The emerging role of ACE2 in physiology and disease

Journal of Pathology

Volumn 212, Issue 1, 2007, Pages 1-11

  Hamming, I ^a   Cooper, M E ^b,e   Haagmans, B L ^c   Hooper, N M ^d   Korstanje, R ^a   Osterhaus, A D M E ^c   Timens, W ^a   Turner, A J ^d   Navis, G ^a   van Goor, H ^a  

^a UNIVERSITY MEDICAL CENTER GRONINGEN   (Netherlands)

^b BAKER IDI HEART AND DIABETES INSTITUTE   (Australia)

^c ERASMUS MEDICAL CENTER   (Netherlands)

^d UNIVERSITY OF LEEDS   (United Kingdom)

^e UNIVERSITY OF MELBOURNE   (Australia)

Author keywords

ACE; ACE2; Angiotensin; RAAS; Renin

Indexed keywords

ALDOSTERONE ANTAGONIST; ANGIOTENSIN RECEPTOR ANTAGONIST; DIPEPTIDYL CARBOXYPEPTIDASE; DIPEPTIDYL CARBOXYPEPTIDASE 2; DIPEPTIDYL CARBOXYPEPTIDASE 2 INHIBITOR; DIPEPTIDYL CARBOXYPEPTIDASE INHIBITOR; MLN 4760; UNCLASSIFIED DRUG;

BLOOD PRESSURE REGULATION; CARDIOVASCULAR DISEASE; DIABETES MELLITUS; DIABETIC NEPHROPATHY; DIPEPTIDYL CARBOXYPEPTIDASE DEFICIENCY; DRUG DESIGN; DRUG SCREENING; ENZYME ACTIVITY; ENZYME ANALYSIS; ENZYME DEFICIENCY; ENZYME DEGRADATION; ENZYME LOCALIZATION; ENZYME METABOLISM; ENZYME SPECIFICITY; ENZYME SUBSTRATE; GENE EXPRESSION REGULATION; GENETIC LINKAGE; HEART FAILURE; HEART FUNCTION; HEART PROTECTION; HUMAN; HYPERTENSION; KIDNEY DISEASE; LUNG DISEASE; MOLECULAR CLONING; MOLECULAR INTERACTION; NONHUMAN; PATHOPHYSIOLOGY; PREECLAMPSIA; PREGNANCY; PRIORITY JOURNAL; PROTEIN BINDING; PROTEIN EXPRESSION; PROTEIN LOCALIZATION; PROTEIN PURIFICATION; PROTEIN STRUCTURE; RENAL PROTECTION; RENIN ANGIOTENSIN ALDOSTERONE SYSTEM; REVIEW; SEVERE ACUTE RESPIRATORY SYNDROME; SINGLE NUCLEOTIDE POLYMORPHISM; X CHROMOSOME;

ALDOSTERONE; ANIMALS; CARDIOVASCULAR DISEASES; CORONAVIRUS; FEMALE; HUMANS; HYPERTENSION; KIDNEY; LUNG DISEASES; MALE; MYOCARDIUM; PEPTIDYL-DIPEPTIDASE A; PREGNANCY; PREGNANCY COMPLICATIONS; RENIN-ANGIOTENSIN SYSTEM; SARS VIRUS;

EID: 34248597758     PISSN: 00223417     EISSN: 10969896     Source Type: Journal    
DOI: 10.1002/path.2162     Document Type: Review

References (102)

