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The three main structures phospholipids form in solution; the liposome (a closed bilayer), the micelle and the bilayer.

A protocell (or protobiont) is a self-organized, endogenously ordered, spherical collection of lipids proposed as a stepping-stone to the origin of life.[1] A central question in evolution is how simple protocells first arose and began the competitive process that drove the evolution of life. Although a functional protocell has not yet been achieved in a laboratory setting, the goal to understand the process appears well within reach.[2][3][4][5]


  • Selectivity for compartmentalization 1
    • Energy gradient 1.1
  • Vesicles and micelles 2
    • Geothermal ponds and clay 2.1
    • Montmorillonite bubbles 2.2
    • Membrane transport 2.3
  • Artificial models 3
    • Langmuir-Blodgett deposition 3.1
    • Jeewanu 3.2
  • Ethics and controversy 4
  • Endosymbiotic theory 5
  • See also 6
  • References 7
  • External links 8

Selectivity for compartmentalization

Self-assembled vesicles are essential components of primitive cells.[1] The

  • "Living Chemistry & A Natural History of Protocells." Synth-ethic: Art and Synthetic Biology Exhibition (2013) at the Natural History Museum, Vienna, Austria.
  • "Protocells: Bridging Nonliving and Living Matter." Edited by Steen Rasmussen, Mark A. Bedau, Liaochai Chen, David Deamer, David Krakauer, Norman, H.Packard and Peter F. Stadler. MIT Press, Cambridge, Massachusetts. 2008.

