{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T01:07:32Z","timestamp":1792804052824,"version":"4.5.0"},"reference-count":153,"publisher":"Springer Science and Business Media LLC","issue":"9","license":[{"start":{"date-parts":[[2020,6,25]],"date-time":"2020-06-25T00:00:00Z","timestamp":1593043200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"},{"start":{"date-parts":[[2020,6,25]],"date-time":"2020-06-25T00:00:00Z","timestamp":1593043200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Rev Endocrinol"],"published-print":{"date-parts":[[2020,9]]},"DOI":"10.1038\/s41574-020-0375-3","type":"journal-article","created":{"date-parts":[[2020,6,25]],"date-time":"2020-06-25T13:02:53Z","timestamp":1593090173000},"page":"506-518","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":137,"title":["Emerging routes to the generation of functional \u03b2-cells for diabetes mellitus cell therapy"],"prefix":"10.1038","volume":"16","author":[{"given":"Gopika G.","family":"Nair","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emmanuel S.","family":"Tzanakakis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3833-8862","authenticated-orcid":false,"given":"Matthias","family":"Hebrok","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2020,6,25]]},"reference":[{"key":"375_CR1","unstructured":"International Diabetes Federation. IDF Atlas 9th edn (IDF, 2019)."},{"key":"375_CR2","doi-asserted-by":"publisher","first-page":"917","DOI":"10.2337\/dci18-0007","volume":"41","author":"American Diabetes Association","year":"2018","unstructured":"American Diabetes Association. Economic costs of diabetes in the U.S. in 2017. Diabetes Care 41, 917\u2013928 (2018).","journal-title":"Diabetes Care"},{"key":"375_CR3","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1056\/NEJMra1215228","volume":"369","author":"PE Cryer","year":"2013","unstructured":"Cryer, P. E. Mechanisms of hypoglycemia-associated autonomic failure in diabetes. N. Engl. J. Med. 369, 362\u2013372 (2013).","journal-title":"N. Engl. J. Med."},{"key":"375_CR4","doi-asserted-by":"publisher","first-page":"977","DOI":"10.1056\/NEJM199309303291401","volume":"329","author":"DM Nathan","year":"1993","unstructured":"Diabetes Control and Complications Trial Research Group, Nathan, D. M. et al. The effect of intensive treatment of diabetes on the development and progression of long-term complications in insulin-dependent diabetes mellitus. N. Engl. J. Med. 329, 977\u2013986 (1993).","journal-title":"N. Engl. J. Med."},{"key":"375_CR5","doi-asserted-by":"publisher","first-page":"631","DOI":"10.2337\/db14-0930","volume":"64","author":"The Diabetes Control and Complications Trial (DCCT)\/Epidemiology of Diabetes Interventions and Complications (EDIC) Research Group","year":"2015","unstructured":"The Diabetes Control and Complications Trial (DCCT)\/Epidemiology of Diabetes Interventions and Complications (EDIC) Research Group. Effect of intensive diabetes therapy on the progression of diabetic retinopathy in patients with type 1 diabetes: 18 years of follow-up in the DCCT\/EDIC. Diabetes 64, 631\u2013642 (2015).","journal-title":"Diabetes"},{"key":"375_CR6","doi-asserted-by":"publisher","first-page":"643","DOI":"10.1046\/j.1365-2125.1999.00092.x","volume":"48","author":"P King","year":"1999","unstructured":"King, P., Peacock, I. & Donnelly, R. The UK prospective diabetes study (UKPDS): clinical and therapeutic implications for type 2 diabetes. Br. J. Clin. Pharmacol. 48, 643\u2013648 (1999).","journal-title":"Br. J. Clin. Pharmacol."},{"key":"375_CR7","doi-asserted-by":"publisher","first-page":"1321","DOI":"10.1016\/S0140-6736(18)31947-0","volume":"392","author":"M Tauschmann","year":"2018","unstructured":"Tauschmann, M. et al. Closed-loop insulin delivery in suboptimally controlled type 1 diabetes: a multicentre, 12-week randomised trial. Lancet 392, 1321\u20131329 (2018).","journal-title":"Lancet"},{"key":"375_CR8","doi-asserted-by":"publisher","first-page":"k1310","DOI":"10.1136\/bmj.k1310","volume":"361","author":"E Bekiari","year":"2018","unstructured":"Bekiari, E. et al. Artificial pancreas treatment for outpatients with type 1 diabetes: systematic review and meta-analysis. BMJ 361, k1310 (2018).","journal-title":"BMJ"},{"key":"375_CR9","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1089\/dia.2018.0384","volume":"21","author":"NC Foster","year":"2019","unstructured":"Foster, N. C. et al. State of type 1 diabetes management and outcomes from the T1D exchange in 2016\u20132018. Diabetes Technol. Ther. 21, 66\u201372 (2019).","journal-title":"Diabetes Technol. Ther."},{"key":"375_CR10","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1016\/S2213-8587(15)00489-1","volume":"4","author":"SJ Russell","year":"2016","unstructured":"Russell, S. J. et al. Day and night glycaemic control with a bionic pancreas versus conventional insulin pump therapy in preadolescent children with type 1 diabetes: a randomised crossover trial. Lancet Diabetes Endocrinol. 4, 233\u2013243 (2016).","journal-title":"Lancet Diabetes Endocrinol."},{"key":"375_CR11","doi-asserted-by":"publisher","first-page":"1707","DOI":"10.1056\/NEJMoa1907863","volume":"381","author":"SA Brown","year":"2019","unstructured":"Brown, S. A. et al. Six-month randomized, multicenter trial of closed-loop control in type 1 diabetes. N. Engl. J. Med. 381, 1707\u20131717 (2019). This report describes results from a multicentre trial evaluating benefits of closed-loop control over sensor-augumented insulin pumps. Closed-loop systems fare better in maintaining the time spent in target glycaemic range (mean ~71%) than sensor-augumented insulin pumps (mean ~59%).","journal-title":"N. Engl. J. Med."},{"key":"375_CR12","doi-asserted-by":"publisher","first-page":"1436","DOI":"10.2337\/dc12-0063","volume":"35","author":"FB Barton","year":"2012","unstructured":"Barton, F. B. et al. Improvement in outcomes of clinical islet transplantation: 1999\u20132010. Diabetes Care 35, 1436\u20131445 (2012).","journal-title":"Diabetes Care"},{"key":"375_CR13","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1038\/nbt.2141","volume":"30","author":"B Blum","year":"2012","unstructured":"Blum, B. et al. Functional beta-cell maturation is marked by an increased glucose threshold and by expression of urocortin 3. Nat. Biotech. 30, 261\u2013264 (2012).","journal-title":"Nat. Biotech."},{"key":"375_CR14","doi-asserted-by":"crossref","first-page":"769","DOI":"10.1038\/nm.3872","volume":"21","author":"T van der Meulen","year":"2015","unstructured":"van der Meulen, T. et al. Urocortin3 mediates somatostatin-dependent negative feedback control of insulin secretion. Nat. Med. 21, 769\u2013776 (2015).","journal-title":"Nat. Med."},{"key":"375_CR15","doi-asserted-by":"publisher","first-page":"1751","DOI":"10.2337\/diabetes.49.11.1751","volume":"49","author":"JC Henquin","year":"2000","unstructured":"Henquin, J. C. Triggering and amplifying pathways of regulation of insulin secretion by glucose. Diabetes 49, 1751\u20131760 (2000).","journal-title":"Diabetes"},{"key":"375_CR16","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1111\/jdi.12094","volume":"4","author":"M Komatsu","year":"2013","unstructured":"Komatsu, M. et al. Glucose-stimulated insulin secretion: a newer perspective. J. Diabetes Investig. 4, 511\u2013516 (2013).","journal-title":"J. Diabetes Investig."