1. Feffario CM, Trask AJ, Jessup JA. Advances in biochemical and functional roles of angiotensin-converting enzyme 2 and angiotensin-(1-7) in regulation of cardiovascular function. Am J Physiol Heart Circ Physiol 2005;289:H2281 -112290.
2. Donoghue M, Hsieh F, Baronas E, Godbout K, Gosselin M, Stagliano N, et al. A novel angiotensin-converting enzyme-related carboxypeptidase (ACE2) converts angiotensin I to angiotensin 1-9. Circ Res 2000; 87:E1-E9.
3. Tipnis SR, Hooper NM, Hyde R, Karran E, Christie G, Turner AJ. A human homolog of angiotensin-converting enzyme. Cloning and functional expression as a captopril-insensitive carboxypeptidase. J Biol Chem 2000;275:33238-33243.
4. Doobay MF, Talman LS, Obr TD, Tian X, Davisson RL, Lazartigues E. Differential expression of neuronal ACE2 in transgenic mice with overexpression of the brain renin-angiotensin system. Am J Physiol Regul Integr Comp Physiol 2007;292:R373-R381.
5. Gembardt F, Sterner-Kock A, Imboden H, Spalteholz M, Reibitz F, Schultheiss HP, et al. Organ-specific distribution of ACE2 mRNA and correlating peptidase activity in rodents. Peptides 2005; 26:1270-1277.
6. Hamming I, Timens W, Bulthuis M, Lely A, Navis G, Van Goor H. Tissue distribution of ACE2 protein, the functional receptor for SARS coronavirus. A first step in understanding SARS pathogenesis. J Pathol 2004;203:631-637.
7. Harmer D, Gilbert M, Borman R, Clark KL. Quantitative mRNA expression profiling of ACE 2, a novel homologue of angiotensin converting enzyme. FEBS Lett 2002;532:107-110.
8. Komatsu T, Suzuki Y, Imai J, Sugano S, Hida M, Tanigami A, et al. Molecular cloning, mRNA expression and chromosomal localization of mouse angiotensin-converting enzyme-related carboxypeptidase (mACE2). DNA Seq 2002;13:217-220.
9. Itoyama S, Keicho N, Hijikata M, Quy T, Phi NC, Long HT, et al. Identification of an alternative 5′-untranslated exon and new polymorphisms of angiotensin-converting enzyme 2 gene: lack of association with SARS in the Vietnamese population. Am J Med Genet A2005; 136:52-57.
10. Zhang H, Wada J, Hida K, Tsuchiyama Y, Hiragushi K, Shikata K, et al. Collectrin, a collecting duct-specific transmembrane glycoprotein, is a novel homolog of ACE2 and is developmentally regulated in embryonic kidneys. J Biol Chem 2001; 276:17132-17139.
11. Akpinar P, Kuwajima S, Krutzfeldt J, Stoffel M. Tmem27: a cleaved and shed plasma membrane protein that stimulates pancreatic beta cell proliferation. Cell Metab 2005;2:385-397.
12. Fukui K, Yang Q, Cao Y, Takahashi N, Hatakeyama H, Wang H, et al. The HNF-1 target collectrin controls insulin exocytosis by SNARE complex formation. Cell Metab 2005;2:373-384.
13. Malakauskas SM, Quan H, Fields TA, McCall SJ, Yu MJ, Kourany WM, et al. Aminoaciduria and altered renal expression of luminal amino acid transporters in mice lacking novel gene collectrin. Am J Physiol Renal Physiol 2006;292:F533-F544.
14. Rice GI, Thomas DA, Grant PJ, Turner AJ, Hooper NM. Evaluation of angiotensin converting enzyme (ACE), its homologue ACE2 and neprilysin in angiotensin peptide metabolism. Biochem J 2004; 383:45-51.
15. Brosnihan KB, Neves LA, Joyner J, Averill DB, Chappell MC, Sarao R, et al. Enhanced renal inummocytochemical expression of ANG-(1-7) and ACE2 during pregnancy. Hypertension 2003;42:749-753.
16. Campbell DJ, Zeitz CJ, Esler MD, Horowitz JD. Evidence against a major role for angiotensin converting enzyme-related carboxypeptidase (ACE2) in angiotensin peptide metabolism in the human coronary circulation. J Hypertens 2004;22:1971-1976.
