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We have investigatedthe kinetic mechanism of phosPE' N-methyltransferase is an enzyme of the endoplasmic phatidylethanolamine PE ; N-methyltransferase pu- reticulum that is responsible for the synthesis of 20-30% of rified from rat liver using PE, phosphatidyl-N-mono- hepatic PC 1 ; .The kinetic mechanism for PE methylation methylethanolamine PMME ; , and phosphatidyl-N, N- appears to be a complex pathway involving three separate dimethylethanolamine PDME ; as substrates. We pre- methylation events. Until recently, the number of enzymes viously reported Ridgway, N. D., and Vance, D. E. involved in theprocess has been in doubt. The purification of 1987 ; Biol. Chem. 262, J. 17231-17239 ; initial a low molecular weight methyltransferase, and the finding that velocity curves with PE, PMME, and PDME at a fixed that it catalyzes the complete conversion of PE to concentration of Triton X-100 were sigmoidal, thus casts doubt on reports that two enzymes convert PE to PC generating nonlinear inverse plots. Comparison with liver 3, 4 ; . With the availability of purified PE N-methylother integral membrane enzymes suggested this re- transferase, a detailed kinetic mechanism can now be consponse resulted from the enzyme's requirement for a structed based on substrate initial velocity and product inhicomplete boundary layer of phospholipid. Hence, the bition patterns according to the method of Cleland 5 ; . An effect of a nonsubstrate phospholipid on initial velocity interesting feature of PE N-methyltransferase kinetics is the patterns for PE, PMME, and PDME was examined. apparent cooperative effect of PE, PMME, and PDMEwhen The sigmoidicity of initial velocity curves at constant their surface concentration in a Triton X-100 mixed micelle and PE, Triton X-100 concentration increasing was varied 2 ; . Such apparent cooperativity is a common PMME, and PDME were converted to the more famil- feature of catalysis in mixed micelles for both integral and iar hyperbolic response by the addition of egg phos- extrinsic membrane proteins 6, 7 ; . However, it is not coopphatidylcholine PC ; . Hill coefficients for PE, PMME, erativity in the true sense, but is more readily explained by and PDME at a fixed Triton concentration were 3 6 an effect on activity due to noncooperative binding of lipid to ., 2.5, and 4.7, respectively, but with the addition of 30 portions of integral membrane proteins in contact with the or 40 mol 70of egg PC, coefficients were close to unity micelle or phospholipid membrane 8 ; - The activation by egg of PE, PMME, and 0912. PC In this report we provide kinetic evidence that the 18.3PDME methylation indicates that a secondary phos- kDa hepatic PE N-methyltransferase converts PE to PCby pholipid binding site s ; plays a role in catalysis in a concerted mechanism. In addition, the influence of micelle mixed micelles. This site s ; may representa transmem- concentration on enzyme activity and initial velocity curves brane segment s ; in close association with a boundary at fixed substrate and increasing Triton X-100 were further investigated. The observed lack of saturable binding by PE layer of phospholipid. Kinetic analysisof initial velocity and product inhi- N-methyltransferase to mixed micelles is a furtherindication bition patterns for PMME and PDME methylation fit of its integral nature. Effects of a nonsubstrate phospholipid an ordered Bi Bi mechanism. Phospholipid substrates egg PC ; on apparent cooperativity exerted by the lipid suband products were the first to bind and the last to strates was analyzed and interpreted in terms of a boundary dissociate from the active site, respectively. As well, layer of phospholipid necessary for full catalytic activity. PE, PMME, and PDME compete for a single active site. The overall kinetic scheme for the methylation of PE EXPERIMENTALPROCEDURES to PC in mixed micelles involves the initial binding of Materials-PMME and PDMEwere purchased from Avanti Polar PE, followed by successive steps where S-adenosyl-L- Lipids, Birmingham, AL. Egg PC, AdoMet, AdoHcy, Triton X-100, methionine is bound, the sulfonium methyl group is and DTT were from Sigma. [methyl-3H]AdoMetand [l-3H]ethanoltransferred, and S-adenosyl-L-homocysteine is re- amine were purchased from Amersham Corp. Silica Gel 60 thin-layer plates were from Merck. All other materials were of reagent grade. leased and revlimid.
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References Abramovitz, Moises, "Resource and Output Trends in the United States since 1870, " American Economic Review Papers and Proceedings, 46, 5-23, 1956. Arrow, Kenneth J., "Economic Welfare and the Allocation of Resources for Inventions." In R. Nelson ed. ; , The Rate and Direction of Inventive Activity, Princeton University Press, 609-25, 1962. Avnimelech, Gil and Morris Teubal, "Evolutionary Innovation and High Tech Policy: What can we learn from Israel's targeting of venture capital?" The Samuel Neaman Institute STE Working Paper Series, 25, 2005. Blomstrom, M. and Kokko, A., "How Foreign Investment Affects Host Countries", Policy Research Working Paper, No. 1745, The World Bank, 1999. Branscomb, Lewis M. and Philip E. Auerswald, "Between Invention and Innovation, an Analysis of Funding for Early-Stage Technology Development." The Advanced Technology Program, National Institute of Standards and Technology, November 2002. Bresnahan, T. and Trajtenberg, M., "General Purpose Technologies - Engines of Growth?" Journal of Econometrics, 65 1 ; , pp. 83-108, 1995. Coe, David T and Helpman, Elhanan. "International R&D Spillovers." European Economic Review. Vol. 39 5 ; . 859-87. May 1995. Coe, David T., Helpman, E. and Hoffmaister, A.W., "North-South R&D Spillovers", Economic Journal, Vol.107, No.440, pp. 134-49, 1997. Cohen, Wesley M. and Levinthal, Daniel A, "Innovation and Learning: The Two Faces of R&D." The Economic Journal, 99, pp. 569-96, 1989. Comin, Diego and Bart Hobijn, "Cross-Country Technological Adoption: New Evidence and Questions." Journal of Monetary Economics, forthcoming 2005. Griliches, Zvi, "An Exploration in the Economics of Technical Change." Econometrica, 25 4 ; , 1957. Griliches, Zvi ed. ; , R&D, Patents, and Productivity, University of Chicago Press, 1984. Griliches, Zvi, Technology, Education, and Productivity, New York: Basil Blackwell, 1988. Grossman, Gene M. and Helpman, Elhanan, Innovation and Growth in the Global Economy. Cambridge, MA: The MIT Press, 1991 and reyataz.
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