Coupled Growth and Division of Model Protocell Membranes
Abstract
The generation of synthetic forms of cellular life requires solutions to the problem of how biological processes such as cyclic growth and division could emerge from purely physical and chemical systems. Small unilamellar fatty acid vesicles grow when fed with fatty acid micelles and can be forced to divide by extrusion, but this artificial division process results in significant loss of protocell contents during each division cycle. Here we describe a simple and efficient pathway for model protocell membrane growth and division. The growth of large multilamellar fatty acid vesicles fed with fatty acid micelles, in a solution where solute permeation across the membranes is slow, results in the transformation of initially spherical vesicles into long thread-like vesicles, a process driven by the transient imbalance between surface area and volume growth. Modest shear forces are then sufficient to cause the thread-like vesicles to divide into multiple daughter vesicles without loss of internal contents. In an environment of gentle shear, protocell growth and division are thus coupled processes. We show that model protocells can proceed through multiple cycles of reproduction. Encapsulated RNA molecules, representing a primitive genome, are distributed to the daughter vesicles. Our observations bring us closer to the laboratory synthesis of a complete protocell consisting of a self-replicating genome and a self-replicating membrane compartment. In addition, the robustness and simplicity of this pathway suggests that similar processes might have occurred under the prebiotic conditions of the early Earth.
Introduction
Materials and Methods
Preparation of Large Monodisperse Multilamellar Vesicles
Adding Micelles and Imaging
Vesicle Growth and Division
Vesicle Counting
FRET Assay
Preparation of Unilamellar Fatty Acid Vesicles
Quantifying the Critical Shear Rate for Vesicle Division
Cycles of Vesicle Growth and Division
Measuring RNA Leakage
Results
Vesicle Growth and Division
Cycles of Growth and Division
Growth and Division of RNA-containing Model Protocells
Prebiotically Plausible Protocell Membranes
Mechanism of Initiation of Filamentous Vesicle Growth
Discussion
Supporting Information
Text S1 with references, Figures S1−S6, and movies S1−S3. This material is available free of charge via the Internet at http://pubs.acs.org.
Terms & Conditions
Most electronic Supporting Information files are available without a subscription to ACS Web Editions. Such files may be downloaded by article for research use (if there is a public use license linked to the relevant article, that license may permit other uses). Permission may be obtained from ACS for other uses through requests via the RightsLink permission system: http://pubs.acs.org/page/copyright/permissions.html.
Acknowledgment
We thank R. Bruckner, I. Chen, S. Chung, M. Elenko, R. Irwin, P. L. Luisi, A. Luptak, S. Mansy, D. Treco, C. Wong, and S. Zhou for helpful discussions and comments on the manuscript. This work was supported in part by grant EXB02-0031-0018 from the NASA Exobiology Program. J.W.S. is an Investigator of the Howard Hughes Medical Institute.
References
This article references 38 other publications.
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8Hargreaves, W. R. and Deamer, D. W. Biochemistry 1978, 17, 3759– 3768Google ScholarThere is no corresponding record for this reference.
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10Deamer, D. W. Nature (London) 1985, 317, 792– 794Google Scholar10https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaL2MXmtV2gsbw%253D&md5=d9e1446212d70d2f1c6e4ccbb31c7793Boundary structures are formed by organic components of the Murchison carbonaceous chondriteDeamer, David W.Nature (London, United Kingdom) (1985), 317 (6040), 792-4CODEN: NATUAS; ISSN:0028-0836.Formation of boundary structures by compds. extd. from the Murchison carbonaceous chondrite by org. solvents was investigated. A yellow, fluorescent ext. was obtained which appeared to contain carboxylate and phenolic groups and hydrocarbons. Two types of boundary structures were obsd. One class of microstructures was a viscous fluid droplet at alk. pH and a solid structure at neutral and acidic pH. The 2nd class of microstructure, formed only at alk. pH, was a thin boundary which functioned as a barrier to free diffusion, i.e., a membrane. The implications of these observations with respect to prebiotic evolution are discussed.