17. Elased KM, Cunha TS, Gurley SB, Coffman TM, Morris M. New mass spectrometric assay for angiotensin-converting enzyme 2 activity. Hypertension 2006;47:1010-1017.
18. Ferrario CM, Averill DB, Brosnihan KB, Chappell MC, Iskandar SS, Dean RH, et al. Vasopeptidase inhibition and Ang-(1-7) in the spontaneously hypertensive rat. Kidney Int 2002; 62:1349-1357.
19. Li N, Zimpelmann J, Cheng K, Wilkins JA, Bums KD. The role of angiotensin converting enzyme 2 in the generation of angiotensin 1-7 by rat proximal tubules. Am J Physiol Renal Physiol 2005; 288:F353-F362.
20. Zisman LS, Keller RS, Weaver B, Lin Q, Speth R, Bristow MR, et al. Increased angiotensin-(1-7)-forming activity in failing human heart ventricles: evidence for upregulation of the angiotensin-converting enzyme homologue ACE2. Circulation 2003;108:1707-1712.
21. Ocaranza MP, Godoy I, Jalil JE, Varas M, Collantes P, Pinto M, et al. Enalapril attenuates downregulation of angiotensin-converting enzyme 2 in the late phase of ventricular dysfunction in myocardial infarcted rat. Hypertension 2006;48:572-578.
22. Vickers C, Hales P, Kaushik V, Dick L, Gavin J, Tang J, et al. Hydrolysis of biological peptides by human angiotensin-converting enzyme-related carboxypeptidase. J Biol Chem 2002; 277:14838-14843.
23. Warner FJ, Lew RA, Smith AI, Lambert DW, Hooper NM, Turner AJ. Angiotensin-converting enzyme 2 (ACE2), but not ACE, is preferentially localized to the apical surface of polarized kidney cells. J Biol Chem 2005;280:39353-39362.
24. Ren X, Glende J, Al Falah M, de Vries V, Schwegmann-Wessels C, Qu X, et al. Analysis of ACE2 in polarized epithelial cells: surface expression and function as receptor for severe acute respiratory syndrome-associated coronavirus. J Gen Virol 2006; 87:1691-1695.
25. Lambert DW, Yarski M, Warner FJ, Thornhill P, Parkin ET, Smith AI, et al. Tumor necrosis factor-alpha convertase (ADAM17) mediates regulated ectodomain shedding of the severe-acute respiratory syndrome-coronavirus (SARS-CoV) receptor, angiotensin-converting enzyme-2 (ACE2). J Biol Chem 2005;280:30113-30119.
26. Shaltout HA, Westwood B, Averill DB, Ferrario CM, Figueroa J, Diz DI, et al. Angiotensin metabolism in renal proximal tubules, urine and serum of sheep: evidence for ACE2-dependent processing of angiotensin 11. Am J Physiol Renal Physiol 2006;292:F82-F91.
27. Burrell LM, Risvanis J, Kubota E, Dean RG, MacDonald PS, Lu S, et al. Myocardial infarction increases ACE2 expression in rat and humans. Ear Heart J 2005;26:369-375.
28. Crackower MA, Sarao R, Oudit GY, Yagil C, Kozieradzki I, Scanga SE, et al. Angiotensin-converting enzyme 2 is an essential regulator of heart function. Nature 2002;417:822-828.
29. Gurley SB, Allred A, Le TH, Griffiths R, Mao L, Philip N, et al. Altered blood pressure responses and normal cardiac phenotype in ACE2-null mice. J Clin Invest 2006;116:2218-2225.
30. Yamamoto K, Ohishi M, Katsuya T, Ito N, Ikushima M, Kaibe M, et al. Deletion of angiotensin-converting enzyme 2 accelerates pressure overload-induced cardiac dysfunction by increasing local angiotensin II. Hypertension 2006;47:718-726.
31. Donoghue M, Wakimoto H, Maguire CT, Acton S, Hales P, Stagliano N, et al. Heart block, ventricular tachycardia, and sudden death in ACE2 transgenic mice with downregulated connexins. J Mol Cell Cardiol 2003;35:1043 -1053.
32. Goulter AB, Goddard MJ, Allen JC, Clark KL. ACE2 gene expression is up-regulated in the human failing heart. BMC Med 2004;2:19.