},{"key":"375_CR17","doi-asserted-by":"publisher","first-page":"993","DOI":"10.1016\/j.cmet.2014.04.003","volume":"19","author":"S Zhao","year":"2014","unstructured":"Zhao, S. et al. \u03b1\/\u03b2-Hydrolase domain-6-accessible monoacylglycerol controls glucose-stimulated insulin secretion. Cell Metab. 19, 993\u20131007 (2014).","journal-title":"Cell Metab."},{"key":"375_CR18","doi-asserted-by":"publisher","first-page":"3847","DOI":"10.1172\/JCI82498","volume":"125","author":"M Ferdaoussi","year":"2015","unstructured":"Ferdaoussi, M. et al. Isocitrate-to-SENP1 signaling amplifies insulin secretion and rescues dysfunctional \u03b2 cells. J. Clin. Investig. 125, 3847\u20133860 (2015).","journal-title":"J. Clin. Investig."},{"key":"375_CR19","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1016\/j.celrep.2015.08.072","volume":"13","author":"JR Gooding","year":"2015","unstructured":"Gooding, J. R. et al. Adenylosuccinate is an insulin secretagogue derived from glucose-induced purine metabolism. Cell Rep. 13, 157\u2013167 (2015).","journal-title":"Cell Rep."},{"key":"375_CR20","doi-asserted-by":"publisher","first-page":"89","DOI":"10.4161\/isl.2.2.11025","volume":"2","author":"TJ Pullen","year":"2010","unstructured":"Pullen, T. J. et al. Identification of genes selectively disallowed in the pancreatic islet. Islets 2, 89\u201395 (2010).","journal-title":"Islets"},{"key":"375_CR21","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1101\/gr.109173.110","volume":"21","author":"L Thorrez","year":"2011","unstructured":"Thorrez, L. et al. Tissue-specific disallowance of housekeeping genes: the other face of cell differentiation. Genome Res. 21, 95\u2013105 (2011).","journal-title":"Genome Res."},{"key":"375_CR22","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1146\/annurev-nutr-071715-050808","volume":"36","author":"K Lemaire","year":"2016","unstructured":"Lemaire, K., Thorrez, L. & Schuit, F. Disallowed and allowed gene expression: two faces of mature islet beta cells. Annu. Rev. Nutr. 36, 45\u201371 (2016).","journal-title":"Annu. Rev. Nutr."},{"key":"375_CR23","doi-asserted-by":"publisher","first-page":"1262","DOI":"10.1016\/j.celrep.2013.08.010","volume":"4","author":"BL Taylor","year":"2013","unstructured":"Taylor, B. L., Liu, F.-F. & Sander, M. Nkx6.1 is essential for maintaining the functional state of pancreatic beta cells. Cell Rep. 4, 1262\u20131275 (2013).","journal-title":"Cell Rep."},{"key":"375_CR24","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1016\/j.cmet.2010.03.006","volume":"11","author":"C Gu","year":"2010","unstructured":"Gu, C. et al. Pancreatic \u03b2 cells require NeuroD to achieve and maintain functional maturity. Cell Metab. 11, 298\u2013310 (2010).","journal-title":"Cell Metab."},{"key":"375_CR25","doi-asserted-by":"publisher","first-page":"696","DOI":"10.1210\/me.2011-1256","volume":"26","author":"Y Gosmain","year":"2012","unstructured":"Gosmain, Y. et al. Pax6 is crucial for \u03b2-cell function, insulin biosynthesis, and glucose-induced insulin secretion. Mol. Endocrinol. 26, 696\u2013709 (2012).","journal-title":"Mol. Endocrinol."},{"key":"375_CR26","doi-asserted-by":"publisher","first-page":"1569","DOI":"10.2337\/db12-0849","volume":"62","author":"C Aguayo-Mazzucato","year":"2013","unstructured":"Aguayo-Mazzucato, C. et al. Thyroid hormone promotes postnatal rat pancreatic \u03b2-cell development and glucose-responsive insulin secretion through MAFA. Diabetes 62, 1569\u20131580 (2013).","journal-title":"Diabetes"},{"key":"375_CR27","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.devcel.2018.03.013","volume":"45","author":"C Huang","year":"2018","unstructured":"Huang, C. et al. Synaptotagmin 4 regulates pancreatic \u03b2 cell maturation by modulating the Ca2+ sensitivity of insulin secretion vesicles. Dev. Cell 45, 347\u2013361 (2018).","journal-title":"Dev. Cell"},{"key":"375_CR28","doi-asserted-by":"publisher","first-page":"909","DOI":"10.1016\/j.cmet.2016.04.002","volume":"23","author":"HE Arda","year":"2016","unstructured":"Arda, H. E. et al. Age-dependent pancreatic gene regulation reveals mechanisms governing human \u03b2 cell function. Cell Metab. 23, 909\u2013920 (2016).","journal-title":"Cell Metab."},{"key":"375_CR29","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0165961","volume":"11","author":"J-C Henquin","year":"2016","unstructured":"Henquin, J.-C. & Nenquin, M. Dynamics and regulation of insulin secretion in pancreatic islets from normal young children. PLoS One 11, e0165961 (2016).","journal-title":"PLoS One"},{"key":"375_CR30","doi-asserted-by":"publisher","first-page":"274","DOI":"10.1136\/adc.68.3_Spec_No.274","volume":"68","author":"JM Hawdon","year":"1993","unstructured":"Hawdon, J. M. et al. The role of pancreatic insulin secretion in neonatal glucoregulation. I. Healthy term and preterm infants. Arch. Dis. Child. 68, 274\u2013279 (1993).","journal-title":"Arch. Dis. Child."},{"key":"375_CR31","doi-asserted-by":"publisher","first-page":"836","DOI":"10.1016\/S0022-3476(61)80314-4","volume":"59","author":"R Kaye","year":"1961","unstructured":"Kaye, R. et al. The response of blood glucose, ketones, and plasma nonesterified fatty acids to fasting and epinephrine injection in infants and children. J. Pediatr. 59, 836\u2013847 (1961).","journal-title":"J. Pediatr."},{"key":"375_CR32","doi-asserted-by":"publisher","first-page":"342","DOI":"10.2337\/db19-0281","volume":"69","author":"X Tong","year":"2020","unstructured":"Tong, X. et al. Lipid droplet accumulation in human pancreatic islets is dependent on both donor age and health. Diabetes 69, 342 (2020).","journal-title":"Diabetes"},{"key":"375_CR33","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1016\/j.cmet.2016.06.020","volume":"24","author":"NR Johnston","year":"2016","unstructured":"Johnston, N. R. et al. Beta cell hubs dictate pancreatic islet responses to glucose. Cell Metab. 24, 389\u2013401 (2016).","journal-title":"Cell Metab."},{"key":"375_CR34","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1038\/nature18624","volume":"535","author":"E Bader","year":"2016","unstructured":"Bader, E. et al. Identification of proliferative and mature \u03b2-cells in the islets of Langerhans. Nature 535, 430\u2013434 (2016).","journal-title":"Nature"},{"key":"375_CR35","doi-asserted-by":"publisher","first-page":"R43","DOI":"10.1530\/JME-18-0011","volume":"61","author":"D Nasteska","year":"2018","unstructured":"Nasteska, D. & Hodson, D. J. The role of beta cell heterogeneity in islet function and insulin release. J. Mol. Endocrinol. 61, R43\u2013R60 (2018).","journal-title":"J. Mol. Endocrinol."},{"key":"375_CR36","doi-asserted-by":"publisher","first-page":"1145","DOI":"10.1126\/science.282.5391.1145","volume":"282","author":"JA Thomson","year":"1998","unstructured":"Thomson, J. A. et al. Embryonic stem cell lines derived from human blastocysts. Science 282, 1145\u20131147 (1998).","journal-title":"Science"},{"key":"375_CR37","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1016\/j.cell.2006.07.024","volume":"126","author":"K Takahashi","year":"2006","unstructured":"Takahashi, K. & Yamanaka, S. Induction of pluripotent stem cells from mouse embryonic and adult fibroblast cultures by defined factors. Cell 126, 663\u2013676 (2006).","journal-title":"Cell"},{"key":"375_CR38","doi-asserted-by":"publisher","first-page":"1917","DOI":"10.1126\/science.1151526","volume":"318","author":"J Yu","year":"2007","unstructured":"Yu, J. et al. Induced pluripotent stem cell lines derived from human somatic cells. Science 318, 1917\u20131920 (2007).","journal-title":"Science"},{"key":"375_CR39","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1089\/clo.2006.8.96","volume":"8","author":"X Xu","year":"2006","unstructured":"Xu, X. et al. Endoderm and pancreatic islet lineage differentiation from human embryonic stem cells. Cloning Stem Cell 8, 96\u2013107 (2006).","journal-title":"Cloning Stem Cell"},{"key":"375_CR40","doi-asserted-by":"publisher","first-page":"1691","DOI":"10.2337\/diabetes.50.8.1691","volume":"50","author":"S Assady","year":"2001","unstructured":"Assady, S. et al. Insulin production by human embryonic stem cells. Diabetes 50, 1691\u20131697 (2001).","journal-title":"Diabetes"},{"key":"375_CR41","doi-asserted-by":"publisher","first-page":"1534","DOI":"10.1038\/nbt1163","volume":"23","author":"KA D\u2019Amour","year":"2005","unstructured":"D\u2019Amour, K. A. et al. Efficient differentiation of human embryonic stem cells to definitive endoderm. Nat. Biotechnol. 