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11Deamer, D. W. and Pashley, R. M. Origins Life Evol. Biosphere 1989, 19, 21– 38Google Scholar11https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaL1MXkslegurs%253D&md5=d67bcc8337efb31d277d4adaea706953Amphiphilic components of the Murchison carbonaceous chondrite: surface properties and membrane formationDeamer, D. W.; Pashley, R. M.Origins of Life and Evolution of the Biosphere (1989), 19 (1), 21-38CODEN: OLEBEM; ISSN:0169-6149.Physicochem. properties of amphiphilic compds. in carbonaceous meteorites were investigated. The primary aim was to det. whether such materials represent plausible sources of lipid-like compds. that could have been involved as membrane components in primitive cells. Samples of the Murchison CM2 chondrite were extd. with chloroform-methanol, and the chloroform-sol. material was sepd. by 2-dimensional TLC. Fluorescence, I stains, and charring were used to identify major components on the plates. These were then scraped and eluted as specific fractions which were investigated by fluorescence and absorption spectra, surface chem. methods, gas chromatog.-mass spectrometry, and electron microscopy. Fraction 5 was strongly fluorescent, and contained pyrene and fluoranthene, the major polycyclic arom. hydrocarbons of the Murchison chondrite. This fraction was also present in exts. from the Murray and Mighei CM2 chondrites. Fraction 3 was surface active, forming apparent monomol. films at air-water interfaces. Surface force measurements suggested that fraction 3 contained acidic groups. Fraction 1 was also surface active, and certain components could self-assemble into membranous vesicles which encapsulated polar solutes. The observations reported demonstrate that org. compds. plausibly available on the primitive Earth through meteoritic infall are surface active, and have the ability to self-assemble into membranes.
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12McCollom, T. M., Ritter, G., and Simoneit, B. R. Origins Life Evol. Biosphere 1999, 29, 153– 166Google Scholar12https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaK1MXivFSrt7Y%253D&md5=48c775a8553772475d313adf50a3fb25Lipid synthesis under hydrothermal conditions by Fischer-Tropsch-type reactionsMccollom, Thomas M.; Ritter, Gilles; Simoneit, Bernd R. T.Origins of Life and Evolution of the Biosphere (1999), 29 (2), 153-166CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)Ever since their discovery in the late 1970's, mid-ocean-ridge hydrothermal systems have received a great deal of attention as a possible site for the origin of life on Earth (and environments analogous to mid-ocean-ridge hydrothermal systems are postulated to have been sites where life could have originated on Mars and elsewhere as well). Because no modern-day terrestrial hydrothermal systems are free from the influence of org. compds. derived from biol. processes, lab. expts. provide the best opportunity for confirmation of the potential for org. synthesis in hydrothermal systems. Here we report on the formation of lipid compds. during Fischer-Tropsch-type synthesis from aq. solns. of formic acid or oxalic acid. Optimum synthesis occurs in stainless steel vessels by heating at 175° for 2-3 days and produces lipid compds. ranging from C2 to >C35 which consist of n-alkanols, n-alkanoic acids, n-alkenes, n-alkanes and alkanones. The precursor carbon sources used are either formic acid or oxalic acid, which disproportionate to H2, CO2 and probably CO. Both carbon sources yield the same lipid classes with essentially the same ranges of compds. The synthesis reactions were confirmed by using 13C labeled precursor acids.
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13Naraoka, H., Shimoyama, A., and Harada, K. Origins Life Evol. Biosphere 1999, 29, 187– 201Google Scholar13https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaK1MXivFSrtL4%253D&md5=984508d8e139f6c272c1862a9b8108dcMolecular distribution of monocarboxylic acids in Asuka carbonaceous chondrites from AntarcticaNaraoka, Hiroshi; Shimoyama, Akira; Harada, KaoruOrigins of Life and Evolution of the Biosphere (1999), 29 (2), 187-201CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)Mol. distribution of low-mol.-wt. monocarboxylic acids was studied in three CM2 Asuka carbonaceous chondrites (A-881280, A-881334 and A-881458), which were recovered from Antarctica by the 29th Japanese Antarctic Research Expedition in 1988. GC and GC/MS analyses identified more than 30 monocarboxylic acids in A-881458, including aliph. and arom. acids with various structural isomers. Isomeric phenolic compds. were also identified. The aliph. carboxylic acids have straight-chain structures having 2 to 12 carbon atoms (C2 to C12), and branched-chain structures (C4 to C9). The quantities of straight-chain acids decrease logarithmically with increasing carbon no. At the same carbon no., a straight-chain isomer is always predominant compared to branched-chain isomers. All of the 14 possible C4, C5 and C6 aliph. monocarboxylic acids (not including optical isomers) have been identified, although all the isomers were not reported in Murchison and Y-791198 meteorites. Of the 17 possible isomeric C7 acids, at least 14 isomers were tentatively identified by mass spectra (EI and CI mode). At C8 or above, peaks of branched-chain isomers become obscure, probably due to the large no. of isomers and small concns. Branched-chain monocarboxylic acids over C6 have never been reported in Murchison. Although occurrence of aliph. acids are similar between A-881458 and Murchison at C4, C5 and C6 acids, a major difference is that A-881458 as well as Y-791198 have straight-chain predominance among isomers in contrast to Murchison being branched-chain predominant. In the case of isomeric arom. compds. such as toluic acids and cresols, m-toluic acid and p-cresol are more abundant among their isomers, resp. The mol. distribution may not reflect thermodn. equil. but rather a kinetically controlled process for their formation mechanism. The other two CM2 chondrites (A-881280 and A-881334) were depleted in carboxylic acids in spite of similar carbon contents. The depletion is not due to weathering on ice, because the degrees of weathering are small and similar among the three chondrites. Probably those latter two chondrites may have been subjected to aq. alteration or metamorphism on their meteorite parent bodies.