External links

  1. ^ a b c d Chen, Irene A.; Walde, Peter (July 2010). "From Self-Assembled Vesicles to Protocells" (PDF). Cold Spring Harb Perspect Biol. 2 (7).  
  2. ^ National Science Foundation (2013). "Exploring Life's Origins - Protocells". Retrieved 2014-03-18. 
  3. ^ a b c d Chen, Irene A. (8 December 2006). "The Emergence of Cells During the Origin of Life". Science 314 (5805): 1558–1559.  
  4. ^ a b c d Zimmer, Carl (26 June 2004). "What Came Before DNA?". Discover Magazine: 1–5. 
  5. ^ Rasmussen, Steen (2 July 2014). "Scientists Create Possible Precursor to Life". A Letters Journal Exploring the Frontiers of Physics. Volume 107, Number 2, July 2014. (Astrobiology Web). Retrieved 2014-10-24. 
  6. ^ Shapiro, Robert (12 February 2007). "A Simpler Origin for Life". Scientific American. 
  7. ^ Vodopich, Darrell S.; Moore., Randy (2002). "The Importance of Membranes". Biology Laboratory Manual, 6/a. McGraw-Hill. Retrieved 2014-03-17. 
  8. ^ Chang, Thomas Ming Swi (2007). Artificial cells : biotechnology, nanomedicine, regenerative medicine, blood substitutes, bioencapsulation, cell/stem cell therapy. Hackensack, N.J.: World Scientific.  
  9. ^ a b Knowles, JR (1980). "Enzyme-catalyzed phosphoryl transfer reactions". Annu. Rev. Biochem. 49: 877–919.  
  10. ^ a b Campbell, Neil A.; Williamson, Brad; Heyden, Robin J. (2006). Biology: Exploring Life. Boston, Massachusetts: Pearson Prentice Hall.  
  11. ^ a b Clavin, Whitney (13 March 2014). "How Did Life Arise? Fuel Cells May Have Answers".  
  12. ^ Walsby, AE (1994). "Gas vesicles". Microbiological reviews 58 (1): 94–144.  
  13. ^ Szostak, Jack W. (3 September 2004). "Battle of the Bubbles May Have Sparked Evolution".  
  14. ^ a b National Science Foundation (2013). "Membrane Lipids of Past and Present". Exploring Life's Origins Project - A timeline of Life's Evolution. Retrieved 2014-03-17. 
  15. ^ a b Switek, Brian (13 February 2012). "Debate bubbles over the origin of life". Nature - News. 
  16. ^ Szostak, Jack W. (4 June 2008). "Researchers Build Model Protocell Capable of Copying DNA". Howard Huges Medical Institute. 
  17. ^ Cohen, Philip (23 October 2003). "Clay's matchmaking could have sparked life". New Scientist. Journal reference: Science (vol 302, p 618 ) 
  18. ^ a b Stone, Howard A. (7 February 2011). "Clay-armored bubbles may have formed first protocells". Harvard School of Engineering and Applied Sciences. 
  19. ^ Müller, A. W. (June 2006). "Re-creating an RNA world". Cell Mol Life Sci. 63 (11): 1278–93.  
  20. ^ Ma, Wentao; Yu, Chunwu; Zhang, Wentao; Hu., Jiming (Nov 2007). "Nucleotide synthetase ribozymes may have emerged first in the RNA world". RNA 13 (11): 2012–2019.  
  21. ^ Demanèche, S; Bertolla, F; Buret, F; et. al (August 2001). "Laboratory-scale evidence for lightning-mediated gene transfer in soil". Appl. Environ. Microbiol. 67 (8): 3440–4.  
  22. ^ Neumann, E; Schaefer-Ridder, M; Wang, Y; Hofschneider, PH (1982). "Gene transfer into mouse lyoma cells by electroporation in high electric fields". EMBO J. 1 (7): 841–5.  
  23. ^ Norris, V.; Raine, D.J. (October 1998). "A fission-fussion origin for life". Orig Life Evol Biosph 28 (4-4): 523–537.  
  24. ^ a b "Scientists Create Artificial Cell Membranes". Astrobiology Magazine. 4 October 2014. Retrieved 2014-05-07. 
  25. ^ a b Matosevic, Sandro; Paegel, Brian M. (29 September 2013). "Layer-by-layer cell membrane assembly". Nature Chemistry 5 (11): 958–963.  
  26. ^ a b c Grote, M (September 2011). "'"Jeewanu, or the 'particles of life (PDF). Journal of Biosciences 36 (4): 563–570.  
  27. ^ a b Gupta, V. K.; Rai, R. K. (2013). "Histochemical localisation of RNA-like material in photochemically formed self-sustaining, abiogenic supramolecular assemblies 'Jeewanu'". Int. Res. J. of Science & Engineering 1 (1): 1-4.  
  28. ^ Ponnamperuma, Cyril (1967). "'"A review of some experiments on the synthesis of 'Jeewanu (PDF). NASA Technical Memorandum X-1439 (Moffett Field, California: Ames Research Center). 
  29. ^ Dworkin, Jason P.; Deamer, David W.; Sandford, Scott A.; Allamandola, Louis J. (30 January 2001). "Self-assembling amphiphilic molecules: Synthesis in simulated interstellar/precometary ices". Proceedings of the National Academy of Sciences of the United States of America 98 (3): 815–9.  
  30. ^ Mullen, L (5 September 2005). "Building Life from Star-Stuff". Astrobiology Magazine. 
  31. ^ "Life after the synthetic cell". Nature 465 (7297): 422–424. 27 May 2010.  
  32. ^ Beadau, Mark A.; Parke, Emily C. (2009). The ethics of protocells moral and social implications of creating life in the laboratory (Online ed.). Cambridge, Mass.: MIT Press.  
  33. ^ Wallin, IE (1923). "The Mitochondria Problem". The American Naturalist 57 (650): 255–61.  
  34. ^ Wallin, I.E. (1927). Symbionticism and the origin of species. Baltimore: Williams & Wilkins Company. p. 171. 
  35. ^ Schimper, AFW (1883). "Über die Entwicklung der Chlorophyllkörner und Farbkörper". Bot. Zeitung (in German) 41: 105–14, 121–31, 137–46, 153–62. 
  36. ^ Andersson, SG; Karlberg, O; Canbäck, B; Kurland, CG (January 2003). "On the origin of mitochondria: a genomics perspective". Philosophical Transactions of the Royal Society B 358 (1429): 165–77; discussion 177–9.  
  37. ^ "Mitochondria Share an Ancestor With SAR11, a Globally Significant Marine Microbe". ScienceDaily. 25 July 2011. 
  38. ^ Thrash, J. Cameron; et. al (2011). "Phylogenomic evidence for a common ancestor of mitochondria and the SAR11 clade".  
  39. ^ Henze, K; Martin, William (2003). "Evolutionary biology: essence of mitochondria". Nature 426 (6963): 127–8.  


