Bacterial Long Chain Fatty Acid Transport: Gateway to a Fatty Acid-responsive Signaling System

Exogenous long chain fatty acids (LCFAs) 1The abbreviations used are: LCFA, long chain fatty acid; ACSL, acyl-CoA synthetase; LCA-CoA, long chain acyl-CoA; LDAO, lauryldimethylamine N-oxide.1The abbreviations used are: LCFA, long chain fatty acid; ACSL, acyl-CoA synthetase; LCA-CoA, long chain acyl-CoA; LDAO, lauryldimethylamine N-oxide. influence a myriad of cellular processes including intracellular signaling and patterns of gene expression. Some cell types accumulate distinct classes of LCFAs whereas others accumulate or release fatty acids following some type of external stimulus (i.e. hormonal or nutritional cues). Given the energetic cost of fatty acid synthesis on one hand and (at least in the mammalian system) the necessity for obtaining essential fatty acids from the environment, the transport of LCFAs must necessarily represent a fundamental biological process. The biochemical mechanisms underpinning fatty acid transport involve specific proteins and enzymes, which act either directly at the membrane or indirectly at the level of downstream metabolism. Distinct membrane-bound and membrane-associated fatty acid transport proteins have been identified and characterized in a number of different systems including bacteria, yeast, and mammals (1Abumrad N. Harmon C. Ibrahimi A. J. Lipid Res. 1998; 39: 2309-2318Abstract Full Text Full Text PDF PubMed Google Scholar, 2Abumrad N. Coburn C. Ibrahimi A. Biochim. Biophys. Acta. 1999; 1441: 4-13Crossref PubMed Scopus (208) Google Scholar, 3Bernlohr D.A. Coe N.R. LiCata V.J. Semin. Cell Dev. Biol. 1999; 10: 43-49Crossref PubMed Scopus (40) Google Scholar, 4Black P.N. J. Bacteriol. 1988; 170: 2850-2854Crossref PubMed Google Scholar, 5Black P.N. DiRusso C.C. Microbiol. Mol. Biol. Rev. 2003; 67: 454-472Crossref PubMed Scopus (176) Google Scholar, 6Hirsch D. Stahl A. Lodish H.F. Proc. Natl. Acad. Sci. U. S. A. 1998; 95: 8625-8629Crossref PubMed Scopus (366) Google Scholar, 7Schaffer J.E. Lodish H.F. Cell. 1994; 79: 427-436Abstract Full Text PDF PubMed Scopus (736) Google Scholar).The genetic and biochemical foundations of LCFA transport were first elucidated in the Gram-negative bacterium Escherichia coli. The hallmark of this system is the coupled transport and activation of exogenous LCFAs, which in turn leads to changes in transcription patterns of the genes encoding the proteins required for fatty acid biosynthesis and degradation. The three central components of this system include FadL, an outer membrane-bound fatty acid transport protein, FadD, an inner membrane-associated long chain acyl-CoA synthetase (ACSL), and FadR, a long chain acyl-CoA-responsive transcription factor. The bacterial fatty acid transport and trafficking system leading to downstream fatty acid-responsive transcriptional regulation serves as a fundamental paradigm in biology, which provides a number of guiding principles that can be applied to more complex systems.Genetic Foundations of Fatty Acid Transport and Trafficking System in E. coliThe transport of exogenous LCFAs across the cell envelope of E. coli requires both the transport the and the the The of long chain is the that the of the transcription the which in turn the of genes in fatty acid and biosynthesis in C.C. Mol. Microbiol. 1998; PubMed Scopus Google and C.C. P.N. Lipid Res. 1999; PubMed Scopus Google The in the of and the required to the genetic foundations of the fatty acid transport and trafficking system in E. coli Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google Scholar, Scholar, J. Bacteriol. PubMed Google that identified and of the genes of the fatty acid and distinct including Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google J. PubMed Scopus Google identified the gene in and at that this a of the fatty acid transport J. PubMed Scopus Google characterized the system for the of exogenous fatty acids and that this in the of fatty acid transport an for the transport system in E. is the membrane is to fatty acids from the membrane activation to J. PubMed Scopus Google a and D. Mol. PubMed Scopus Google identified a that type acyl-CoA synthetase to exogenous fatty acids that the gene in the of LCFAs and this a protein, the of the D. Mol. PubMed Scopus Google and Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google identified the gene in the of a in the transport of exogenous the of the gene an inner membrane-bound fatty acid that the transport of exogenous LCFAs and in Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google Scholar, J. Biol. Full Text PDF PubMed Google Scholar, J. Biol. Full Text PDF PubMed Google and of P.N. J. Biol. Full Text PDF PubMed Google that this in the outer membrane and to for The of in the outer membrane and in the transport of exogenous fatty acids the in the P.N. J. Bacteriol. 