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14Nooner, D. W., Gibert, J. M., Gelpi, E., and Oro, J. Geochim. Cosmochim. Acta 1976, 40, 915– 924Google ScholarThere is no corresponding record for this reference.
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31Karlsson, M., Sott, K., Davidson, M., Cans, A. S., Linderholm, P., Chiu, D., and Orwar, O. Proc. Natl. Acad. Sci. U.S.A. 2002, 99, 11573– 11578Google ScholarThere is no corresponding record for this reference.
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1Szostak, J. W., Bartel, D. P., and Luisi, P. L. Nature (London) 2001, 409, 387– 3901https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD3MXmslSjtw%253D%253D&md5=a1df371d9efa0c022efc96bf360114d8Synthesizing lifeSzostak, Jack W.; Bartel, David P.; Luisi, P. LuigiNature (London) (2001), 409 (6818), 387-390CODEN: NATUAS; ISSN:0028-0836. (Nature Publishing Group)Advances in directed evolution and membrane biophysics make the synthesis of simple living cells, if not yet foreseeable reality, an imaginable goal. Overcoming the many scientific challenges along the way will deepen our understanding of the essence of cellular life and its origin on Earth.
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2Johnston, W. K., Unrau, P. J., Lawrence, M. S., Glasner, M. E., and Bartel, D. P. Science 2001, 292, 1319– 1325There is no corresponding record for this reference.
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3Hanczyc, M. M., Fujikawa, S. M., and Szostak, J. W. Science 2003, 302, 618– 622There is no corresponding record for this reference.
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4Hanczyc, M. M. and Szostak, J. W. Curr. Opin. Chem. Biol. 2004, 8, 660– 6644https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD2cXhtVWjtrrF&md5=76d9d0f6826342fab1cbef2e27e4ac6aReplicating vesicles as models of primitive cell growth and divisionHanczyc, Martin M.; Szostak, Jack W.Current Opinion in Chemical Biology (2004), 8 (6), 660-664CODEN: COCBF4; ISSN:1367-5931. (Elsevier Ltd.)A review and discussion. Primitive cells, lacking the complex bio-machinery present in modern cells, would have had to rely on the self-organizing properties of their components and on interactions with their environment to achieve basic cellular functions such as growth and division. Many bilayer-membrane vesicles, depending on their compn. and environment, can exhibit complex morphol. changes such as growth, fusion, fission, budding, internal vesicle assembly, and vesicle-surface interactions. The rich dynamic properties of these vesicles provide interesting models of how primitive cellular replication might have occurred in response to purely phys. and chem. forces.
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5Mansy, S. S., Schrum, J. P., Krishnamurthy, M., Tobe, S., Treco, D. A., and Szostak, J. W. Nature (London) 2008, 454, 122– 1255https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD1cXotVertLw%253D&md5=6752e68f267b7e5a29a48e51e3b78b9cTemplate-directed synthesis of a genetic polymer in a model protocellMansy, Sheref S.; Schrum, Jason P.; Krishnamurthy, Mathangi; Tobe, Sylvia; Treco, Douglas A.; Szostak, Jack W.Nature (London, United Kingdom) (2008), 454 (7200), 122-125CODEN: NATUAS; ISSN:0028-0836. (Nature Publishing Group)Contemporary phospholipid-based cell membranes are formidable barriers to the uptake of polar and charged mols. ranging from metal ions to complex nutrients. Modern cells therefore require sophisticated protein channels and pumps to mediate the exchange of mols. with their environment. The strong barrier function of membranes has made it difficult to understand the origin of cellular life and has been thought to preclude a heterotrophic lifestyle for primitive cells. Although nucleotides can cross dimyristoyl phosphatidylcholine membranes through defects formed at the gel-to-liq. transition temp., phospholipid membranes lack the dynamic properties required for membrane growth. Fatty acids and their corresponding alcs. and glycerol monoesters are attractive candidates for the components of protocell membranes because they are simple amphiphiles that form bilayer membrane vesicles that retain encapsulated oligonucleotides and are capable of growth and division. Here we show that such membranes allow the passage of charged mols. such as nucleotides, so that activated nucleotides added to the outside of a model protocell spontaneously cross the membrane and take part in efficient template copying in the protocell interior. The permeability properties of prebiotically plausible membranes suggest that primitive protocells could have acquired complex nutrients from their environment in the absence of any macromol. transport machinery; i.e., they could have been obligate heterotrophs.
