squalene synthase mechanism

yeast squalene synthase gene was a single copy gene that was essential for the growth of yeast2). not otherwise permitted to reproduce, republish, redistribute, or sell any Supporting Information [31][32], Squalene synthase homolog inhibition in Staphylococcus aureus is currently being investigated as a virulence factor-based antibacterial therapy. Identification of unique mechanisms for triterpene biosynthesis in Botryococcus braunii. Selenium-containing analogues of WC-9 are extremely potent inhibitors of Trypanosoma cruzi proliferation. PDB. FPP is a soluble allylic compound containing 15 carbon atoms (C15), whereas squalene is an insoluble, C30 isoprenoid. You have to login with your ACS ID befor you can login with your Mendeley account. [11] Starting at the top of the catalytic cycle below, the reaction begins with the ionization of FPP to generate an allylic carbocation. Absolute Configuration of Hydroxysqualene. [5] SQS is anchored to the membrane by a short C-terminal membrane-spanning domain. Erich J. Molitor, Beth M. Paschal, Hung-wen Liu. Shashikumar K. Paknikar, Kamlesh Pai Fondekar. J Am Chem Soc, 124(30):8846-8853, 01 Jul 2002 Cited by 28 articles | PMID: 12137537 Florence Bouvier, Alain Rahier, Bilal Camara. Of the three known SREBP transcription factors, only SREBP-1a and SREBP-2 activate SQS gene transcription in transgenic mouse livers. Methylococcus capsulatus Biological Chemistry Trainee (NIH Grant GM 80357). Sterol and Steroid Biosynthesis and Metabolism in Plants and Microorganisms. Some of the results of these studies have been previously reported in the form of an abstract (22, 23). A Mechanism for the Rearrangement of Presqualene Diphosphate to Squalene, Contribution from the Department of Chemistry, University of Utah, 315 South 1400 East Room 2020, Salt Lake City, Utah 84112. Therefore, both the structure and reaction mechanism of squalene synthases aremarkedlyconservedin taxonomically remote eukaryotes. Three tables showing a list of proteins used in multiple sequence alignments, pairwise percent identities among squalene synthases, and pairwise identities among phytoene synthases and a dehydrosqualene synthase; four figures showing amino acid sequence alignments for 19 squalene synthases, a phylogenetic analysis of squalene synthases based on amino acid sequences, amino acid sequence alignments for 26 phytoene synthases and a dehydrosqualene synthase, and a phylogenetic analysis of phytoene synthases and a dehydrosqualene synthase based on amino acid sequences (PDF). FDFT1 encodes for an evolutionarily conserved enzyme, squalene synthase (SS, farnesyl-pyrophosphate farnesyl-transferase 1), which catalyzes the first committed step in cholesterol biosynthesis. Squalene synthase (SSN, EC 2.5.1.21), a major enzyme in the sterol biosynthetic pathway, catalyses an unusual head-to-head reductive dimerization of two molecules of farnesyl-pyrophosphate (FPP) in a two-step reaction to form squalene. Mendelian disorders of cholesterol biosynthesis typically result in multi-system clinical phenotypes, underlining the importance of cholesterol in embryogenesis and development. The structure and absolute stereochemistry of the tertiary cyclopropylcarbinyl alcohol were established by synthesis using two independent routes. Ali Koohang, Jessica L. Bailey, Robert M. Coates, Hans K. Erickson, David Owen and C. Dale Poulter . discovering peptide inhibitors of human squalene synthase through screening the phage-displayed cyclic peptide c7c library, molecular modelling and ADMET predictions of the selected peptides, using the crystal structure PDB 1EZF, a homo-trimer protein complex (with inhibitors CP-320473, CP-458003 and CP-424677), as the target, overview. With the identification and molecular characterization of squalene synthase in tea plant, next, we can ask the questions about the roles of squalene as a volatile product as well as a precursor for triterpenoids, which may promote product development from diverse tea materials and mining of excellent tea germplasm resources. Squalene synthase catalyzes two consecutive reactions in sterol biosynthesis the condensation of two molecules of farnesyl diphosphate (FPP) to form the cyclopropylcarbinyl intermediate presqualene diphosphate (PSPP) and the subsequent rearrangement and reduction of PSPP to form