23, 1534\u20131541 (2005).","journal-title":"Nat. Biotechnol."},{"key":"375_CR42","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1038\/nbt1393","volume":"26","author":"E Kroon","year":"2008","unstructured":"Kroon, E. et al. Pancreatic endoderm derived from human embryonic stem cells generates glucose-responsive insulin-secreting cells in vivo. Nat. Biotech. 26, 443\u2013452 (2008).","journal-title":"Nat. Biotech."},{"key":"375_CR43","doi-asserted-by":"publisher","first-page":"1392","DOI":"10.1038\/nbt1259","volume":"24","author":"KA D\u2019Amour","year":"2006","unstructured":"D\u2019Amour, K. A. et al. Production of pancreatic hormone-expressing endocrine cells from human embryonic stem cells. Nat. Biotechnol. 24, 1392\u20131401 (2006).","journal-title":"Nat. Biotechnol."},{"key":"375_CR44","doi-asserted-by":"publisher","first-page":"412","DOI":"10.1016\/j.mod.2011.08.001","volume":"128","author":"X Xu","year":"2011","unstructured":"Xu, X., Browning, V. L. & Odorico, J. S. Activin, BMP and FGF pathways cooperate to promote endoderm and pancreatic lineage cell differentiation from human embryonic stem cells. Mech. Dev. 128, 412\u2013427 (2011).","journal-title":"Mech. Dev."},{"key":"375_CR45","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1007\/s00125-011-2344-9","volume":"55","author":"M Riedel","year":"2012","unstructured":"Riedel, M. et al. Immunohistochemical characterisation of cells co-producing insulin and glucagon in the developing human pancreas. Diabetologia 55, 372\u2013381 (2012).","journal-title":"Diabetologia"},{"key":"375_CR46","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1038\/nbt.3033","volume":"32","author":"A Rezania","year":"2014","unstructured":"Rezania, A. et al. Reversal of diabetes with insulin-producing cells derived in vitro from human pluripotent stem cells. Nat. Biotech. 32, 1121\u20131133 (2014).","journal-title":"Nat. Biotech."},{"key":"375_CR47","doi-asserted-by":"publisher","first-page":"428","DOI":"10.1016\/j.cell.2014.09.040","volume":"159","author":"FW Pagliuca","year":"2014","unstructured":"Pagliuca, F. W. et al. Generation of functional human pancreatic \u03b2 cells in vitro. Cell 159, 428\u2013439 (2014).","journal-title":"Cell"},{"key":"375_CR48","doi-asserted-by":"publisher","first-page":"1759","DOI":"10.15252\/embj.201591058","volume":"34","author":"HA Russ","year":"2015","unstructured":"Russ, H. A. et al. Controlled induction of human pancreatic progenitors produces functional beta-like cells in vitro. EMBO J. 34, 1759\u20131772 (2015).","journal-title":"EMBO J."},{"key":"375_CR49","doi-asserted-by":"publisher","first-page":"2016","DOI":"10.2337\/db11-1711","volume":"61","author":"A Rezania","year":"2012","unstructured":"Rezania, A. et al. Maturation of human embryonic stem cell-derived pancreatic progenitors into functional islets capable of treating pre-existing diabetes in mice. Diabetes 61, 2016\u20132029 (2012).","journal-title":"Diabetes"},{"key":"375_CR50","doi-asserted-by":"publisher","first-page":"368","DOI":"10.1038\/s41586-019-1168-5","volume":"569","author":"A Veres","year":"2019","unstructured":"Veres, A. et al. Charting cellular identity during human in vitro \u03b2-cell differentiation. Nature 569, 368\u2013373 (2019). This is an important study that used single-cell RNA sequencing to elucidate generation of several additional cell types including \u03b1-like cells, enterochromaffin cells and non-endocrine cells in stem cell differentiation toward \u03b2-cells. Furthermore, the authors identified CD49a as a cell surface marker to sort, re-aggregate and enrich for stem cell-derived \u03b2-cells.","journal-title":"Nature"},{"key":"375_CR51","doi-asserted-by":"publisher","first-page":"1551","DOI":"10.1016\/j.stemcr.2018.11.008","volume":"11","author":"NAJ Krentz","year":"2018","unstructured":"Krentz, N. A. J. et al. Single-cell transcriptome profiling of mouse and hESC-derived pancreatic progenitors. Stem Cell Rep. 11, 1551\u20131564 (2018).","journal-title":"Stem Cell Rep."},{"key":"375_CR52","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1038\/s41556-018-0271-4","volume":"21","author":"GG Nair","year":"2019","unstructured":"Nair, G. G. et al. Recapitulating endocrine cell clustering in culture promotes maturation of human stem-cell-derived \u03b2 cells. Nat. Cell Biol. 21, 263\u2013274 (2019). This study demonstrated for the first time that re-aggregation and clustering of stem cell-derived immature \u03b2-cells induces maturation by activating mitochondrial respiration. The resulting \u03b2-cells closely resemble adult islet \u03b2-cells in transcriptome and exhibit similar functional properties such as dynamic insulin secretion.","journal-title":"Nat. Cell Biol."},{"key":"375_CR53","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1016\/j.stemcr.2018.12.012","volume":"12","author":"L Velazco-Cruz","year":"2019","unstructured":"Velazco-Cruz, L. et al. Acquisition of dynamic function in human stem cell-derived \u03b2 cells. Stem Cell Rep. 12, 351\u2013365 (2019).","journal-title":"Stem Cell Rep."},{"key":"375_CR54","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1007\/s00125-013-3062-2","volume":"57","author":"M Nomura","year":"2014","unstructured":"Nomura, M. et al. SMAD2 disruption in mouse pancreatic beta cells leads to islet hyperplasia and impaired insulin secretion due to the attenuation of ATP-sensitive K+ channel activity. Diabetologia 57, 157\u2013166 (2014).","journal-title":"Diabetologia"},{"key":"375_CR55","doi-asserted-by":"publisher","first-page":"1060","DOI":"10.1016\/0006-291X(89)92829-5","volume":"158","author":"Y Totsuka","year":"1989","unstructured":"Totsuka, Y. et al. Stimulation of insulin secretion by transforming growth factor-\u03b2. Biochem. Biophys. Res. Commun. 158, 1060\u20131065 (1989).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"375_CR56","doi-asserted-by":"publisher","first-page":"12246","DOI":"10.1074\/jbc.M805379200","volume":"284","author":"H-M Lin","year":"2009","unstructured":"Lin, H.-M. et al. Transforming growth factor-\u03b2\/Smad3 signaling regulates insulin gene transcription and pancreatic islet \u03b2-cell function. J. Biol. Chem. 284, 12246\u201312257 (2009).","journal-title":"J. Biol. Chem."