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6Mansy, S. S. and Szostak, J. W. Proc. Natl. Acad. Sci. U.S.A. 2008, 105, 13351– 13355There is no corresponding record for this reference.
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7Gebicki, J. M. and Hicks, M. Nature (London) 1973, 243, 232– 2347https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaE3sXktl2msLc%253D&md5=4977bbc7f4107e8f177f00ffe4aec312Ufasomes are stable particles surrounded by unsaturated fatty acid membranesGebicki, J. M.; Hicks, M.Nature (London, United Kingdom) (1973), 243 (5404), 232-4CODEN: NATUAS; ISSN:0028-0836.For the purpose of studying ufasomes, thin layer of various fatty acids and acid mixts. were deposited in glass vessels by evapn. of chloroform solns. and the film resuspended by hand or mech. shaking in water buffered at pH 8-9. The suspensions contained spherical particles visible under a microscope but showing no fine structure. The spheres were unstable to drying and the technique of neg. staining for electron microscopy was unsuccessful. Freeze-etched specimens, however, showed clearly under an electron microscope that the spheres consist of an aq. interior surrounded by 1 or more concentric layers of fatty acid membranes. A sphere made from oleic acid had at least 12 layers. Some spheres had complex interior structures with many orientated sheets of incomplete membranes and others consisted of collections of small spheres within a common envelope. The sepn. between individual bilayers varied between 20 and 30 nm. This suggests that here, as in liposomes, aq, shells are interposed between individual membranes. The bilayers were stabilized by ionization of a small proportion of the weak acid groups at the slightly alk. pH. Because of the obligatory presence of unsatd. fatty acids the particles have been called ufasomes. The presence of aq. spaces within the ufasomes was confirmed by forming them in the presence of glucose (I), removing exterior I by dialysis, and measuring the amt. of sugar released by spheres disrupted with Triton X-100 detergent. An interesting feature of ufasomes is the possibility that they can form from mols. synthesized in the absence of living organisms.
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8Hargreaves, W. R. and Deamer, D. W. Biochemistry 1978, 17, 3759– 3768There is no corresponding record for this reference.
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9Walde, P., Wick, R., Fresta, M., Mangone, A., and Luisi, P. L. J. Am. Chem. Soc. 1994, 116, 11649– 11654There is no corresponding record for this reference.
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10Deamer, D. W. Nature (London) 1985, 317, 792– 79410https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaL2MXmtV2gsbw%253D&md5=d9e1446212d70d2f1c6e4ccbb31c7793Boundary structures are formed by organic components of the Murchison carbonaceous chondriteDeamer, David W.Nature (London, United Kingdom) (1985), 317 (6040), 792-4CODEN: NATUAS; ISSN:0028-0836.Formation of boundary structures by compds. extd. from the Murchison carbonaceous chondrite by org. solvents was investigated. A yellow, fluorescent ext. was obtained which appeared to contain carboxylate and phenolic groups and hydrocarbons. Two types of boundary structures were obsd. One class of microstructures was a viscous fluid droplet at alk. pH and a solid structure at neutral and acidic pH. The 2nd class of microstructure, formed only at alk. pH, was a thin boundary which functioned as a barrier to free diffusion, i.e., a membrane. The implications of these observations with respect to prebiotic evolution are discussed.