See also

The mitochondrion (plural mitochondria) is a eukaryotic cells (the cells that make up plants, animals, fungi, and many other forms of life).[39] A mitochondrion produces adenosine triphosphate (ATP) and it is closely related to the adenosine nucleotide, a monomer of RNA. ATP is often called the "molecular unit of currency" of intracellular energy transfer.[9] ATP transports chemical energy within cells for metabolism. ATP is one of the end products of photophosphorylation, cellular respiration, and fermentation and used by enzymes and structural proteins in many cellular processes, including biosynthetic reactions, motility, and cell division.[10]

The endosymbiotic theory states that several key bacteria that were taken inside another cell as an endosymbiont. As evidence, the mitochondrion has its own independent mitochondrial DNA genome. Further, its DNA shows substantial similarity to bacterial genomes,[36] particularly, molecular and biochemical evidence suggest that the mitochondrion developed from proteobacteria.[37][38]

Molecular and biochemical evidence suggest that the mitochondrion developed from proteobacteria

Endosymbiotic theory

Protocell research has created controversy and opposing opinions, including critics of the vague definition of "artificial life".[31] The creation of a basic unit of life is the most pressing ethical concern, although the most widespread worry about protocells is their potential threat to human health and the environment through uncontrolled replication.[32]

Ethics and controversy

In a similar synthesis experiment using light, led by Jason Dworkin in 2000,[29] he exposed a frozen mixture of water, micelles when immersed in water. Dworkin considered these globules to resemble cell membranes that enclose and concentrate the chemistry of life, separating their interior from the outside world. The globules were between 10 to 40 micrometres (0.00039 to 0.00157 in), or about the size of red blood cells. Remarkably, the globules fluoresced, or glowed, when exposed to UV light. Absorbing UV and converting it into visible light in this way was considered one possible way of providing energy to a primitive cell. If such globules played a role in the origin of life, the fluorescence could have been a precursor to primitive photosynthesis. Such fluorescence also provides the benefit of acting as a sunscreen, diffusing any damage that otherwise would be inflicted by UV radiation. Such a protective function would have been vital for life on the early Earth, since the ozone layer, which blocks out the sun's most destructive UV rays, did not form until after photosynthetic life began to produce oxygen.[30]

The sunlight, it is still reported to have some metabolic capabilities, the presence of semipermeable membrane, amino acids, phospholipids, carbohydrates and RNA-like molecules.[26][27] However, the nature and properties of the Jeewanu remains to be clarified.[28][26][27]


Starting with a technique commonly used to deposit molecules on a solid surface, Langmuir-Blodgett deposition, scientist are able to assemble phospholipid membranes layer by layer of arbitrary complexity.[24][25] These artificial phospholipid membranes support functional insertion both of purified and of in situ expressed membrane proteins.[25] The technique could help astrobiologists understand how the first living cells originated.[24]

Surfactant molecules arranged on an air – water interface

Langmuir-Blodgett deposition

Artificial models

Some molecules or particles are too large or too hydrophilic to pass through a lipid bilayer, but can be moved across the cell membrane through fusion or budding of vesicles.[23] This may have eventually led to mechanisms that facilitate movement of molecules to the inside (endocytosis) or to release its contents into the extracellular space (exocytosis).


It has been proposed that electroporation resulting from lightning strikes could be a mechanism of natural horizontal gene transfer.[21] Electroporation is the rapid increase in bilayer permeability induced by the application of a large artificial electric field across the membrane. During electroporation in laboratory procedures, the lipid molecules are not chemically altered but simply shift position, opening up a pore (hole) that acts as the conductive pathway through the bilayer as it is filled with water. The mechanism is the creation of nanometer sized water-filled holes in the membrane. Experimentally, electroporation is used to introduce hydrophilic molecules into cells. It is a particularly useful technique for large highly charged molecules such as DNA and RNA, which would never passively diffuse across the hydrophobic bilayer core.[22] Because of this, electroporation is one of the key methods of transfection as well as bacterial transformation.