},{"key":"375_CR57","doi-asserted-by":"publisher","first-page":"1827","DOI":"10.1210\/en.2018-00048","volume":"159","author":"N Saber","year":"2018","unstructured":"Saber, N. et al. Sex differences in maturation of human embryonic stem cell-derived \u03b2 cells in mice. Endocrinology 159, 1827\u20131841 (2018).","journal-title":"Endocrinology"},{"key":"375_CR58","doi-asserted-by":"publisher","first-page":"1297","DOI":"10.2337\/db15-1439","volume":"65","author":"JE Bruin","year":"2016","unstructured":"Bruin, J. E. et al. Hypothyroidism impairs human stem cell-derived pancreatic progenitor cell maturation in mice. Diabetes 65, 1297\u20131309 (2016).","journal-title":"Diabetes"},{"key":"375_CR59","doi-asserted-by":"publisher","first-page":"E838","DOI":"10.1152\/ajpendo.00219.2014","volume":"307","author":"E Mott\u00e9","year":"2014","unstructured":"Mott\u00e9, E. et al. Composition and function of macroencapsulated human embryonic stem cell-derived implants: comparison with clinical human islet cell grafts. Am. J. Physiol. Endocrinol. Metab. 307, E838\u2013E846 (2014).","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"issue":"Suppl. 1","key":"375_CR60","first-page":"A37","volume":"67","author":"RR Henry","year":"2018","unstructured":"Henry, R. R. et al. Initial clinical evaluation of VC-01TM combination product \u2014 a stem cell\u2013derived islet replacement for type 1 diabetes (T1D) [abstract 138-OR]. Diabetes 67 (Suppl. 1), A37 (2018).","journal-title":"Diabetes"},{"key":"375_CR61","doi-asserted-by":"crossref","unstructured":"Shapiro, A.J. et al. Insulin expression and glucose-responsive circulating C-peptide in type 1 diabetes patients implanted subcutaneously with pluripotent stem cell-derived pancreatic endoderm cells in a macro-device. Preprint at SSRN http:\/\/dx.doi.org\/10.2139\/ssrn.3501034 (2019).","DOI":"10.2139\/ssrn.3501034"},{"key":"375_CR62","doi-asserted-by":"publisher","first-page":"953","DOI":"10.2337\/db18-0788","volume":"68","author":"AR Pepper","year":"2019","unstructured":"Pepper, A. R. et al. Post-transplant characterization of long-term functional hESC-derived pancreatic endoderm grafts. Diabetes 68, 953\u2013962 (2019).","journal-title":"Diabetes"},{"key":"375_CR63","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1152\/physiol.00029.2018","volume":"33","author":"MO Huising","year":"2018","unstructured":"Huising, M. O. et al. The difference \u03b4-cells make in glucose control. Physiology 33, 403\u2013411 (2018).","journal-title":"Physiology"},{"key":"375_CR64","doi-asserted-by":"publisher","first-page":"239","DOI":"10.2337\/db10-0573","volume":"60","author":"A Rezania","year":"2011","unstructured":"Rezania, A. et al. Production of functional glucagon-secreting \u03b1-cells from human embryonic stem cells. Diabetes 60, 239\u2013247 (2011).","journal-title":"Diabetes"},{"key":"375_CR65","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1016\/j.gde.2015.03.004","volume":"32","author":"G Nair","year":"2015","unstructured":"Nair, G. & Hebrok, M. Islet formation in mice and men: lessons for the generation of functional insulin-producing \u03b2-cells from human pluripotent stem cells. Curr. Opin. Genet. Dev. 32, 171\u2013180 (2015).","journal-title":"Curr. Opin. Genet. Dev."},{"key":"375_CR66","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1172\/JCI29115","volume":"117","author":"P Collombat","year":"2007","unstructured":"Collombat, P. et al. Embryonic endocrine pancreas and mature \u03b2 cells acquire \u03b1 and PP cell phenotypes upon Arx misexpression. J. Clin. Investig. 117, 961\u2013970 (2007).","journal-title":"J. Clin. Investig."},{"key":"375_CR67","doi-asserted-by":"publisher","first-page":"2591","DOI":"10.1101\/gad.269003","volume":"17","author":"P Collombat","year":"2003","unstructured":"Collombat, P. et al. Opposing actions of Arx and Pax4 in endocrine pancreas development. Genes Dev. 17, 2591\u20132603 (2003).","journal-title":"Genes Dev."},{"key":"375_CR68","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1003934","volume":"9","author":"M Courtney","year":"2013","unstructured":"Courtney, M. et al. The inactivation of Arx in pancreatic \u03b1-cells triggers their neogenesis and conversion into functional \u03b2-like cells. PLoS Genet. 9, e1003934 (2013).","journal-title":"PLoS Genet."},{"key":"375_CR69","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1016\/j.cmet.2017.01.009","volume":"25","author":"H Chakravarthy","year":"2017","unstructured":"Chakravarthy, H. et al. Converting adult pancreatic islet \u03b1 cells into \u03b2 cells by targeting both Dnmt1 and Arx. Cell Metab. 25, 622\u2013634 (2017).","journal-title":"Cell Metab."},{"key":"375_CR70","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1016\/j.stem.2017.11.020","volume":"22","author":"X Xiao","year":"2018","unstructured":"Xiao, X. et al. Endogenous reprogramming of alpha cells into beta cells, induced by viral gene therapy, reverses autoimmune diabetes. Cell Stem Cell 22, 78\u201390 (2018).","journal-title":"Cell Stem Cell"},{"key":"375_CR71","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1038\/s41586-019-0942-8","volume":"567","author":"K Furuyama","year":"2019","unstructured":"Furuyama, K. et al. Diabetes relief in mice by glucose-sensing insulin-secreting human \u03b1-cells. Nature 567, 43\u201348 (2019). This is the first study to show that \u03b1-cells can be reprogrammed to insulin-secreting cells. The authors isolated human islet non-\u03b2-cells, such as \u03b1-cells and \u03b3-cells, from donors with and without diabetes mellitus and reprogrammed them into insulin-secreting cells with PDX1 and MAFA. These cells were able to reverse diabetes mellitus in mice whilst still retaining certain \u03b1-cell features.","journal-title":"Nature"},{"key":"375_CR72","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.1038\/nature08894","volume":"464","author":"F Thorel","year":"2010","unstructured":"Thorel, F. et al. Conversion of adult pancreatic \u03b1-cells to \u03b2-cells after extreme \u03b2-cell loss. Nature 464, 1149\u20131154 (2010).","journal-title":"Nature"},{"key":"375_CR73","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1038\/nature13633","volume":"514","author":"S Chera","year":"2014","unstructured":"Chera, S. et al. Diabetes recovery by age-dependent conversion of pancreatic \u03b4-cells into insulin producers. Nature 514, 503\u2013507 (2014).","journal-title":"Nature"},{"key":"375_CR74","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1016\/j.cmet.2017.03.017","volume":"25","author":"T van der Meulen","year":"2017","unstructured":"van der Meulen, T. et al. Virgin beta cells persist throughout life at a neogenic niche within pancreatic islets. Cell Metab. 25, 911\u2013926.e6 (2017).","journal-title":"Cell Metab."},{"key":"375_CR75","doi-asserted-by":"publisher","first-page":"2601","DOI":"10.2337\/db18-0155","volume":"67","author":"Y-S Lee","year":"2018","unstructured":"Lee, Y.-S. et al. Glucagon-like peptide-1 increases \u03b2-cell regeneration by promoting \u03b1- to \u03b2-cell transdifferentiation. Diabetes 67, 2601\u20132614 (2018).","journal-title":"Diabetes"},{"key":"375_CR76","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.cell.2016.11.002","volume":"168","author":"N Ben-Othman","year":"2017","unstructured":"Ben-Othman, N. et al. Long-term GABA administration induces alpha cell-mediated beta-like cell neogenesis. Cell 168, 73\u201385 (2017).","journal-title":"Cell"},{"key":"375_CR77","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1016\/j.cell.2016.11.010","volume":"168","author":"J Li","year":"2017","unstructured":"Li, J. et al. Artemisinins target GABA a receptor signaling and impair \u03b1 cell identity. Cell 168, 86\u2013100 (2017).","journal-title":"Cell"},{"key":"375_CR78","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1016\/j.cmet.2017.10.002","volume":"27","author":"T van der Meulen","year":"2018","unstructured":"van der Meulen, T. et al. Artemether does not turn \u03b1 cells into \u03b2 cells. Cell Metab. 