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11Deamer, D. W. and Pashley, R. M. Origins Life Evol. Biosphere 1989, 19, 21– 3811https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaL1MXkslegurs%253D&md5=d67bcc8337efb31d277d4adaea706953Amphiphilic components of the Murchison carbonaceous chondrite: surface properties and membrane formationDeamer, D. W.; Pashley, R. M.Origins of Life and Evolution of the Biosphere (1989), 19 (1), 21-38CODEN: OLEBEM; ISSN:0169-6149.Physicochem. properties of amphiphilic compds. in carbonaceous meteorites were investigated. The primary aim was to det. whether such materials represent plausible sources of lipid-like compds. that could have been involved as membrane components in primitive cells. Samples of the Murchison CM2 chondrite were extd. with chloroform-methanol, and the chloroform-sol. material was sepd. by 2-dimensional TLC. Fluorescence, I stains, and charring were used to identify major components on the plates. These were then scraped and eluted as specific fractions which were investigated by fluorescence and absorption spectra, surface chem. methods, gas chromatog.-mass spectrometry, and electron microscopy. Fraction 5 was strongly fluorescent, and contained pyrene and fluoranthene, the major polycyclic arom. hydrocarbons of the Murchison chondrite. This fraction was also present in exts. from the Murray and Mighei CM2 chondrites. Fraction 3 was surface active, forming apparent monomol. films at air-water interfaces. Surface force measurements suggested that fraction 3 contained acidic groups. Fraction 1 was also surface active, and certain components could self-assemble into membranous vesicles which encapsulated polar solutes. The observations reported demonstrate that org. compds. plausibly available on the primitive Earth through meteoritic infall are surface active, and have the ability to self-assemble into membranes.
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12McCollom, T. M., Ritter, G., and Simoneit, B. R. Origins Life Evol. Biosphere 1999, 29, 153– 16612https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaK1MXivFSrt7Y%253D&md5=48c775a8553772475d313adf50a3fb25Lipid synthesis under hydrothermal conditions by Fischer-Tropsch-type reactionsMccollom, Thomas M.; Ritter, Gilles; Simoneit, Bernd R. T.Origins of Life and Evolution of the Biosphere (1999), 29 (2), 153-166CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)Ever since their discovery in the late 1970's, mid-ocean-ridge hydrothermal systems have received a great deal of attention as a possible site for the origin of life on Earth (and environments analogous to mid-ocean-ridge hydrothermal systems are postulated to have been sites where life could have originated on Mars and elsewhere as well). Because no modern-day terrestrial hydrothermal systems are free from the influence of org. compds. derived from biol. processes, lab. expts. provide the best opportunity for confirmation of the potential for org. synthesis in hydrothermal systems. Here we report on the formation of lipid compds. during Fischer-Tropsch-type synthesis from aq. solns. of formic acid or oxalic acid. Optimum synthesis occurs in stainless steel vessels by heating at 175° for 2-3 days and produces lipid compds. ranging from C2 to >C35 which consist of n-alkanols, n-alkanoic acids, n-alkenes, n-alkanes and alkanones. The precursor carbon sources used are either formic acid or oxalic acid, which disproportionate to H2, CO2 and probably CO. Both carbon sources yield the same lipid classes with essentially the same ranges of compds. The synthesis reactions were confirmed by using 13C labeled precursor acids.
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13Naraoka, H., Shimoyama, A., and Harada, K. Origins Life Evol. Biosphere 1999, 29, 187– 20113https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaK1MXivFSrtL4%253D&md5=984508d8e139f6c272c1862a9b8108dcMolecular distribution of monocarboxylic acids in Asuka carbonaceous chondrites from AntarcticaNaraoka, Hiroshi; Shimoyama, Akira; Harada, KaoruOrigins of Life and Evolution of the Biosphere (1999), 29 (2), 187-201CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)Mol. distribution of low-mol.-wt. monocarboxylic acids was studied in three CM2 Asuka carbonaceous chondrites (A-881280, A-881334 and A-881458), which were recovered from Antarctica by the 29th Japanese Antarctic Research Expedition in 1988. GC and GC/MS analyses identified more than 30 monocarboxylic acids in A-881458, including aliph. and arom. acids with various structural isomers. Isomeric phenolic compds. were also identified. The aliph. carboxylic acids have straight-chain structures having 2 to 12 carbon atoms (C2 to C12), and branched-chain structures (C4 to C9). The quantities of straight-chain acids decrease logarithmically with increasing carbon no. At the same carbon no., a straight-chain isomer is always predominant compared to branched-chain isomers. All of the 14 possible C4, C5 and C6 aliph. monocarboxylic acids (not including optical isomers) have been identified, although all the isomers were not reported in Murchison and Y-791198 meteorites. Of the 17 possible isomeric C7 acids, at least 14 isomers were tentatively identified by mass spectra (EI and CI mode). At C8 or above, peaks of branched-chain isomers become obscure, probably due to the large no. of isomers and small concns. Branched-chain monocarboxylic acids over C6 have never been reported in Murchison. Although occurrence of aliph. acids are similar between A-881458 and Murchison at C4, C5 and C6 acids, a major difference is that A-881458 as well as Y-791198 have straight-chain predominance among isomers in contrast to Murchison being branched-chain predominant. In the case of isomeric arom. compds. such as toluic acids and cresols, m-toluic acid and p-cresol are more abundant among their isomers, resp. The mol. distribution may not reflect thermodn. equil. but rather a kinetically controlled process for their formation mechanism. The other two CM2 chondrites (A-881280 and A-881334) were depleted in carboxylic acids in spite of similar carbon contents. The depletion is not due to weathering on ice, because the degrees of weathering are small and similar among the three chondrites. Probably those latter two chondrites may have been subjected to aq. alteration or metamorphism on their meteorite parent bodies.