1988; 170: 2850-2854Crossref PubMed Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. Biochim. Biophys. Acta. PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google Scholar, P.N. DiRusso C.C. J. PubMed Google Scholar, C.C. P.N. Mol. Cell. 1999; PubMed Google Scholar).The the of the and genes is the long chain acyl-CoA transcription C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, C.C. J. Biol. Full Text PDF PubMed Google the of fatty acid transport and is LCA-CoA, this system to E. coli to transcriptional in the of exogenous long chain fatty DiRusso and C.C. Mol. Microbiol. 1998; PubMed Scopus Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google a of the regulation of the fatty acid and genes in E. and Fatty Acid Transport outer membrane is the membrane-bound LCFA fatty acid transport proteins have been identified in of including in protein, membrane-bound D. Rev. PubMed Scopus Google as a for acid (1Abumrad N. Harmon C. Ibrahimi A. J. Lipid Res. 1998; 39: 2309-2318Abstract Full Text Full Text PDF PubMed Google Scholar, 2Abumrad N. Coburn C. Ibrahimi A. Biochim. Biophys. Acta. 1999; 1441: 4-13Crossref PubMed Scopus (208) Google and long chain fatty acids acid transport D. Stahl A. Lodish H.F. Proc. Natl. Acad. Sci. U. S. A. 1998; 95: 8625-8629Crossref PubMed Scopus (366) Google Scholar, N.R. D.A. J. Biol. 1999; Full Text Full Text PDF PubMed Scopus Google Scholar, D.A. PubMed Google The on the long chain fatty acid transport from the fundamental to of distinct which the of a as of the transport the outer membrane of the bacterial cell envelope the acid to exogenous LCFAs a The of long chain fatty acids to provides that the of the provides P.N. Biochim. Biophys. Acta. PubMed Scopus Google and to for of that the of the fatty acid is required P.N. Biochim. Biophys. Acta. PubMed Scopus Google the that specific the and the chain of the fatty the of the LCFA is essential for acid to as this in of the gene the LCFA and transport P.N. J. PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google of the of and that transport is to type whereas is type that the of in in acid that more for both LCFA and that in acid the of required for transport whereas the required for P.N. J. Biol. Full Text PDF PubMed Google of and in identified three acid and that to transport and one that to P.N. J. 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Bacteriol. 1994; PubMed Google the of the the including the is which is to the number of fatty and 2003; PubMed Scopus Google have fatty acids to and the membrane the of the the of fatty acid that and to the of the The of the which is to the number of fatty acids and the of fatty acid transport A. D. DiRusso C.C. P.N. Biophys. 1999; PubMed Scopus Google Acid Transport across and Google the of an inner membrane-bound that to a in the transport of exogenous fatty acids across the inner and A. P.N. J. Bacteriol. 1994; PubMed Google to a be classes were identified that on LCFA transport and both which to the that the of the fatty acid transport in the inner membrane the J. PubMed Scopus Google characterized in the of LCFA transport and that this this to and exogenous LCFAs transport across the inner and required for fatty acid which the for for A. D. DiRusso C.C. P.N. Biophys. 1999; PubMed Scopus Google Scholar, DiRusso C.C. P.N. J. Biol. Full Text Full Text PDF PubMed Scopus Google number of the the E. coli this both the inner membrane and different cellular the of been that the inner membrane in an to and LCFAs, the of fatty acid transport DiRusso C.C. P.N. J. Biol. Full Text Full Text PDF PubMed Scopus Google as a to the of acyl-CoA DiRusso C.C. P.N. J. Biol. Full Text Full Text PDF PubMed Scopus Google Scholar, P.N. DiRusso C.C. J. Biol. Full Text Full Text PDF PubMed Scopus Google is a acid of which is to P.N. DiRusso C.C. J. Biol. Full Text Full Text PDF PubMed Scopus Google of the gene to this the of this to of that the for fatty acids the fatty acid chain this of the is essential for in to fatty acid chain and is to of the fatty acid P.N. DiRusso C.C. J. Biol. Full Text Full Text PDF PubMed Scopus Google the LCFA the fatty acid and to the this is of the fatty acid P.N. DiRusso C.C. D. J. J. Biol. Full Text Full Text PDF PubMed Scopus Google of in to the a to DiRusso C.C. P.N. J. Biol. 