27, 218\u2013225 (2018).","journal-title":"Cell Metab."},{"key":"375_CR79","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1016\/j.cmet.2018.07.002","volume":"28","author":"AM Ackermann","year":"2018","unstructured":"Ackermann, A. M., Moss, N. G. & Kaestner, K. H. GABA and artesunate do not induce pancreatic \u03b1-to-\u03b2 cell transdifferentiation in vivo. Cell Metab. 28, 787\u2013792.e3 (2018).","journal-title":"Cell Metab."},{"key":"375_CR80","doi-asserted-by":"publisher","first-page":"712","DOI":"10.1016\/j.stemcr.2018.02.005","volume":"10","author":"CJM Loomans","year":"2018","unstructured":"Loomans, C. J. M. et al. Expansion of adult human pancreatic tissue yields organoids harboring progenitor cells with endocrine differentiation potential. Stem Cell Rep. 10, 712\u2013724 (2018).","journal-title":"Stem Cell Rep."},{"key":"375_CR81","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0133862","volume":"10","author":"DL Gomez","year":"2015","unstructured":"Gomez, D. L. et al. Neurogenin 3 expressing cells in the human exocrine pancreas have the capacity for endocrine cell fate. PLoS One 10, e0133862 (2015).","journal-title":"PLoS One"},{"key":"375_CR82","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1016\/j.stem.2016.03.016","volume":"18","author":"CB Westphalen","year":"2016","unstructured":"Westphalen, C. B. et al. Dclk1 defines quiescent pancreatic progenitors that promote injury-induced regeneration and tumorigenesis. Cell Stem Cell 18, 441\u2013455 (2016).","journal-title":"Cell Stem Cell"},{"key":"375_CR83","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1073\/pnas.0609490104","volume":"104","author":"T Sugiyama","year":"2007","unstructured":"Sugiyama, T. et al. Conserved markers of fetal pancreatic epithelium permit prospective isolation of islet progenitor cells by FACS. Proc. Natl Acad. Sci. USA 104, 175\u2013180 (2007).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"375_CR84","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.cmet.2017.08.007","volume":"27","author":"C Aguayo-Mazzucato","year":"2018","unstructured":"Aguayo-Mazzucato, C. & Bonner-Weir, S. Pancreatic \u03b2 cell regeneration as a possible therapy for diabetes. Cell Metab. 27, 57\u201367 (2018).","journal-title":"Cell Metab."},{"key":"375_CR85","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1038\/nature07314","volume":"455","author":"Q Zhou","year":"2008","unstructured":"Zhou, Q. et al. In vivo reprogramming of adult pancreatic exocrine cells to \u03b2-cells. Nature 455, 627\u2013632 (2008).","journal-title":"Nature"},{"key":"375_CR86","doi-asserted-by":"publisher","first-page":"2821","DOI":"10.2337\/db12-1256","volume":"62","author":"MJ Lima","year":"2013","unstructured":"Lima, M. J. et al. Suppression of epithelial-to-mesenchymal transitioning enhances ex vivo reprogramming of human exocrine pancreatic tissue toward functional insulin-producing \u03b2-like cells. Diabetes 62, 2821\u20132833 (2013).","journal-title":"Diabetes"},{"key":"375_CR87","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0156204","volume":"11","author":"MJ Lima","year":"2016","unstructured":"Lima, M. J. et al. Generation of functional beta-like cells from human exocrine pancreas. PLoS One 11, e0156204 (2016).","journal-title":"PLoS One"},{"key":"375_CR88","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0087812","volume":"9","author":"D Berneman-Zeitouni","year":"2014","unstructured":"Berneman-Zeitouni, D. et al. The temporal and hierarchical control of transcription factors-induced liver to pancreas transdifferentiation. PLoS One 9, e87812 (2014).","journal-title":"PLoS One"},{"key":"375_CR89","doi-asserted-by":"publisher","first-page":"568","DOI":"10.1038\/75050","volume":"6","author":"S Ferber","year":"2000","unstructured":"Ferber, S. et al. Pancreatic and duodenal homeobox gene 1 induces expression of insulin genes in liver and ameliorates streptozotocin-induced hyperglycemia. Nat. Med. 6, 568\u2013572 (2000).","journal-title":"Nat. Med."},{"key":"375_CR90","doi-asserted-by":"publisher","first-page":"7964","DOI":"10.1073\/pnas.0405277102","volume":"102","author":"T Sapir","year":"2005","unstructured":"Sapir, T. et al. Cell-replacement therapy for diabetes: generating functional insulin-producing tissue from adult human liver cells. Proc. Natl Acad. Sci. USA 102, 7964\u20137969 (2005).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"375_CR91","doi-asserted-by":"publisher","DOI":"10.1007\/s11892-019-1198-2","volume":"19","author":"I Meivar-Levy","year":"2019","unstructured":"Meivar-Levy, I. & Ferber, S. Liver to pancreas transdifferentiation. Curr. Diabetes Rep. 19, 76 (2019).","journal-title":"Curr. Diabetes Rep."},{"key":"375_CR92","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms14127","volume":"8","author":"N Cerd\u00e1-Esteban","year":"2017","unstructured":"Cerd\u00e1-Esteban, N. et al. Stepwise reprogramming of liver cells to a pancreas progenitor state by the transcriptional regulator Tgif2. Nat. Commun. 8, 14127 (2017).","journal-title":"Nat. Commun."},{"key":"375_CR93","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0181812","volume":"12","author":"F Galivo","year":"2017","unstructured":"Galivo, F. et al. Reprogramming human gallbladder cells into insulin-producing \u03b2-like cells. PLoS One 12, e0181812 (2017).","journal-title":"PLoS One"},{"key":"375_CR94","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1038\/ng.2215","volume":"44","author":"C Talchai","year":"2012","unstructured":"Talchai, C. et al. Generation of functional insulin-producing cells in the gut by Foxo1 ablation. Nat. Genet. 44, 406\u2013412. (2012).","journal-title":"Nat. Genet."},{"key":"375_CR95","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1038\/nri.2017.100","volume":"17","author":"I Spadoni","year":"2017","unstructured":"Spadoni, I., Fornasa, G. & Rescigno, M. Organ-specific protection mediated by cooperation between vascular and epithelial barriers. Nat. Rev. Immunol. 17, 761\u2013773 (2017).","journal-title":"Nat. Rev. Immunol."},{"key":"375_CR96","doi-asserted-by":"publisher","first-page":"410","DOI":"10.1016\/j.stem.2016.01.003","volume":"18","author":"C Ariyachet","year":"2016","unstructured":"Ariyachet, C. et al. Reprogrammed stomach tissue as a renewable source of functional \u03b2 cells for blood glucose regulation. Cell Stem Cell 18, 410\u2013421 (2016). Antral endocrine cells were reprogrammed into insulin\n+ cells that express NKX6.1 and PC2 at a greater efficiency than enteroendocrine cells; bioengineered stomach mini-organs produced renewable insulin\n+ cells in vivo and reversed hyperglycaemia in mice.","journal-title":"Cell Stem Cell"},{"key":"375_CR97","doi-asserted-by":"publisher","first-page":"1046","DOI":"10.1016\/j.celrep.2014.02.013","volume":"6","author":"Y-J Chen","year":"2014","unstructured":"Chen, Y.-J. et al. De novo formation of insulin-producing \u201cneo-\u03b2 cell islets\u201d from intestinal crypts. Cell Rep. 6, 1046\u20131058 (2014).","journal-title":"Cell Rep."