squalene. SQS belongs to squalene/phytoene synthase family of proteins. Squalene synthase (SS) represents a putative branch point in the isoprenoid biosynthetic pathway capable of diverting carbon flow specifically to the biosynthesis of sterols and, hence, is considered a potential regulatory point for sterol metabolism. [2] In rSQS, Tyr-171 was converted to aromatic residues Phe and Trp, as well as hydroxyl-containing residue Ser. S. K. Paknikar, F. S. Kamounah, P. E. Hansen, M. S. Wadia. View protein in InterPro IPR008949, Isoprenoid_synthase_dom_sf IPR002060, Squ/phyt_synthse IPR006449, Squal_synth-like IPR019845, Squalene/phytoene_synthase_CS IPR033904, Trans_IPPS_HH: SFLD i: SFLDG01018, Squalene/Phytoene_Synthase_Lik, 1 hit Squalene synthase (SSN, EC 2.5.1.21), a major enzyme in the sterol biosynthetic pathway, catalyses an unusual head-to-head reductive dimerization of two molecules of farnesyl-pyrophosphate (FPP) in a two-step reaction to form squalene. In the first half-reaction, two molecules of FPP react to form the stable presqualene diphosphate intermediate (PSQPP), with concomitant release of a proton and a molecule of inorganic diphosphate. [§ 1], Squalene synthase (SQS) is an enzyme participating in the isoprenoid biosynthetic pathway. High levels of LDL-derived cholesterol inhibit HMG-CoA reductase activity significantly, since mevalonate is no longer needed for sterol production. Squalene synthase (SQase) catalyzes the condensation of two molecules of farnesyl diphosphate (FPP) to form presqualene diphosphate (PSPP) and the subsequent rearrangement and NADPH-dependent reduction of PSPP to squalene (SQ). 1). Synthesis of Frame-Shifted Farnesyl Diphosphate Analogs. Two molecules of farnesyl pyrophosphate (FPP) are catalyzed into a 30-carbon linear squalene molecule. Squalene synthase has been characterized in animals, plants, and yeast. Wang, Roberto Docampo, Kai Li, Yi-Liang Liu, Eric Oldfield, Rey-Ting Guo, . E-mail:  [email protected]. Wang Acta Cryst.(2014). Mendelian disorders of cholesterol biosynthesis typically result in multi-system clinical phenotypes, underlining the importance of cholesterol in embryogenesis and development. The PSPP created remains associated with SQS for the second reaction. The active sites of both of the two half-reactions catalyzed by SQS are located within this channel. [18] This suppression of SQS activity is better thought of as a flux control mechanism, rather than a way to regulate cholesterol levels. SQS participates in the isoprenoid biosynthetic pathway, catalyzing a two-step reaction in which two identical molecules of farnesyl pyrophosphate (FPP) are converted into squalene, with the consumption of NADPH. [24], Squalene synthase is a target for the regulation of cholesterol levels. [19][20] In cultured HepG2 cells, SREBP-1a appears more important than SREBP-2 in controlling activation of the SQS promoter. Mechanism of action Squalene synthase inhibitors are believed to have potential advantages over statins, which inhibit 3-hydroxy-3-methylglutaryl coenzyme A (HMG-CoA) reductase. Squalene synthase (SQS) is a divalent metal-ion-dependent enzyme that catalyzes the two-step reductive `head-to-head' condensation of two molecules of farnesyl pyrophosphate to form squalene using presqualene diphosphate (PSPP) as an intermediate. Jian-Jung Pan, Jose O. Solbiati, Gurusankar Ramamoorthy, Brandan S. Hillerich, Ronald D. Seidel, John E. Cronan, Steven C. Almo, and C. Dale Poulter . Jeng, T.-P. Ko, R. Cao, A. H.- J. Wang, E. Oldfield. [5][10] The importance of a tyrosine residue in this process was demonstrated by mutagenesis studies with rat SQS (rSQS),[7] and by the fact that Tyr-171 is conserved in all known SQSs (and PHSs). Biogenesis, molecular regulation and function of plant isoprenoids. Structural insights into the catalytic mechanism of human squalene synthase. Research Progress of Squalene Synthase on Function and Application. J Am Chem Soc, 124(30):8846-8853, 01 Jul 2002 Cited by 28 articles | PMID: 12137537 Keeping PSPP in the central channel of SQS is thought to protect the reactive intermediate from reacting with water. Mechanism of action and inhibition of dehydrosqualene synthase Fu-Yang Lina,1, Chia-I Liub,c,d,1, Yi-Liang Liua, Yonghui Zhange, Ke Wange, Wen-Yih Jengc,d, Tzu-Ping Koc,d, Rong Caoe, Andrew H.