graz, Austria.

Robert Zimmermann (R)

Institute of Molecular Biosciences, Biochemistry, University of Graz, Graz, Austria.
Field of Excellence BioHealth - University of Graz, Graz, Austria.
BioTechMed-Graz, Graz, Austria.

Ulrich Stelzl (U)

Institute of Pharmaceutical Sciences, Pharmaceutical Chemistry, University of Graz, Graz, Austria. ulrich.stelzl@uni-graz.at.
Field of Excellence BioHealth - University of Graz, Graz, Austria. ulrich.stelzl@uni-graz.at.
BioTechMed-Graz, Graz, Austria. ulrich.stelzl@uni-graz.at.

Classifications MeSH