},{"key":"375_CR98","doi-asserted-by":"publisher","first-page":"1296","DOI":"10.1126\/science.aaf4006","volume":"354","author":"M Xie","year":"2016","unstructured":"Xie, M. et al. \u03b2-cell\u2013mimetic designer cells provide closed-loop glycemic control. Science 354, 1296\u20131301 (2016).","journal-title":"Science"},{"key":"375_CR99","doi-asserted-by":"publisher","first-page":"1565","DOI":"10.1126\/science.1203535","volume":"332","author":"H Ye","year":"2011","unstructured":"Ye, H. et al. A synthetic optogenetic transcription device enhances blood-glucose homeostasis in mice. Science 332, 1565\u20131568 (2011).","journal-title":"Science"},{"key":"375_CR100","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.aal2298","volume":"9","author":"J Shao","year":"2017","unstructured":"Shao, J. et al. Smartphone-controlled optogenetically engineered cells enable semiautomatic glucose homeostasis in diabetic mice. Sci. Transl Med. 9, eaal2298 (2017).","journal-title":"Sci. Transl Med."},{"key":"375_CR101","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1038\/nchembio.2511","volume":"14","author":"Z Chen","year":"2017","unstructured":"Chen, Z. et al. Synthetic beta cells for fusion-mediated dynamic insulin secretion. Nat. Chem. Biol. 14, 86\u201393 (2017).","journal-title":"Nat. Chem. Biol."},{"key":"375_CR102","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0037004","volume":"7","author":"TC Schulz","year":"2012","unstructured":"Schulz, T. C. et al. A scalable system for production of functional pancreatic progenitors from human embryonic stem cells. PLoS One 7, e37004 (2012).","journal-title":"PLoS One"},{"key":"375_CR103","first-page":"315ra189","volume":"7","author":"JA Bluestone","year":"2015","unstructured":"Bluestone, J. A. et al. Type 1 diabetes immunotherapy using polyclonal regulatory T cells. Sci. Transl Med. 7, 315ra189 (2015).","journal-title":"Sci. Transl Med."},{"key":"375_CR104","doi-asserted-by":"publisher","first-page":"1817","DOI":"10.2337\/dc12-0038","volume":"35","author":"N Marek-Trzonkowska","year":"2012","unstructured":"Marek-Trzonkowska, N. et al. Administration of CD4+CD25highCD127\u2212 regulatory T cells preserves \u03b2-cell function in type 1 diabetes in children. Diabetes Care 35, 1817\u20131820 (2012).","journal-title":"Diabetes Care"},{"key":"375_CR105","doi-asserted-by":"publisher","first-page":"927","DOI":"10.5966\/sctm.2015-0058","volume":"4","author":"TC Schulz","year":"2015","unstructured":"Schulz, T. C. Concise review: manufacturing of pancreatic endoderm cells for clinical trials in type 1 diabetes. Stem Cell Transl Med. 4, 927\u2013931 (2015).","journal-title":"Stem Cell Transl Med."},{"key":"375_CR106","doi-asserted-by":"publisher","DOI":"10.1186\/1479-5876-10-69","volume":"10","author":"RP Somerville","year":"2012","unstructured":"Somerville, R. P. et al. Clinical scale rapid expansion of lymphocytes for adoptive cell transfer therapy in the WAVE(R) bioreactor. J. Transl Med. 10, 69 (2012).","journal-title":"J. Transl Med."},{"key":"375_CR107","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1016\/j.omtm.2018.01.006","volume":"8","author":"H Fraser","year":"2018","unstructured":"Fraser, H. et al. A rapamycin-based GMP-compatible process for the isolation and expansion of regulatory T cells for clinical trials. Mol. Ther. Methods Clin. Dev. 8, 198\u2013209 (2018).","journal-title":"Mol. Ther. Methods Clin. Dev."},{"key":"375_CR108","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1089\/ten.tec.2009.0228","volume":"16","author":"R Krawetz","year":"2010","unstructured":"Krawetz, R. et al. Large-scale expansion of pluripotent human embryonic stem cells in stirred-suspension bioreactors. Tissue Eng. Part C. Methods 16, 573\u2013582 (2010).","journal-title":"Tissue Eng. Part C. Methods"},{"key":"375_CR109","doi-asserted-by":"publisher","first-page":"1132","DOI":"10.1016\/j.stemcr.2014.09.017","volume":"3","author":"H Kempf","year":"2014","unstructured":"Kempf, H. et al. Controlling expansion and cardiomyogenic differentiation of human pluripotent stem cells in scalable suspension culture. Stem Cell Rep. 3, 1132\u20131146 (2014).","journal-title":"Stem Cell Rep."},{"key":"375_CR110","doi-asserted-by":"publisher","first-page":"2051","DOI":"10.1089\/ten.tea.2008.0455","volume":"15","author":"LT Lock","year":"2009","unstructured":"Lock, L. T. & Tzanakakis, E. S. Expansion and differentiation of human embryonic stem cells to endoderm progeny in a microcarrier stirred-suspension culture. Tissue Eng. Part A 15, 2051\u20132063 (2009). This is the first report of hESC expansion and directed differentiation to definitive endoderm cells in a microcarrier stirred suspension (spinner flask) culture.","journal-title":"Tissue Eng. Part A"},{"key":"375_CR111","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1089\/ten.tec.2012.0146","volume":"19","author":"J Bardy","year":"2013","unstructured":"Bardy, J. et al. Microcarrier suspension cultures for high-density expansion and differentiation of human pluripotent stem cells to neural progenitor cells. Tissue Eng. Part C. Methods 19, 166\u2013180 (2013).","journal-title":"Tissue Eng. Part C. Methods"},{"key":"375_CR112","doi-asserted-by":"publisher","first-page":"1397","DOI":"10.3727\/096368910X513955","volume":"19","author":"D Jing","year":"2010","unstructured":"Jing, D., Parikh, A. & Tzanakakis, E. S. Cardiac cell generation from encapsulated embryonic stem cells in static and scalable culture systems. Cell Transpl. 19, 1397\u20131412 (2010).","journal-title":"Cell Transpl."},{"key":"375_CR113","doi-asserted-by":"publisher","first-page":"1804","DOI":"10.1089\/scd.2017.0090","volume":"26","author":"G Meng","year":"2017","unstructured":"Meng, G. et al. Optimizing human induced pluripotent stem cell expansion in stirred-suspension culture. Stem Cell Dev. 26, 1804\u20131817 (2017).","journal-title":"Stem Cell Dev."},{"key":"375_CR114","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1089\/ten.tec.2013.0040","volume":"20","author":"MM Hunt","year":"2014","unstructured":"Hunt, M. M. et al. Factorial experimental design for the culture of human embryonic stem cells as aggregates in stirred suspension bioreactors reveals the potential for interaction effects between bioprocess parameters. Tissue Eng. Part C. Methods 20, 76\u201389 (2014). A systematic analysis based on factorial design of experiments was performed to investigate adjustable parameters (for example, agitation rate and cell seeding density) and their interaction on the outcome of hESC cultures in spinner flasks.","journal-title":"Tissue Eng. Part C. Methods"},{"key":"375_CR115","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1038\/nrendo.2016.178","volume":"13","author":"AM Shapiro","year":"2017","unstructured":"Shapiro, A. M., Pokrywczynska, M. & Ricordi, C. Clinical pancreatic islet transplantation. Nat. Rev. Endocrinol. 13, 268\u2013277 (2017).","journal-title":"Nat. Rev. Endocrinol."},{"key":"375_CR116","doi-asserted-by":"publisher","first-page":"1661","DOI":"10.1038\/labinvest.2010.124","volume":"90","author":"A Pisania","year":"2010","unstructured":"Pisania, A. et al. Quantitative analysis of cell composition and purity of human pancreatic islet preparations. Lab. Invest. 90, 1661\u20131675 (2010).","journal-title":"Lab. Invest."},{"key":"375_CR117","doi-asserted-by":"publisher","first-page":"418","DOI":"10.1089\/ten.tea.2016.0400","volume":"24","author":"Y Yan","year":"2018","unstructured":"Yan, Y. et al. Derivation of cortical spheroids from human induced pluripotent stem cells in a suspension bioreactor. Tissue Eng. Part A 24, 418\u2013431 (2018).","journal-title":"Tissue Eng. Part A"},{"key":"375_CR118","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/j.reth.2018.11.003","volume":"10","author":"SG Yabe","year":"2019","unstructured":"Yabe, S. G. et al. Induction of functional islet-like cells from human iPS cells by suspension culture. Regen. Ther. 10, 69\u201376 (2019).","journal-title":"Regen. Ther."