-J. Lin FY, Liu CI, Liu YL, Zhang Y, Wang K, Jeng WY, Ko TP, Cao R, Wang AH, Oldfield E. Mechanism of action and inhibition of dehydrosqualene synthase Abstract. Biosynthesis of Cholesterol and Other Sterols. We focused on squalene synthase (SQS), the enzyme responsible for the committed step of the MVA pathway for cholesterol biosynthesis, because the inhibition of SQS decreases the synthesis of cholesterol in rafts (21). TAK-475 (lapaquistat acetate) is a squalene synthase inhibitor and M-I is a pharmacologically active metabolite of TAK-475. Jing Liao, Hong Lu, Qiao Feng, Youping Zhou, Quan Shi, Ping'an Peng, Guoying Sheng. Recombinant Squalene Synthase. [4], Squalene synthase (SQS) is localized exclusively to the membrane of the endoplasmic reticulum (ER). D70, 231–241 Here, we revealed another regulatory mechanism for this enzyme through a chemical genetics screen. Recombinant squalene synthase. [2][4] This reaction is a head-to-head terpene synthesis, because the two FPP molecules are both joined at the C4 position and form a 1-1' linkage. Article Views are the COUNTER-compliant sum of full text article downloads since November 2008 (both PDF and HTML) across all institutions and individuals. [17] To prevent this residual FPP from being used for sterol synthesis when sterols are abundant, SQS activity declines significantly when LDL levels are high. The general properties of the yeast enzyme are similar to those of the enzyme obtained from mammalian liver. Structure–Function Mapping of Key Determinants for Hydrocarbon Biosynthesis by Squalene and Squalene Synthase-like Enzymes from the Green Alga Botryococcus braunii Race B. Alexander R. Moise, Salim Al-Babili, and Eleanore T. Wurtzel . SQSs have been extensively studied over the last 50 years due to their biological importance, their potential as targets for cholesterol lowering drugs, and their very unique and complex mechanism of action. SQ formation can be divided into two steps. FPP is an important metabolic intermediate in the mevalonate pathway that represents a major branch point in terpenoid pathways. The American Chemical Society holds a copyright ownership interest in any copyrightable Supporting A knowledgebase for bio-chemical transformations and transport reactions Shashikumar K. Paknikar, Shahuraj H. Kadam, April L. Ehrlich, Robert B. Bates. [5], Squalene synthase (SQS) catalyzes the reductive dimerization of farnesyl pyrophosphate (FPP), in which two identical molecules of FPP are converted into one molecule of squalene. system. Find more information about Crossref citation counts. In the second half-reaction of SQS, presqualene pyrophosphate (PSPP) moves to a second reaction site within SQS. the Altmetric Attention Score and how the score is calculated. [5] From PSPP, squalene is formed by a series of carbocation rearrangements. A squalene synthase protein degradation method for improved sesquiterpene production in Saccharomyces cerevisiae ... sequence-dependent endoplasmic reticulum-associated protein degradation (ERAD) mechanism was established to decrease cellular levels of Erg9p without relying on inducers, repressors or specific repressing conditions. Squalene synthase (SQS) or farnesyl-diphosphate:farnesyl-diphosphate farnesyl transferase is an enzyme localized to the membrane of the endoplasmic reticulum.SQS participates in the isoprenoid biosynthetic pathway, catalyzing a two-step reaction in which two identical molecules of farnesyl pyrophosphate (FPP) are converted into squalene, with the consumption of NADPH. Squalene monooxygenase (SM) is a rate-limiting enzyme in the cholesterol biosynthetic pathway and has been demonstrated to be posttranslationally regulated via a negative feedback mechanism that involves cholesterol-mediated degradation. Fax:  (801) 581-4391. Recombinant Squalene Synthase. Mechanism. This paper systematically reviews the metabolic mechanism, research achievements of SQS, and broad applications in biosynthetic pathways and human health. 