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14Nooner, D. W., Gibert, J. M., Gelpi, E., and Oro, J. Geochim. Cosmochim. Acta 1976, 40, 915– 924There is no corresponding record for this reference.
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15Rushdi, A. I. and Simoneit, B. R. Origins Life Evol. Biosphere 2001, 31, 103– 11815https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD3MXjtVyhtL8%253D&md5=8aa116c6d991b696f4a7758f3932065fLipid formation by aqueous Fischer-Tropsch-type synthesis over a temperature range of 100 to 400°CRushdi, Ahmed I.; Simoneit, Bernd R. T.Origins of Life and Evolution of the Biosphere (2001), 31 (1-2), 103-118CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)The formation of lipid compds. during an aq. Fischer-Tropsch-type reaction was studied with solns. of oxalic acid as the carbon and hydrogen source. The reactions were conducted in stainless steel vessels by heating the oxalic acid soln. at discrete temps. from 100 to 400°C, at intervals of 50°C for two days each. The max. lipid yield, esp. for oxygenated compds., is in the window of 150-250°C. At a temp. of 100°C only a trace amt. of lipids was detected. At temps. above 150°C the lipid components ranged from C12 to > C33 and included n-alkanols, n-alkanoic acids, n-alkyl formates, n-alkanals, n-alkanones, n-alkanes, and n-alkenes, all with essentially no carbon no. preference. The n-alkanes increased in concn. over the oxygenated compds. at temps. of 200°C and above, with a slight redn. in their carbon no. ranges due to cracking. The n-alkanoic acids increased while n-alkanols decreased with increasing temp. above 200°C. At temps. above 300°C synthesis competes with cracking and reforming reactions. At 400°C significant cracking was obsd. and polynuclear arom. hydrocarbons and their alkylated homologs were detected. The formation of lipid compds. by aq. FTT reactions proceeds by insertion of a CO group at the terminal end of a carboxylic acid functionality to form n-oxoalkanoic acids, followed by redn. to n-alkanoic acids, to n-alkanals, then to n-alkanols. The n-alkenes are intermediate homologs for n-alkan-2-ones and n-alkanes. This proposed mechanism for aq. FTT synthesis differs from the surface-catalyzed stepwise FT process (i.e., gaseous) of polymn. of methylene reported in the literature.
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16Yuen, G. U. and Kvenvold, K. A. Nature (London) 1973, 246, 301– 30216https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaE2cXmt1aqsA%253D%253D&md5=399e3bbbc675bc4f5f7da4601368184fMonocarboxylic acids in Murray and Murchison carbonaceous meteoritesYuen, George U.; Kvenvolden, Keith A.Nature (London, United Kingdom) (1973), 246 (5431), 301-3CODEN: NATUAS; ISSN:0028-0836.Straight and branched-chain C2-C8 monocarboxylic acids were found in Murray and Murchison carbonaceous meteorites. Fifteen acids were identified by gas chromatog.-mass spectrometry, 10 pos. and 5 tentatively. The concns. 10-60 μg/g, and distribution of the acids suggested that synthesis was random chem. and nonbiol.
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17Chen, I. A., Roberts, R. W., and Szostak, J. W. Science 2004, 305, 1474– 1476There is no corresponding record for this reference.
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18Chen, I. A. and Szostak, J. W. Biophys. J. 2004, 87, 988– 998There is no corresponding record for this reference.
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19Chen, I. A. and Szostak, J. W. Proc. Natl. Acad. Sci. U.S.A. 2004, 101, 7965– 7970There is no corresponding record for this reference.
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20Sacerdote, M. G. and Szostak, J. W. Proc. Natl. Acad. Sci. U.S.A. 2005, 102, 6004– 6008There is no corresponding record for this reference.
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21Berclaz, N., Muller, M., Walde, P., and Luisi, P. L. J. Phys. Chem. B 2001, 105, 1056– 1064There is no corresponding record for this reference.