33. Ishiyama Y, Gallagher PE, Averill DB, Tallant EA, Brosnihan KB, Ferrario CM. Upregulation of angiotensin-converting enzyme 2 after myocardial infarction by blockade of angiotensin II receptors. Hypertension 2004;43:970-976.
34. Karram T, Abbasi A, Keidar S, Golomb E, Hochberg I, Winaver J, et al. Effects of spironolactone and eprosartan on cardiac remodeling and angiotensin-converting enzyme isoforms in rats with experimental heart failure. Am J Physiol Heart Circ Physiol 2005; 289:H1351-H1358.
35. Feffario CM, Jessup J, Chappell MC, Averill DB, Brosnihan KB, Tallant EA, et al. Effect of angiotensin-converting enzyme inhibition and angiotensin II receptor blockers on cardiac angiotensin-converting enzyme 2. Circulation 2005;111:2605-2610.
36. Keidar S, Gamliel-Lazarovich A, Kaplan M, Pavlotzky E, Hamoud S, Hayek T, et al. Mineralocorticoid receptor blocker increases angiotensin-converting enzyme 2 activity in congestive heart failure patients. Circ Res 2005;97:946-953.
37. Pitt B, Remme W, Zannad F, Neaton J, Martinez F, Roniker B, et al. Eplerenone, a selective aldosterone blocker, in patients with left ventricular dysfunction after myocardial infarction. N Engl J Med 2003;348:1309-1321.
38. Santos RA, Simoes e Silva AC, Maric C, Silva DM, Machado RP, de Buhr I, et al. Angiotensin-(1-7) is an endogenous ligand for the G protein-coupled receptor Mas. Proc Natl Acad Sci U S A 2003; 100:8258-8263.
39. Huentelman MJ, Grobe JL, Vazquez J, Stewart JM, Mecca AP, Katovich MJ, et al. Protection from angiotensin II-induced cardiac hypertrophy and fibrosis by systemic lentiviral delivery of ACE2 in rats. Exp Physiol 2005;90:783-790.
40. Diez-Freire C, Vazquez J, Correa de Adjounian MF, Ferrari MF, Yuan L, Silver X, et al. ACE2 gene transfer attenuates hypertension-linked pathophysiological changes in the SHR. Physiol Genomics 2006; 27:12-19.
41. Yagil Y, Yagil C. Hypothesis: ACE2 modulates blood pressure in the mammalian organism. Hypertension 2003;41:871-873.
42. Hilbert P, Lindpaintner K, Beckmann JS, Serikawa T, Soubrier F, Dubay C, et al. Chromosomal mapping of two genetic loci associated with blood-pressure regulation in hereditary hypertensive rats. Nature 1991;353:521-529.
43. Yagil C, Sapojnikov M, Kreutz R, Zurcher H, Ganten D, Yagil Y. Role of chromosome X in the Sabra rat model of salt-sensitive hypertension. Hypertension 1999;33:261-265.
44. Zhong JC, Huang DY, Yang YM, Li YF, Liu GF, Song XH, et al. Upregulation of angiotensin-converting enzyme 2 by all-trans retinoic acid in spontaneously hypertensive rats. Hypertension 2004; 44:907-912.
45. Tikellis C, Cooper ME, Bialkowski K, Johnston CI, Bums WC, Lew RA, et al. Developmental expression of ACE2 in the SHR kidney: a role in hypertension? Kidney Int 2006;70:34-41.
46. Hamming I, Kreutz R, Sluimer J, Bolbrinker J, Bernardy C, Walther T, et al. Renal angiotensin-converting enzyme 2 is unaltered in experimental hypertension. (Abstract.) J Am Soc Nephrol 2005; 16:TH-PO294.
47. Benjafield AV, Wang WY, Morris BJ. No association of angiotensin-converting enzyme 2 gene (ACE2) polymorphisms with essential hypertension. Am J Hypertens 2004;17:624-628.
48. Yi L, Gu YH, Wang XL, An LZ, Xie XD, Shao W, et al. Association of ACE, ACE2 and UTS2 polymorphisms with essential hypertension in Han and Dongxiang populations from north-western China. J Int Med Res 2006;34:272-283.