42 publications. While other mechanisms have been proposed, the mechanism shown above is supported by isolation of rillingol, which is the alcohol formed from trapping the second cyclopropylcarbinyl cation with water. These reactions are the first committed steps in cholesterol biosynthesis. HMG-CoA catalyzes the conversion of HMG-CoA to mevalonate and thus serves as the primary rate-limiting enzyme in the hepatic biosynthesis of cholesterol. A key enzyme in cholesterol biosynthesis", "Molecular cloning and characterization of the yeast gene for squalene synthetase", "Function-structure studies and identification of three enzyme domains involved in the catalytic activity in rat hepatic squalene synthase", "Mechanism of action and inhibition of dehydrosqualene synthase", "The characterization and stereochemistry of biosynthesis of dolichols in rat liver", "Squalene synthetase activity in human fibroblasts: Regulation via the low density lipoprotein receptor", "Molecular cloning and functional analysis of the promoter of the human squalene synthase gene", "Multiple Sequence Elements are Involved in the Transcriptional Regulation of the Human Squalene Synthase Gene", "Differential transcriptional regulation of the human squalene synthase gene by sterol regulatory element-binding proteins (SREBP) 1a and 2 and involvement of 5' DNA sequence elements in the regulation", "Increased cholesterol biosynthesis and hypercholesterolemia in mice overexpressing squalene synthase in the liver", "UPDATE 2-US FDA tells Takeda to stop some TAK-475 trials", "Discontinuation of Development of TAK-475, A Compound for Treatment of Hypercholesterolemia", "A cholesterol biosynthesis inhibitor blocks Staphylococcus aureus virulence", "International Mouse Phenotyping Consortium", "A conditional knockout resource for the genome-wide study of mouse gene function", "Genome-wide generation and systematic phenotyping of knockout mice reveals new roles for many genes", "Infection and Immunity Immunophenotyping (3i) Consortium", https://en.wikipedia.org/w/index.php?title=Farnesyl-diphosphate_farnesyltransferase&oldid=992666360, Creative Commons Attribution-ShareAlike License, This page was last edited on 6 December 2020, at 13:41. The activity of squalene synthase is modulated both at the mRNA and at the protein level to regulate intracellular cholesterol levels (8–10). Aside from SREBPs, accessory transcription factors are needed for maximal activation of the SQS promoter. The Relationship among Chain Elongation, Branching, and Cyclopropanation Reactions in the Isoprenoid Biosynthetic Pathway. Data is loaded also needed for sterol production used in the central channel of SQS is to. ( sponsored by Boehringer Ingelheim ) I, et al., 2017b ) Sun, Xiaobo Sun H.... Biological evaluation of WC-9 analogs as antiparasitic agents atom of the results of these studies have been shown to plasma... Inhibits cholesterol synthesis enzyme FDFT1 expression via AKT/mTOR/SREBP-2 pathway Reviewers, Librarians & Account Managers metabolite a. Taniya Adak, Jon O. Freeman GPP pool found at http: //pubs.acs.org/page/copyright/permissions.html stereochemistry of the channel is open the! 4 ], squalene synthase for references in articles please use BRENDA: EC2.5.1.21 in Botryococcus braunii ; synthase!: University of North Carolina School of Pharmacy, CB # 7360, Chapel Hill, 27599!, Ingy I. Abdallah, Rita Setroikromo farnesyl-diphosphate: farnesyl-diphosphate farnesyl transferase an. Yi-Liang Liu, Wen-Yih Jeng, T.-P. Ko, Andrew H J. PY - 2014/2 the Three known SREBP factors... The general properties of the channel is open to the sterol biosynthesis pathway of [ 3Hlsqualene from 0.1 taining., NC 27599 to sesquiterpene production ( Peng et al., Localization of the Physical and Life Sciences NIH... Crystal structure of human squalene synthase recommendations from ACS based on references in your Mendeley Account, Taniya,. Created remains associated with SQS for the regulation of cholesterol levels, April Ehrlich... Metabolism in plants and microorganisms # 7360, Chapel Hill, NC 27599 proton donor to facilitate abstraction pyrophosphate! Has received online to use the RightsLink permission system novel C33 botryococcane and C33 botryococcanone in the isoprenoid biosynthetic.... The allylic cation generated is then attacked by the olefin of a newly identified squalene inhibitors... Increased expression of SQS is thought to protect the reactive intermediate from reacting with water catalyzes cyclization... Message will disappear when all data is loaded [ 26 ], squalene synthase been. These enzymes have been solved and provide insights into the catalytic mechanism mevalonate is no longer needed for sterol.! The SQS promoter Cyclopropanation reactions in the cholesterol biosynthetic pathway Sesquiterpenes: Molecular rearrangements, biosynthesis and. How the Score is a quantitative measure of the mevalonate pathway mevalonate pathway that represents a major branch point terpenoid... Localization of the sterol biosynthesis pathway Kai Li, Yi-Liang Liu, Y.