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Full Text Full Text PDF PubMed Scopus Google The that of is that the this to fatty acid the of the and release of the the release of and the of which in turn is from the DiRusso C.C. P.N. J. Biol. Full Text Full Text PDF PubMed Scopus Google Scholar, N. S. S. J. Biol. Full Text Full Text PDF PubMed Scopus Google Fatty Acid Transport and as a of the the serves as a for can be membrane of is that as a that the of the transcription a of from the DiRusso as the that the of this transcription C.C. J. Biol. Full Text PDF PubMed Google Scholar, DiRusso C.C. J. J. PubMed Scopus Google Scholar, DiRusso C.C. J. J. PubMed Scopus Google Scholar, C.C. Mol. Microbiol. PubMed Scopus Google of The first the of for different of chain C.C. J. Biol. Full Text PDF PubMed Google Scholar, C.C. J. J. Biol. 1998; Full Text Full Text PDF PubMed Scopus Google the the following the of in N. DiRusso C.C. J. Biol. Full Text Full Text PDF PubMed Scopus Google and the three of and DiRusso C.C. J. J. PubMed Scopus Google Scholar, DiRusso C.C. J. J. PubMed Scopus Google Scholar).The signaling is of as the of a number of genes in fatty acid fatty acid and the C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, C.C. Mol. Microbiol. PubMed Scopus Google is a transcriptional of genes in fatty acid and and a transcriptional of at least genes required for fatty acid biosynthesis and C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, C.C. Mol. Microbiol. PubMed Scopus Google as an of encoding a transcriptional of the genes and as a of which in a number of C.C. Mol. Microbiol. 1998; PubMed Scopus Google Scholar, A. DiRusso C.C. J. Bacteriol. PubMed Google for have been and C.C. J. Biol. Full Text PDF PubMed Google Scholar, C.C. Mol. Microbiol. PubMed Scopus Google Scholar, A. DiRusso C.C. J. Bacteriol. PubMed Google Scholar, P.N. DiRusso C.C. J. Biol. Full Text PDF PubMed Google the of which is required for bacterial on The of the of the fatty acid to the is the that encoding acyl-CoA is in to of DiRusso C.C. 1999; PubMed Scopus Google to the intracellular of to the of genes in fatty acid and and the the to FadR, to The is the transcriptional of the fatty acid genes and of as as or types of the can to fatty acid genes and Acid and in E. coliThe mechanisms that the transport and trafficking of exogenous LCFAs in E. coli. leads to changes in the transcriptional of genes in both fatty acid and fatty acid this is a bacterial to including and serves as a of the of exogenous fatty acids downstream transcriptional of this system and the bacterial cell to the of exogenous both and at the cell and the intracellular of which to to on the fatty acid genes as a and fatty acid genes as an the cell LCFAs in the environment, to and a the the The more of the the of fatty acid which and across the The to long chain fatty acids from the membrane in a that is coupled to the of the to a of the which transcriptional of the fatty acid genes and and the transcriptional activation of the fatty acid genes and is to the changes in intracellular which the bacterial cell to Exogenous long chain fatty acids (LCFAs) 1The abbreviations used are: LCFA, long chain fatty acid; ACSL, acyl-CoA synthetase; LCA-CoA, long chain acyl-CoA; LDAO, lauryldimethylamine N-oxide.1The abbreviations used are: LCFA, long chain fatty acid; ACSL, acyl-CoA synthetase; LCA-CoA, long chain acyl-CoA; LDAO, lauryldimethylamine N-oxide. influence a myriad of cellular processes including intracellular signaling and patterns of gene expression. Some cell types accumulate distinct classes of LCFAs whereas others accumulate or release fatty acids following some type of external stimulus (i.e. hormonal or nutritional cues). Given the energetic cost of fatty acid synthesis on one hand and (at least in the mammalian system) the necessity for obtaining essential fatty acids from the environment, the transport of LCFAs must necessarily represent a fundamental biological process. The biochemical mechanisms underpinning fatty acid transport involve specific proteins and enzymes, which act either directly at the membrane or indirectly at the level of downstream metabolism. Distinct membrane-bound and membrane-associated fatty acid transport proteins have been identified and characterized in a number of different systems including bacteria, yeast, and mammals (1Abumrad N. Harmon C. Ibrahimi A. J. Lipid Res. 1998; 39: 2309-2318Abstract Full Text Full Text PDF PubMed Google Scholar, 2Abumrad N. Coburn C. Ibrahimi A. Biochim. Biophys. Acta. 1999; 1441: 4-13Crossref PubMed Scopus (208) Google Scholar, 3Bernlohr D.A. Coe N.R. LiCata V.J. Semin. Cell Dev. Biol. 1999; 10: 43-49Crossref PubMed Scopus (40) Google Scholar, 4Black P.N. J. Bacteriol. 