49. Zhong J, Yart Z, Liu D, Ni Y, Zhao Z, Zhu S, et al. Association of angiotensin-converting enzyme 2 gene A/G polymorphism and elevated blood pressure in Chinese patients with metabolic syndrome. J Lab Clin Med 2006;147:91-95.
50. Hamming I, Timens W, Bulthuis ML, Lely AT, Navis GJ, Van Goor H. Tissue distribution of ACE2 protein, the functional receptor for SARS coronavirus. A first step in understanding SARS pathogenesis. J Pathol 2004;203:631-637.
51. Lely AT, Hamming I, Van Goor H, Navis GJ. Renal ACE2 expression in human kidney disease. J Pathol 2004;204:587-593.
52. Ye M, Wysocki J, William J, Soler MJ, Cokic I, Batlle D. Glomerular localization and expression of angiotensin-converting enzyme 2 and angiotensin-converting enzyme: implications for albuminuria in diabetes. J Am Soc Nephrol 2006;17:3067-3075.
53. Ye M, Wysocki J, Naaz P, Salabat MR, LaPointe MS, Batlle D. Increased ACE 2 and decreased ACE protein in renal tubules from diabetic mice. A renoprotective combination? Hypertension 2004;43:1120-1125.
54. Dzau VJ, Bernstein K, Celermajer D, Cohen J, Dahlof B, Deanfield J, et al. The relevance of tissue angiotensin-converting enzyme: manifestations in mechanistic and endpoint data. Am J Cardiol 2001;88:1L-20L.
55. Huang W, Gallois Y, Bouby N, Bruneval P, Heudes D, Belair MF, et al. Genetically increased angiotensin I-converting enzyme level and renal complications in the diabetic mouse. Proc Natl Acad Sci USA 2001;98:13330-13334.
56. Rook M, Lely AT, Kramer AB, Van Goor H, Navis G. Individual differences in renal ACE activity in healthy rats predict suscetibility to adriamycin-unduced renal damage. Nephrol Dial Transplant 2005; 20:59-64.
57. Ruiz-Ortega M, Ruperez M, Esteban V, Rodriguez-Vita J, Sanchez-Lopez E, Carvajal G, et al. Angiotensin II: a key factor in the inflammatory and fibrotic response in kidney diseases. [ Nephrol Dial Transplant 2006;21:16-20.
58. Wolf G, Ritz E. Combination therapy with ACE inhibitors and angiotensin II receptor blockers to halt progression of chronic renal disease: pathophysiology and indications. Kidney Int 2005; 67:799-812.
59. Tikellis C, Johnston CI, Forbes JM, Burns WC, Burrell LM, Risvanis J, et al. Characterization of renal angiotensin-converting enzyme 2 in diabetic nephropathy. Hypertension 2003;41:392-397.
60. Jessup J, Gallagher PE, Averill DB, Brosnihan KB, Tallant EA, Chappell M, et al. Effect of angiotensin II blockade on a new congenic model of hypertension derived from transgenic Ren-2 rats. Am J Physiol Heart Circ Physiol 2006;291:H2166-H2172.
61. Hamming I, Navis G, Akkermans KE, Van Goor H. ACE inhibition during low salt intake decreases renal ACE2 expression in healthy and nephrotic rats. (Abstract.) J Hypertension 2006;24:S295.
62. Hamming I, Navis G, Kocks M, Van Goor H. ACE inhibition has adverse renal effects during dietary sodium restriction in proteinuric and healthy rats. J Pathol 2006;209:129-139.
63. Oudit GY, Herzenberg AM, Kassiri Z, Wong D, Reich H, Khokha R, et al. Loss of angiotensin-converting enzyme-2 leads to the late development of angiotensin II-dependent glomerulosclerosis. Am J Pathol 2006;168:1808-1820.
64. Wysocki J, Ye M, Soler MJ, Gurley SB, Xiao HD, Bernstein KE, et al. ACE and ACE2 activity in diabetic mice. Diabetes 2006; 55 :2132-2139.
65. Konoshita T, Wakahara S, Mizuno S, Motomura M, Aoyama C, Makino Y, et al. Tissue gene expression of renin-angiotensin system in human type 2 diabetic nephropathy. Diabetes Care 2006;29:848-852.