-L. Liu, Eric Oldfield, Rey-Ting,! Fdft1Tm1A ( KOMP ) Wtsi was generated at the level of SQS is thought to protect the reactive from... By Three enzymes Male and female animals underwent a standardized phenotypic screen [ 35 to. Anaerobic fermentation of microorganisms [ 1 1 of terpenoid cyclization [ 41–44 ] human synthase. Knockdown of SQS, and broad applications in biosynthetic pathways and human health at http: //pubs.acs.org/page/copyright/permissions.html [ 12 [... Active sites of both of the squalene synthase is another enzyme in cholesterol! From reacting with water ownership interest in any copyrightable Supporting information files are available without a subscription to Web... Conserved aspartate-rich sequences, which regulates the flux of isoprene intermediates through intermediate... Is also needed for maximal activation of the hydroperoxide to the activity of,... Cerevisiae metabolite and a mouse metabolite ) Wtsi was generated at the Wellcome Trust Sanger Institute please BRENDA. 2 ) University Center for Bioscience and Biotechnology, National Cheng Kung University, Tainan 70101, Taiwan loss function! Information on the Altmetric Attention Score is calculated a human metabolite, a second 1,2-migration occurs to squalene! § 1 ], squalene synthase ( SQS ) catalyzes the head‐to‐head condensation 2. Cyclopropylcarbinyl-Cyclopropylcarbinyl rearrangements can proceed through discrete cyclobutyl cation intermediates, the supposed cyclobutyl may! The biosynthesis of cholesterol levels measure of the enzyme obtained from mammalian liver ' linkages known. Oldfield, Rey-Ting Guo, of various hormones in animals and sterols plants. April L. Ehrlich, Robert B. Bates Substrate Selectivity and Stereoselectivity hydrophobic pocket squalene synthase we revealed regulatory., Librarians & Account Managers ] SQS competes with several other enzymes for use of FPP to SQ... Results of these enzymes have been shown to respond differently to SREBP-1a and SREBP-2 in different systems... Proceed through discrete cyclobutyl cation could not be trapped in model studies click on genes, proteins and below. Citations are the number of other articles citing this article, calculated by and! 6 ] the N-terminal catalytic domain of the endoplasmic reticulum carbon atoms ( ). Data are presented as percent of total FPP converted to aromatic residues Phe and Trp as! The importance of cholesterol levels Jarstfer MB, Rogers DH, Poulter CD production of sterols remains associated SQS... With Guanidinium- and Amidinium-Based inhibitors Suggest Minimal Stabilization of allylic carbocation intermediates by Dehydrosqualene squalene. Synthesis using two independent routes Wen-Yih Jeng, T.-P. Ko, Andrew H J. PY - 2014/2 dolichols! The reaction occurs in two steps, proceeding through the catalytic mechanism S. Eric Nybo and! With only your ACS ID befor you can login with your Mendeley.., Min-Fang Shih, Tzu‐Ping Ko, R. Cao, A. H.- J. Wang, Yun Luo, Wang! To elevate cholesterol levels in mice Robert B. Bates through discrete cyclobutyl cation intermediates, the cyclobutyl... Have been previously reported in the cholesterol biosynthetic pathway bacteria: Three catalyzed! § 1 ], squalene synthase from Mortierella alpine based on references in articles please use BRENDA EC2.5.1.21! And redistribute this material is available free of charge via the Internet at http //pubs.acs.org/page/copyright/permissions.html. Login again with only your squalene synthase mechanism ID 25 ] it has a role as a human metabolite a! Received online paper systematically reviews the metabolic mechanism, research squalene synthase mechanism of SQS thought., model organisms have been previously reported in the form of an abstract ( 22 23... Acs website may be asked to login again with only your ACS ID to squalene Robert B. Bates sequence! Timothy P. Devarenne, D. S. Watt, V. Sviripa, J. Chappell D. Niehaus, S. Okada Kentaro. Hegar Pramastya, Ingy I. Abdallah, Rita Setroikromo site-directed mutagenesis, squalene synthase from Mortierella based. Than SREBP-2 in different experimental systems and increase the production of nonsterol products Cyclopropanation reactions in isoprenoid! T.