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22Blochliger, E., Blocher, M., Walde, P., and Luisi, P. L. J. Phys. Chem. B 1998, 102, 10383– 10390There is no corresponding record for this reference.
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23Luisi, P. L., Stano, P., Rasi, S., and Mavelli, F. Artif. Life. 2004, 10, 297– 30823https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A280%3ADC%252BD2czkt1Gjtw%253D%253D&md5=1149865f76b82466f116a24fbbdf4f31A possible route to prebiotic vesicle reproductionLuisi Pier Luigi; Rasi Pasquale Stano Silvia; Mavelli FabioArtificial life (2004), 10 (3), 297-308 ISSN:1064-5462.Spherical bounded structures such as those formed by surfactant aggregates (mostly micelles and vesicles), with an inside that is chemically and physically different from the outside medium, can be seen as primitive cell models. As such, they are fundamental structures for the theory of autopoiesis as originally formulated by Varela and Maturana. In particular, since self-reproduction is a very important feature of minimal cellular life, the study of self-reproduction of micelles and vesicles represents a quite challenging bio-mimetic approach. Our laboratory has put much effort in recent years into implementing self-reproduction of vesicles as models for self-reproduction of cellular bounded structures, and this article is a further contribution in this direction. In particular, we deal with the so-called matrix effect of vesicles, related to the fact that when fresh surfactant is added to an aqueous solution containing preformed vesicles of a very narrow size distribution, the newly formed vesicles (instead of being polydisperse, as is usually the case) have dimensions very close to those of the preformed ones. In practice, this corresponds to a mechanism of reproduction of vesicles of the same size. In this article, the matrix effect is re-elaborated in the perspective of the origin of life, and in particular in terms of the prebiotic mechanisms that might permit the growth and reproduction of vesicles. The data are analyzed by dynamic light scattering with a new program that permits the calculation of the number-weighted size distribution. It is shown that, on adding a stoichiometric amount of oleate micelles to preformed oleate vesicles extruded at 50 and 100 nm, the final distribution contains about twice the initial number of particles, centered around 50 and 100 nm. The same holds when oleate is added to preformed phospholipid liposomes. By contrast, when the same amount of oleate is added to an aqueous solution (as a control experiment), a very broad distribution ranging between 20 and 1000 nm is obtained. The data can then be seen as a kind of reproduction of the same size vesicles, and the argument is advanced that this may correspond to a simple prebiotic mechanism of vesicle multiplication in prebiotic times, when only physical forces might be responsible for the basic mechanisms of early protocell growth and division. Preliminary data also show that repeated addition of oleate maintains the same basic initial features, and that surfactants other than oleate also respect the reproductive mode of the matrix effect.
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24Rasi, S., Mavelli, F., and Luisi, P. L. J. Phys. Chem. B 2003, 107, 14068– 14076There is no corresponding record for this reference.
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25Rasi, S., Mavelli, F., and Luisi, P. L. Origins Life Evol. Biosphere 2004, 34, 215– 22425https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD3sXhtVSjs7bL&md5=b162b512a26954629ca5b42ae051a51fMatrix effect in oleate micelles-vesicles transformationRasi, Silvia; Mavelli, Fabio; Luisi, Pier LuigiOrigins of Life and Evolution of the Biosphere (2004), 34 (1-2), 215-224CODEN: OLEBEM; ISSN:0169-6149. (Kluwer Academic Publishers)It is accepted by many authors that the formation of closed mol. structures is a key step in the evolution of life. Oleate vesicles represent a good model system in this framework due to the fact that they self-assemble spontaneously and that fatty acids are considered as possible prebiotic structures. In this contribution, we will focus the attention on the transition from oleate micelles to oleic acid/oleate vesicles induced by a pH change. This transformation is strongly influenced by the presence of pre-formed vesicles. We called this phenomenon the matrix effect. The influence of pre-added POPC liposomes (POPC = 1-palmitoyl-2-oleoyl- sn-glycerol-3-phosphocholine) and oleic acid/oleate vesicles on the process rate and on the final size distribution will be discussed elucidating the main differences between these two systems.
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26Luisi, P. L. The emergence of life: from chemical origins to synthetic biology; Cambridge University Press: Cambridge, 2006.There is no corresponding record for this reference.
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27Stano, P., Wehrli, E., and Luisi, P. L. J. Phys.: Condens. Matter 2006, 18, S2231– S2238There is no corresponding record for this reference.
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28Zhu, T. and Szostak, J.PLoS ONE2009,
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29Karlsson, M., Nolkrantz, K., Davidson, M. J., Stromberg, A., Ryttsen, F., Akerman, B., and Orwar, O. Anal. Chem. 2000, 72, 5857– 5862There is no corresponding record for this reference.