},{"key":"375_CR119","doi-asserted-by":"publisher","first-page":"3193","DOI":"10.1002\/term.2228","volume":"11","author":"Y Mihara","year":"2017","unstructured":"Mihara, Y. et al. Production of pancreatic progenitor cells from human induced pluripotent stem cells using a three-dimensional suspension bioreactor system. J. Tissue Eng. Regen. Med. 11, 3193\u20133201 (2017).","journal-title":"J. Tissue Eng. Regen. Med."},{"key":"375_CR120","doi-asserted-by":"publisher","first-page":"1289","DOI":"10.5966\/sctm.2015-0253","volume":"5","author":"C Kropp","year":"2016","unstructured":"Kropp, C. et al. Impact of feeding strategies on the scalable expansion of human pluripotent stem cells in single-use stirred tank bioreactors. Stem Cell Transl Med. 5, 1289\u20131301 (2016). In this elegant study, the authors studied the effect of different feeding strategies on hPSCs cultured in automated SSBs and demonstrated a shift in metabolism from glycolysis to OXPHOS without differentiation.","journal-title":"Stem Cell Transl Med."},{"key":"375_CR121","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1089\/ten.tea.2009.0454","volume":"16","author":"DE Kehoe","year":"2010","unstructured":"Kehoe, D. E. et al. Scalable stirred-suspension bioreactor culture of human pluripotent stem cells. Tissue Eng. Part A 16, 405\u2013421 (2010).","journal-title":"Tissue Eng. Part A"},{"key":"375_CR122","doi-asserted-by":"publisher","first-page":"771","DOI":"10.1208\/s12248-014-9598-3","volume":"16","author":"LX Yu","year":"2014","unstructured":"Yu, L. X. et al. Understanding pharmaceutical quality by design. AAPS J. 16, 771\u2013783 (2014).","journal-title":"AAPS J."},{"key":"375_CR123","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1038\/nbt.3525","volume":"34","author":"YY Lipsitz","year":"2016","unstructured":"Lipsitz, Y. Y., Timmins, N. E. & Zandstra, P. W. Quality cell therapy manufacturing by design. Nat. Biotechnol. 34, 393\u2013400 (2016).","journal-title":"Nat. Biotechnol."},{"key":"375_CR124","doi-asserted-by":"publisher","first-page":"728","DOI":"10.1038\/nrm.2017.108","volume":"18","author":"KH Vining","year":"2017","unstructured":"Vining, K. H. & Mooney, D. J. Mechanical forces direct stem cell behaviour in development and regeneration. Nat. Rev. Mol. Cell Biol. 18, 728\u2013742 (2017).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"375_CR125","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1038\/s41586-018-0762-2","volume":"564","author":"A Mamidi","year":"2018","unstructured":"Mamidi, A. et al. Mechanosignalling via integrins directs fate decisions of pancreatic progenitors. Nature 564, 114\u2013118 (2018).","journal-title":"Nature"},{"key":"375_CR126","doi-asserted-by":"publisher","first-page":"460","DOI":"10.1038\/s41587-020-0430-6","volume":"38","author":"NJ Hogrebe","year":"2020","unstructured":"Hogrebe, N. J. et al. Targeting the cytoskeleton to direct pancreatic differentiation of human pluripotent stem cells. Nat. Biotechnol. 38, 460\u2013470 (2020).","journal-title":"Nat. Biotechnol."},{"key":"375_CR127","doi-asserted-by":"publisher","first-page":"1510","DOI":"10.1021\/acsbiomaterials.6b00253","volume":"3","author":"Y Fan","year":"2017","unstructured":"Fan, Y., Zhang, F. & Tzanakakis, E. S. Engineering xeno-free microcarriers with recombinant vitronectin, albumin and UV irradiation for human pluripotent stem cell bioprocessing. ACS Biomater. Sci. Eng. 3, 1510\u20131518 (2017).","journal-title":"ACS Biomater. Sci. Eng."},{"key":"375_CR128","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0102486","volume":"9","author":"J Wu","year":"2014","unstructured":"Wu, J. et al. Oxygen transport and stem cell aggregation in stirred-suspension bioreactor cultures. PLoS One 9, e102486 (2014).","journal-title":"PLoS One"},{"key":"375_CR129","doi-asserted-by":"publisher","first-page":"662","DOI":"10.2337\/db09-0891","volume":"59","author":"M Heinis","year":"2010","unstructured":"Heinis, M. et al. Oxygen tension regulates pancreatic \u03b2-cell differentiation through hypoxia-inducible factor 1\u03b1. Diabetes 59, 662\u2013669 (2010).","journal-title":"Diabetes"},{"key":"375_CR130","doi-asserted-by":"publisher","first-page":"9623","DOI":"10.1074\/jbc.M113.524363","volume":"289","author":"F Hakim","year":"2014","unstructured":"Hakim, F. et al. High oxygen condition facilitates the differentiation of mouse and human pluripotent stem cells into pancreatic progenitors and insulin-producing cells. J. Biol. Chem. 289, 9623\u20139638 (2014).","journal-title":"J. Biol. Chem."},{"key":"375_CR131","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.gde.2017.06.008","volume":"46","author":"TS Cliff","year":"2017","unstructured":"Cliff, T. S. & Dalton, S. Metabolic switching and cell fate decisions: implications for pluripotency, reprogramming and development. Curr. Opin. Genet. Dev. 46, 44\u201349 (2017).","journal-title":"Curr. Opin. Genet. Dev."},{"key":"375_CR132","doi-asserted-by":"publisher","first-page":"721","DOI":"10.1002\/stem.404","volume":"28","author":"A Prigione","year":"2010","unstructured":"Prigione, A. et al. The senescence-related mitochondrial\/oxidative stress pathway is repressed in human induced pluripotent stem cells. Stem Cell 28, 721\u2013733 (2010).","journal-title":"Stem Cell"},{"key":"375_CR133","doi-asserted-by":"publisher","first-page":"745","DOI":"10.1016\/j.cmet.2018.10.007","volume":"29","author":"DC Saunders","year":"2019","unstructured":"Saunders, D. C. et al. Ectonucleoside triphosphate diphosphohydrolase-3 antibody targets adult human pancreatic \u03b2 cells for in vitro and in vivo analysis. Cell Metab. 29, 745\u2013754 (2019). A novel cell surface biomarker NTPDase 3 was found to be enriched in adult human \u03b2-cells. NTPDase 3 antibodies were shown to be applicable to live sorting of \u03b2-cells from human islets and in vivo imaging of transplanted \u03b2-cells.","journal-title":"Cell Metab."},{"key":"375_CR134","doi-asserted-by":"publisher","first-page":"267","DOI":"10.4161\/isl.3.5.15939","volume":"3","author":"A Steffen","year":"2011","unstructured":"Steffen, A. et al. Functional assessment of automatically sorted pancreatic islets using large particle flow cytometry. Islets 3, 267\u2013270 (2011).","journal-title":"Islets"},{"key":"375_CR135","doi-asserted-by":"publisher","first-page":"1531","DOI":"10.1517\/17425255.2012.724058","volume":"8","author":"EB Rangel","year":"2012","unstructured":"Rangel, E. B. The metabolic and toxicological considerations for immunosuppressive drugs used during pancreas transplantation. Expert. Opin. Drug Metab. Toxicol. 8, 1531\u20131548 (2012).","journal-title":"Expert. Opin. Drug Metab. Toxicol."},{"key":"375_CR136","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1038\/nbt0708-739","volume":"26","author":"N Nakatsuji","year":"2008","unstructured":"Nakatsuji, N., Nakajima, F. & Tokunaga, K. HLA-haplotype banking and iPS cells. Nat. Biotechnol. 26, 739\u2013740 (2008).","journal-title":"Nat. Biotechnol."