1988; 170: 2850-2854Crossref PubMed Google Scholar, 5Black P.N. DiRusso C.C. Microbiol. Mol. Biol. Rev. 2003; 67: 454-472Crossref PubMed Scopus (176) Google Scholar, 6Hirsch D. Stahl A. Lodish H.F. Proc. Natl. Acad. Sci. U. S. A. 1998; 95: 8625-8629Crossref PubMed Scopus (366) Google Scholar, 7Schaffer J.E. Lodish H.F. Cell. 1994; 79: 427-436Abstract Full Text PDF PubMed Scopus (736) Google The genetic and biochemical foundations of LCFA transport were first elucidated in the Gram-negative bacterium Escherichia coli. The hallmark of this system is the coupled transport and activation of exogenous LCFAs, which in turn leads to changes in transcription patterns of the genes encoding the proteins required for fatty acid biosynthesis and degradation. The three central components of this system include FadL, an outer membrane-bound fatty acid transport protein, FadD, an inner membrane-associated long chain acyl-CoA synthetase (ACSL), and FadR, a long chain acyl-CoA-responsive transcription factor. The bacterial fatty acid transport and trafficking system leading to downstream fatty acid-responsive transcriptional regulation serves as a fundamental paradigm in biology, which provides a number of guiding principles that can be applied to more complex Foundations of Fatty Acid Transport and Trafficking System in E. coliThe transport of exogenous LCFAs across the cell envelope of E. coli requires both the transport the and the the The of long chain is the that the of the transcription the which in turn the of genes in fatty acid and biosynthesis in C.C. Mol. Microbiol. 1998; PubMed Scopus Google and C.C. P.N. Lipid Res. 1999; PubMed Scopus Google The in the of and the required to the genetic foundations of the fatty acid transport and trafficking system in E. coli Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google Scholar, Scholar, J. Bacteriol. PubMed Google that identified and of the genes of the fatty acid and distinct including Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google J. PubMed Scopus Google identified the gene in and at that this a of the fatty acid transport J. PubMed Scopus Google characterized the system for the of exogenous fatty acids and that this in the of fatty acid transport an for the transport system in E. is the membrane is to fatty acids from the membrane activation to J. PubMed Scopus Google a and D. Mol. PubMed Scopus Google identified a that type acyl-CoA synthetase to exogenous fatty acids that the gene in the of LCFAs and this a protein, the of the D. Mol. PubMed Scopus Google and Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google identified the gene in the of a in the transport of exogenous the of the gene an inner membrane-bound fatty acid that the transport of exogenous LCFAs and in Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google Scholar, J. Biol. Full Text PDF PubMed Google Scholar, J. Biol. Full Text PDF PubMed Google and of P.N. J. Biol. Full Text PDF PubMed Google that this in the outer membrane and to for The of in the outer membrane and in the transport of exogenous fatty acids the in the P.N. J. Bacteriol. 1988; 170: 2850-2854Crossref PubMed Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. Biochim. Biophys. Acta. PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google Scholar, P.N. DiRusso C.C. J. PubMed Google Scholar, C.C. P.N. Mol. Cell. 1999; PubMed Google Scholar).The the of the and genes is the long chain acyl-CoA transcription C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, C.C. J. Biol. Full Text PDF PubMed Google the of fatty acid transport and is LCA-CoA, this system to E. coli to transcriptional in the of exogenous long chain fatty DiRusso and C.C. Mol. Microbiol. 1998; PubMed Scopus Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google a of the regulation of the fatty acid and genes in E. coli. The transport of exogenous LCFAs across the cell envelope of E. coli requires both the transport the and the the The of long chain is the that the of the transcription the which in turn the of genes in fatty acid and biosynthesis in C.C. Mol. Microbiol. 1998; PubMed Scopus Google and C.C. P.N. Lipid Res. 1999; PubMed Scopus Google The in the of and the required to the genetic foundations of the fatty acid transport and trafficking system in E. coli Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google Scholar, Scholar, J. Bacteriol. PubMed Google that identified and of the genes of the fatty acid and distinct including Biophys. Res. PubMed Scopus Google Scholar, J. PubMed Scopus Google J. PubMed Scopus Google identified the gene in and at that this a of the fatty acid transport J. PubMed Scopus Google characterized the system for the of exogenous fatty acids and that this in the of fatty acid transport an for the transport system in E. is the membrane is to fatty acids from the membrane activation to J. PubMed Scopus Google a and D. Mol. PubMed Scopus Google identified a that type acyl-CoA synthetase to exogenous fatty acids that the gene in the of LCFAs and this a protein, the of the D. Mol. PubMed Scopus Google and Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google identified the gene in the of a in the transport of exogenous the of the gene an inner membrane-bound fatty acid that the transport of exogenous LCFAs and in Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google Scholar, J. Biol. Full Text PDF PubMed Google Scholar, J. Biol. Full Text PDF PubMed Google and of P.N. J. Biol. Full Text PDF PubMed Google that this in the outer membrane and to for The of in the outer membrane and in the transport of exogenous fatty acids the in the P.N. J. Bacteriol. 