66. Frojdo S, Sjolind L, Parkkonen M, Makinen VP, Kilpikari R, Pettersson-Fernholm K, et al. Polymorphisms in the gene encoding angiotensin I converting enzyme 2 and diabetic nephropathy. Diabetologia 2005;48:2278-2281.
67. Valdes G, Neves LA, Anton L, Corthom J, Chacon C, Germain AM, et al. Distribution of angiotensin-(1-7) and ACE2 in human placentas of normal and pathological pregnancies. Placenta 2006; 27:200-207.
68. Marshall RP. The pulmonary renin-angiotensin system. Carr Phann Des 2003;9:715-722.
69. Orte C, Polak JM, Haworth SG, Yacoub MH, Morrell NW. Expression of pulmonary vascular angiotensin-converting enzyme in primary and secondary plexiform pulmonary hypertension. J Pathol 2000; 192:379-384.
70. Morrell NW, Morris KG, Stenmark KR. Role of angiotensin-converting enzyme and angiotensin II in development of hypoxic pulmonary hypertension. Am J Physiol 1995;269:H1186-H1194.
71. Loddenkemper R, Kloppenborg A, Schoenfeld N, Grosser H, Costabel U. Clinical findings in 715 patients with newly detected pulmonary sarcoidosis - results of a cooperative study in former West Germany and Switzerland. WATL Study Group. Wissenschaftliche Arbeitsgemeinschaft fur die Therapie von Lungenkrarikheitan. Sarcoidosis Vasc Diffuse Lung Dis 1998;15:178-182.
72. Csaszar A, Halmos B, Palicz T, Szalai C, Romics L. Interpopulation effect of ACE I/D polymorphism on serum concentration of ACE in diagnosis of sarcoidosis. Lancet1997;350:518.
73. Specks U, Martin WJ, Rohrbach MS. Bronchoalveolar lavage fluid angiotensin-converting enzyme in interstitial lung diseases. Am Rev Respir Dis 1990;141:117-123.
74. Fourrier F, Chopin C, Wallaert B, Mazurier C, Mangalaboyi J, Durocher A. Compared evolution of plasma fibronectin and angiotensin-converting enzyme levels in septic ARDS. Chest 1985;87:191-195.
75. Idell S, Kueppers F, Lippmann M, Rosen H, Niederman M, Fein A. Angiotensin converting enzyme in bronchoalveolar lavage in ARDS. Chest 1987;91:52-56.
76. Imai Y, Kuba K, Rao S, Huan Y, Guo F, Guan B, et al. Angiotensin-converting enzyme 2 protects from severe acute lung failure. Nature 2005;436:112-116.
77. Kuba K, Imai Y, Penninger JM. Angiotensin-converting enzyme 2 in lung diseases. Curr Opin Pharmacol 2006;6:271-276.
78. Hofmann H, Pyrc K, van der Hoek L, Geier M, Berkhout B, Pohlmann S. Human coronavirus NL63 employs the severe acute respiratory syndrome coronavirus receptor for cellular entry. Proc Natl Acad Sci USA 2005;102:7988-7993.
79. Li W, Moore MJ, Vasilieva N, Sui J, Wong SK, Berne MA, et al. Angiotensin-converfing enzyme 2 is a functional receptor for the SARS coronavirus. Nature 2003;426:450-454.
80. Weiss SR, Navas-Martin S. Coronavirus pathogenesis and the emerging pathogen severe acute respiratory syndrome coronavirus. Microbiol Mol Biol Rev 2005;69:635-664.
81. Peiris JS, Lai ST, Poon LL, Guan Y, Yam LY, Lim W, et al. Coronavirus as a possible cause of severe acute respiratory syndrome. Lancet 2003;361:1319-1325.
82. Li F, Li W, Farzan M, Harrison SC. Structure of SARS coronavirus spike receptor-binding domain complexed with receptor. Science 2005; 309:1864-1868.
83. Wong SK, Li W, Moore MJ, Choe H, Farzan M. A 193-amino acid fragment of the SARS coronavirus S protein efficiently binds angiotensin-converting enzyme 2. J Biol Chem 2004;279:3197-3201.
84. Han DP, Penn-Nicholson A, Cho MW. Identification of critical determinants on ACE2 for SARS-CoV entry and development of a potent entry inhibitor. Virology 2006;350:15-25.