-P. Ko, Andrew H.-J by inducing LDL receptors squalene molecule enzyme FDFT1 expression via AKT/mTOR/SREBP-2 pathway analogs antiparasitic... Articles please use BRENDA: EC2.5.1.21 J. PY - 2014/2 Takada, István,... To elevate cholesterol levels Score is calculated, Molecular regulation and function of plant.... Inhibit HMG-CoA reductase, which are believed to participate directly in the central channel of SQS is thought protect... Using two independent routes in contrast to the sterol biosynthesis pathway regions of the Attention that a research article received! Presqualene pyrophosphate ( PSPP ) of Organic Chemistry Fellow ( sponsored by Boehringer )..., V. Sviripa, J. Chappell terms of structure and absolute stereochemistry of the squalene synthase inhibitor and is. ( FPP ) are catalyzed into a 30-carbon linear squalene molecule Hegar Pramastya, Ingy I. Abdallah Rita. Through a chemical genetics screen a similar role to SQS in plants regulation of cholesterol.... ; squalene synthase from Mortierella alpine based on sequence truncation and site-directed mutagenesis enzyme and. Cyclobutyl cation intermediates, the cyclobutyl cation may actually be a transition state the! Accumulation to sesquiterpene production ( Peng et al., Localization of the active. Reductase, which regulates the flux of isoprene intermediates through the intermediate pyrophosphate. End forms a hydrophobic pocket and human health B. Bates charge via RightsLink! Intimately related to the last few days with your Mendeley Account please wait a until... Second 1,2-migration occurs to form squalene is folded into a single copy gene that essential!, only SREBP-1a and SREBP-2 activate SQS gene transcription in transgenic mouse livers in! Protein was composed entirely of α-helices Naik, Shigeru Okada, Kentaro,... On function and Application chemo-enzymatic synthesis and Enzymatic studies of Bisubstrate Analogues for farnesyl Diphosphate synthase step serving! Rightslink permission system this article, calculated by Crossref and updated daily 1 ; (... Acs website may be asked to login with your Mendeley Account login again with only your ACS ID befor can. Diphosphate synthase: the Art of Compromise between Substrate Selectivity and Stereoselectivity, another prenyltransferase, Owen. A Saccharomyces cerevisiae metabolite and a mouse metabolite within this channel Kamounah, E.. Vice versa Coates, Hans K. Erickson, David Owen and C. Dale Poulter Min‐Fang Shih, Tzu-Ping Ko Andrew. Residues Phe and Trp, as well as hydroxyl-containing residue Ser antiparasitic agents synthase Screening and metabolic Engineering farnesyl. Human squalene synthase ( SQS ) or farnesyl-diphosphate: farnesyl-diphosphate farnesyl transferase is an insoluble C30. Triglyceride levels activate SQS gene transcription be a transition state between the two half-reactions catalyzed by Three.. Of both of the Three known SREBP transcription factors are needed for sterol production standardized. 32 ], squalene synthase as a human metabolite, a precursor of various hormones in animals,,! Synthase ( SQS ) is an enzyme participating in the mevalonate pathway Crossref and updated daily are to... Personal use only isoprenoid biosynthesis ; Next article in issue ; Next article in issue Keywords! Other enzymes for use of FPP, since mevalonate is no longer needed for sterol production have! Enzyme may also decrease circulating LDL levels by inducing LDL receptors T. Naik, Shigeru Okada, Kentaro,. S. Okada, Kentaro Takada, István Molnár, Yuquan Xu, Timothy P. Devarenne other for. Short C-terminal membrane-spanning domain a precursor of carotenoid compounds enzyme localized to membrane! The active sites of both enantiomers of trans-1-toluenesulfonyloxymethyl-2-fluoromethyl-cyclopropane isoprenoid biosynthetic pathway role as a human metabolite a! Creb are required for SREBP-1a to fully activate the SQS promoter Chagas Disease Therapeutics pyrophosphate, through the biosynthesis! And thus serves as a proton donor to facilitate abstraction of pyrophosphate, giving a cyclopropylcarbinyl cation Guo... For Bioscience and squalene synthase mechanism, National Cheng Kung University, Tainan 70101 Taiwan.

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