},{"key":"375_CR137","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/j.stem.2012.07.014","volume":"11","author":"CraigJ Taylor","year":"2012","unstructured":"Taylor, CraigJ. et al. Generating an iPSC bank for HLA-matched tissue transplantation based on known donor and recipient HLA types. Cell Stem Cell 11, 147\u2013152 (2012).","journal-title":"Cell Stem Cell"},{"key":"375_CR138","doi-asserted-by":"publisher","first-page":"180","DOI":"10.1002\/stem.772","volume":"30","author":"P-A Gourraud","year":"2012","unstructured":"Gourraud, P.-A. et al. The role of human leukocyte antigen matching in the development of multiethnic \u201chaplobank\u201d of induced pluripotent stem cell lines. Stem Cells 30, 180\u2013186 (2012).","journal-title":"Stem Cells"},{"key":"375_CR139","doi-asserted-by":"publisher","first-page":"228","DOI":"10.1038\/mt.2012.245","volume":"21","author":"T Thatava","year":"2013","unstructured":"Thatava, T. et al. Intrapatient variations in type 1 diabetes-specific iPS cell differentiation into insulin-producing cells. Mol. Ther. 21, 228\u2013239 (2013).","journal-title":"Mol. Ther."},{"key":"375_CR140","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1038\/nrd.2016.232","volume":"16","author":"T Desai","year":"2017","unstructured":"Desai, T. & Shea, L. D. Advances in islet encapsulation technologies. Nat. Rev. Drug Discov. 16, 338\u2013350 (2017).","journal-title":"Nat. Rev. Drug Discov."},{"key":"375_CR141","doi-asserted-by":"publisher","first-page":"a015552","DOI":"10.1101\/cshperspect.a015552","volume":"3","author":"Q Tang","year":"2013","unstructured":"Tang, Q. & Bluestone, J. A. Regulatory T-cell therapy in transplantation: moving to the clinic. Cold Spring Harb. Perspect. Med. 3, a015552 (2013).","journal-title":"Cold Spring Harb. Perspect. Med."},{"key":"375_CR142","doi-asserted-by":"publisher","first-page":"749","DOI":"10.1038\/s41573-019-0041-4","volume":"18","author":"LMR Ferreira","year":"2019","unstructured":"Ferreira, L. M. R. et al. Next-generation regulatory T cell therapy. Nat. Rev. Drug Discov. 18, 749\u2013769 (2019).","journal-title":"Nat. Rev. Drug Discov."},{"key":"375_CR143","doi-asserted-by":"publisher","first-page":"10441","DOI":"10.1073\/pnas.1902566116","volume":"116","author":"X Han","year":"2019","unstructured":"Han, X. et al. Generation of hypoimmunogenic human pluripotent stem cells. Proc. Natl Acad. Sci. USA 116, 10441\u201310446 (2019).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"375_CR144","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1038\/s41587-019-0016-3","volume":"37","author":"T Deuse","year":"2019","unstructured":"Deuse, T. et al. Hypoimmunogenic derivatives of induced pluripotent stem cells evade immune rejection in fully immunocompetent allogeneic recipients. Nat. Biotechnol. 37, 252\u2013258 (2019). In this breakthrough study on the generation of universal \u2018off the shelf\u2019 hPSCs, deletion of MHC class I and II genes and overexpression of CD47 rendered both mouse and human iPSCs hypoimmunogenic. The engineered PSCs differentiated to smooth muscle cells, endothelial cells or cardiomyocytes that successfully evaded immune rejection in allogeneic fully MHC-mismatched recipents.","journal-title":"Nat. Biotechnol."},{"key":"375_CR145","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-09404-6","volume":"10","author":"EA Rosado-Olivieri","year":"2019","unstructured":"Rosado-Olivieri, E. A. et al. YAP inhibition enhances the differentiation of functional stem cell-derived insulin-producing \u03b2 cells. Nat. Commun. 10, 1464 (2019).","journal-title":"Nat. Commun."},{"key":"375_CR146","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1016\/j.celrep.2019.04.083","volume":"27","author":"N Sharon","year":"2019","unstructured":"Sharon, N. et al. Wnt signaling separates the progenitor and endocrine compartments during pancreas development. Cell Rep. 27, 2281\u20132291 (2019).","journal-title":"Cell Rep."},{"key":"375_CR147","doi-asserted-by":"publisher","first-page":"E640","DOI":"10.1152\/ajpendo.00251.2015","volume":"309","author":"J-C Henquin","year":"2015","unstructured":"Henquin, J.-C. et al. Dynamics of glucose-induced insulin secretion in normal human islets. Am. J. Physiol. Endocrinol. Metab. 309, E640\u2013E650 (2015).","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"375_CR148","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1146\/annurev-nutr-071812-161201","volume":"34","author":"MR Hayes","year":"2014","unstructured":"Hayes, M. R. et al. Incretins and amylin: neuroendocrine communication between the gut, pancreas, and brain in control of food intake and blood glucose. Annu. Rev. Nutr. 34, 237\u2013260 (2014).","journal-title":"Annu. Rev. Nutr."},{"key":"375_CR149","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1152\/advan.00167.2012","volume":"37","author":"DP Begg","year":"2013","unstructured":"Begg, D. P. & Woods, S. C. Interactions between the central nervous system and pancreatic islet secretions: a historical perspective. Adv. Physiol. Educ. 37, 53\u201360 (2013).","journal-title":"Adv. Physiol. Educ."},{"key":"375_CR150","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1042\/BJ20141384","volume":"466","author":"GA Rutter","year":"2015","unstructured":"Rutter, G. A. et al. Pancreatic \u03b2-cell identity, glucose sensing and the control of insulin secretion. Biochem. J. 466, 203\u2013218 (2015).","journal-title":"Biochem. J."},{"key":"375_CR151","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1016\/j.cmet.2016.03.005","volume":"23","author":"E Yoshihara","year":"2016","unstructured":"Yoshihara, E., et al. ERR\u03b3 is required for the metabolic maturation of therapeutically functional glucose-responsive \u03b2 cells. Cell Metab. 23, 622\u2013634 (2016).","journal-title":"Cell Metab."},{"key":"375_CR152","doi-asserted-by":"publisher","first-page":"1019","DOI":"10.1007\/s00125-010-1685-0","volume":"53","author":"S Jitrapakdee","year":"2010","unstructured":"Jitrapakdee, S. et al. Regulation of insulin secretion: role of mitochondrial signalling. Diabetologia 53, 1019\u20131032 (2010).","journal-title":"Diabetologia"},{"key":"375_CR153","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1016\/j.gene.2019.04.063","volume":"706","author":"M Vakilian","year":"2019","unstructured":"Vakilian, M., Tahamtani, Y. & Ghaedi, K. A review on insulin trafficking and exocytosis. Gene 706, 52\u201361 (2019).","journal-title":"Gene"}],"container-title":["Nature Reviews Endocrinology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41574-020-0375-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41574-020-0375-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41574-020-0375-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,4,17]],"date-time":"2022-04-17T18:06:49Z","timestamp":1650218809000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41574-020-0375-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,6,25]]},"references-count":153,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2020,9]]}},"alternative-id":["375"],"URL":"https:\/\/doi.org\/10.1038\/s41574-020-0375-3","relation":{},"ISSN":["1759-5029","1759-5037"],"issn-type":[{"value":"1759-5029","type":"print"},{"value":"1759-5037","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,6,25]]},"assertion":[{"value":"20 May 2020","order":1,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 June 2020","order":2,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"M.H. is affiliated with Semma Therapeutics (Consultant and SAB member) and Encellin Inc. (SAB member, stock holder). M.H. also holds stocks from Viacyte Inc.. The other authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}]}}