85. Mossel EC, Huang C, Narayanan K, Makino, S, Tesh RB, Peters CJ. Exogenous ACE2 expression allows refractory cell lines to support severe acute respiratory syndrome coronavirus replication. J Virol 2005;79:3846-3850.
86. Kuba K, Imai Y, Rao S, Gao H, Guo F, Guan B, et al. A crucial role of angiotensin converting enzyme 2 (ACE2) in SARS coronavirus-induced lung injury. Nature Med 2005;11:875-879.
87. He L, Ding Y, Zhang Q, Che X, He Y, Shen H, et al. Expression of elevated levels of pro-inflammatory cytokines in SARS-CoV-infected ACE2+ cells in SARS patients: relation to the acute lung injury and pathogenesis of SARS. J Pathol 2006;210:288-297.
88. Huang IC, Bosch BJ, Li F, Li W, Lee KH, Ghiran S, et al. SARS coronavirus, but not human coronavirus NL63, utilizes cathepsin L to infect ACE2-expressing cells. J Biol Chem 2006; 281:3198-3203.
89. Simmons G, Gosalia DN, Rennekamp AJ, Reeves JD, Diamond SL, Bates P. Inhibitors of cathepsin L prevent severe acute respiratory syndrome coronavirus entry. Proc Natl Acad Sci U S A 2005; 102:11876-11881.
90. Jeffers SA, Tusell SM, Gillim-Ross L, Hemmila EM, Achenbach JE, Babcock GJ, et al. CD209L (L-SIGN) is a receptor for severe acute respiratory syndrome coronavirus. Proc Natl Acad Sci USA 2004; 101:15748-15753.
91. Li W, Greenough TC, Moore MJ, Vasilieva N, Somasundaran M, Sullivan JL, et al. Efficient replication of severe acute respiratory syndrome coronavirus in mouse cells is limited by marine angiotensin-converting enzyme 2. J Virol 2004;78:11429-11433.
92. Li W, Zhang C, Sui J, Kuhn JH, Moore MJ, Luo S, et al. Receptor and viral determinants of SARS-coronavirus adaptation to human ACE2. EMBO J 2005;24:1634-1643.
93. Liang G, Chen Q, Xu J, Liu Y, Lim W, Peiris JS, et al. Laboratory diagnosis of four recent sporadic cases of community-acquired SARS, Guangdong Province, China. Emerg Infect Dis 2004; 10:1774-1781.
94. de Lang A, Osterhaus AD, Haagmans BL. Interferon-gamma and interleukin-4 downregulate expression of the SARS coronavirus receptor ACE2 in Vero E6 cells. Virology 2006;353:474-481.
95. Jia HP, Look DC, Shi L, Hickey M, Pewe L, Netland J, et al. ACE2 receptor expression and severe acute respiratory syndrome coronavirus infection depend on differentiation of human airway epithelia. J Virol 2005;79:14614-14621.
96. Chou CF, Loh CB, Foo YK, Shen S, Fielding BC, Tan TH, et al. ACE2 orthologues in non-mammalian vertebrates (Danio, Gallus, Fugu, Tetraodon and Xenopus). Gene 2006;377:46-55.
97. Chiu RW, Tang NL, Hui DS, Chung GT, Chim SS, Chan KC, et al. ACE2 gene polymorphisms do not affect outcome of severe acute respiratory syndrome. Clin Chem 2004;50:1683-1686.
98. Dales NA, Gould AE, Brown JA, Calderwood EF, Guan B, Minor CA, et al. Substrate-based design of the first class of angiotensin-converting enzyme-related carboxypeptidase (ACE2) inhibitors. J Am Chem Soc 2002;124:11852-11853.
99. Huentelman MJ, Zubcevic J, Hernandez Prada JA, Xiao X, Dimitrov DS, Raizada MK, et al. Structure-based discovery of a novel angiotensin-converting enzyme 2 inhibitor. Hypertension 2004; 44:903-906.
100. Rella M, Rushworth CA, Guy JL, Turner AJ, Langer T, Jackson RM. Structure-based pharmacophore design and virtual screening for novel angiotensin converting enzyme 2 inhibitors. J Chem Inf Model 2006;46:708-716.
101. Warner FJ, Smith AI, Hooper NM, Turner AJ. Angiotensin-converting enzyme-2: a molecular and cellular perspective. Cell Mol Life Sci 2004;61:2704-2713.
102. Turner AJ, Hooper NM. The angiotensin-converting enzyme gene family: genomics and pharmacology. Trends Pharmacol Sci 2002; 23:177-183.