1988; 170: 2850-2854Crossref PubMed Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. Biochim. Biophys. Acta. PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google Scholar, P.N. DiRusso C.C. J. PubMed Google Scholar, C.C. P.N. Mol. Cell. 1999; PubMed Google The the of the and genes is the long chain acyl-CoA transcription C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, C.C. J. Biol. Full Text PDF PubMed Google the of fatty acid transport and is LCA-CoA, this system to E. coli to transcriptional in the of exogenous long chain fatty DiRusso and C.C. Mol. Microbiol. 1998; PubMed Scopus Google Scholar, C.C. P.N. Lipid Res. 1999; PubMed Scopus Google Scholar, P.N. DiRusso C.C. Biochim. Biophys. Acta. 1994; PubMed Scopus Google a of the regulation of the fatty acid and genes in E. coli. and Fatty Acid Transport outer membrane is the membrane-bound LCFA fatty acid transport proteins have been identified in of including in protein, membrane-bound D. Rev. PubMed Scopus Google as a for acid (1Abumrad N. Harmon C. Ibrahimi A. J. Lipid Res. 1998; 39: 2309-2318Abstract Full Text Full Text PDF PubMed Google Scholar, 2Abumrad N. Coburn C. Ibrahimi A. Biochim. Biophys. Acta. 1999; 1441: 4-13Crossref PubMed Scopus (208) Google and long chain fatty acids acid transport D. Stahl A. Lodish H.F. Proc. Natl. Acad. Sci. U. S. A. 1998; 95: 8625-8629Crossref PubMed Scopus (366) Google Scholar, N.R. D.A. J. Biol. 1999; Full Text Full Text PDF PubMed Scopus Google Scholar, D.A. PubMed Google The on the long chain fatty acid transport from the fundamental to of distinct which the of a as of the transport the outer membrane of the bacterial cell envelope the acid to exogenous LCFAs a The of long chain fatty acids to provides that the of the provides P.N. Biochim. Biophys. Acta. PubMed Scopus Google and to for of that the of the fatty acid is required P.N. Biochim. Biophys. Acta. PubMed Scopus Google the that specific the and the chain of the fatty the of the LCFA is essential for acid to as this in of the gene the LCFA and transport P.N. J. PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google of the of and that transport is to type whereas is type that the of in in acid that more for both LCFA and that in acid the of required for transport whereas the required for P.N. J. Biol. Full Text PDF PubMed Google of and in identified three acid and that to transport and one that to P.N. J. PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google identified of and and that have fatty acid to transport of in an P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google the first that a which is to LCFA The biochemical of the that from the of more the which this in fatty acid P.N. PubMed Scopus Google and of both the is a of a long of The in the the acid of the protein, which a that the on the to as the of three that the of the and a the the of a that the The is specific the of the in to in a in the of that to represent and fatty acid The is a the and the on the of the membrane a is and is to represent the fatty acid of lauryldimethylamine a used in is this least acid in to to this fatty acid to this of to the The to this fatty acid on the of the membrane is to one of the and and is to P.N. 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PubMed Google The on the long chain fatty acid transport from the fundamental to of distinct which the of a as of the transport process. the outer membrane of the bacterial cell envelope the acid to exogenous LCFAs a The of long chain fatty acids to provides that the of the provides P.N. Biochim. Biophys. Acta. PubMed Scopus Google and to for of that the of the fatty acid is required P.N. Biochim. Biophys. Acta. PubMed Scopus Google the that specific the and the chain of the fatty the of the LCFA is essential for acid to as this in of the gene the LCFA and transport P.N. J. PubMed Scopus Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google Scholar, P.N. J. Biol. Full Text PDF PubMed Google of the of and that transport is to type whereas is type that the of in in acid that more for both LCFA and that in acid the of required for transport whereas the required for P.N. J. Biol. Full Text PDF PubMed Google of and in identified three acid and that to transport and one that to P.N. J. 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