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30Karlsson, A., Karlsson, R., Karlsson, M., Cans, A. S., Stromberg, A., Ryttsen, F., and Orwar, O. Nature (London) 2001, 409, 150– 15230https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADC%252BD3MXlvVykug%253D%253D&md5=cca84609cde3fe650b321ab4e34844c5Molecular engineering: Networks of nanotubes and containersKarlsson, Anders; Karlsson, Roger; Karlsson, Mattias; Cans, Ann-Sofie; Stromberg, Anette; Ryttsen, Frida; Orwar, OweNature (London) (2001), 409 (6817), 150-152CODEN: NATUAS; ISSN:0028-0836. (Nature Publishing Group)The authors have constructed complex two-dimensional microscopic networks of phospholipid bilayer nanotubes and containers in which they were able to control the connectivity, container size, nanotube length, and angle between the nanotube extensions. Containers within these networks can be chem. differentiated and materials successfully routed between two containers connected by a common nanotube. These networks will enable model systems to be devised for studying confined biochem. reactions, intracellular transport phenomena and chem. computations.
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31Karlsson, M., Sott, K., Davidson, M., Cans, A. S., Linderholm, P., Chiu, D., and Orwar, O. Proc. Natl. Acad. Sci. U.S.A. 2002, 99, 11573– 11578There is no corresponding record for this reference.
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32Miller, S. L. Science 1953, 117, 528– 9There is no corresponding record for this reference.
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33Miller, S. L. and Urey, H. C. Science 1959, 130, 245– 5133https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaG1MXhtVSjt7Y%253D&md5=def25fcb01ba9073f23ffcc4d6dfc776Organic compound synthesis on the primitive earth. Several questions about the origin of life have been answered, but much remains to be studiedMiller, Stanley L.; Urey, Harold C.Science (Washington, DC, United States) (1959), 130 (), 245-51CODEN: SCIEAS; ISSN:0036-8075.A review with 42 references.
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34Kvenvolden, K., Lawless, J., Pering, K., Peterson, E., Flores, J., Ponnamperuma, C., Kaplan, I. R., and Moore, C. Nature (London) 1970, 228, 923– 634https://chemport.cas.org/services/resolver?origin=ACS&resolution=options&coi=1%3ACAS%3A528%3ADyaE3MXisVCnsg%253D%253D&md5=de3a5fdb67df1b0440c580c808c109e7Evidence for extraterrestrial amino-acids and hydrocarbons in the Murchison meteoriteKvenvolden, Keith A.; Lawless, James; Pering, Katherine; Peterson, Etta; Flores, Jose; Ponnamperuma, Cyril; Kaplan, Isaac R.; Moore, CarletonNature (London, United Kingdom) (1970), 228 (5275), 923-6CODEN: NATUAS; ISSN:0028-0836.The Murchison meteor-ite which fell near Murchison, Victoria, Australia is a type II carbonaceous chondrite. Conventional ion-exchange chromatog. of the hydrolyzed water ext. from 10 g of the sample revealed the presence of glycine (6 μg/g), alanine, glutamic acid, valine, and proline. The presence of amino acids and their enantiomeric distribution were established by capillary gas chromatog. of the N-trifluoroacetyl-D-2-butyl esters of the amino acids. The presence of 2-methylalanine and sarcosine amino acids, not commonly found in biol. systems, was suggested by gas chromatogram. The presence of almost equal amts. of the D and L enantiomers of valine, proline, alanine, and glutamic acid minimizes the possibility of terrestrial contamination from biol. sources in which only the L-form occurs. The results support the contention that the org. mols. identified are abiotic and possible extraterrestrial in origin. A very complex mixt. of alkanes has been isolated from the meteorite. A similarity is established between the aliphatic hydrocarbons from the meteorite and the spark discharge methane.
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35Bozic, B., Gomiscek, G., Kralj-Iglic, V., Svetina, S., and Zeks, B. Eur. Biophys. J. 2002, 31, 487– 96There is no corresponding record for this reference.
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36Dobereiner, H. G. In Giant Vesicles; John Wiley and Sons, Inc.: New York, 2000.There is no corresponding record for this reference.
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37Dobereiner, H. G. Curr. Opin. Colloid Interface Sci. 2000, 5, 256– 263There is no corresponding record for this reference.
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38Bar-Ziv, R. and Moses, E. Phys. Rev. Lett. 1994, 73, 1392– 1395There is no corresponding record for this reference.
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