Instead of the more popular phospholipids of modern cells, the membrane of protocells in the RNA world would be composed of fatty acids,[19] and that such membranes have relatively high permeability to ions and small molecules,[1] such as nucleoside monophosphate (NMP), nucleoside diphosphate (NDP), and nucleoside triphosphatee (NTP), and may withstand millimolar concentrations of Mg2+.[20] Osmotic pressure also plays a significant role in protocell membrane transport.[1]

Schematic showing two possible conformations of the lipids at the edge of a pore. In the top image the lipids have not rearranged, so the pore wall is hydrophobic. In the bottom image some of the lipid heads have bent over, so the pore wall is hydrophilic.

Membrane transport

A team of applied physicists at Harvard's School of Engineering and Applied Sciences say that primitive cells might have formed inside inorganic clay microcompartments, which can provide an ideal container for the synthesis and compartmentalization of complex organic molecules.[18] Clay-armored "bubbles" form naturally when particles of montmorillonite clay collect on the outer surface of air bubbles under water. This creates a semi permeable vesicle from materials that are readily available in the environment. The authors remark that montmorillonite is known to serve as a chemical catalyst, encouraging lipids to form membranes and single nucleotides to join into strands of RNA. Primitive reproduction can be envisioned when the clay bubbles burst, releasing the lipid membrane-bound product into the surrounding medium.[18]

Montmorillonite bubbles

Research has shown that some minerals can catalyze the stepwise formation of hydrocarbon tails of fatty acids from hydrogen and carbon monoxide gases - gases that may have been released from hydrothermal vents or geysers. Fatty acids of various lengths are eventually released into the surrounding water,[14] but vesicle formation requires a higher concentration of fatty acids, so it is suggested that protocell formation started at land-bound hydrothermal vents such as geysers, mud pots, fumaroles and other geothermal features where water evaporates and concentrates the solute.[4][16][17]

In the 1990s biochemist James Ferris of Rensselaer Polytechnic Institute showed that montmorillonite clay can help create RNA chains of as many as 50 nucleotides joined together spontaneously into a single RNA molecule.[4] Then in 2002, Hanczyc, Fujikawa and Szostak discovered that by adding montmorillonite to their solution of fatty acid micelles (lipid spheres), the clay sped up the rate of vesicles formation 100-fold.[4]

Scientists have come to conclude that life began in hydrothermal vents in the deep sea, but a 2012 study led by Armen Mulkidjanian of Germany's University of Osnabrück, suggests that inland pools of condensed and cooled geothermal vapor have the ideal characteristics for the origin of life.[15] The conclusion is based mainly on the chemistry of modern cells, where the cytoplasm is rich in potassium, zinc, manganese, and phosphate ions, which are not widespread in marine environments. Such conditions, the researchers argue, are found only where hot hydrothermal fluid brings the ions to the surface — places such as geysers, mud pots, fumaroles and other geothermal features. Within these fuming and bubbling basins, water laden with zinc and manganese ions could have collected, cooled and condensed in shallow pools.[15]

This fluid lipid bilayer cross section is made up entirely of phosphatidylcholine.

Geothermal ponds and clay

Rather than being made up of phospholipids, however, early membranes may have formed from monolayers or bilayers of fatty acids, which may have formed more readily in a prebiotic environment.[14] Fatty acids have been synthesized in laboratories under a variety of prebiotic conditions and have been found on meteorites, suggesting their natural synthesis in nature.[3]

[13] When

Scheme of a micelle spontaneously formed by phospholipids in an aqueous solution

Vesicles and micelles

A March 2014 study by NASA's Jet Propulsion Laboratory, demonstrated a unique way to study the origins of life: fuel cells.[11] Fuel cells are similar to biological cells in that electrons are also transferred to and from molecules. In both cases, this results in electricity and power. The study states that one important factor was that the Earth provides electrical energy at the seafloor. "This energy could have kick-started life and could have sustained life after it arose. Now, we have a way of testing different materials and environments that could have helped life arise not just on Earth, but possibly on Mars, Europa and other places in the Solar System."[11]

Energy gradient

Researchers Irene A. Chen and Jack W. Szostak (Nobel Prize in Physiology or Medicine 2009) amongst others, demonstrated that simple physicochemical properties of elementary protocells can give rise to essential cellular behaviors, including primitive forms of Darwinian competition and energy storage. Such cooperative interactions between the membrane and encapsulated contents could greatly simplify the transition from replicating molecules to true cells.[3] Furthermore, competition for membrane molecules would favor stabilized membranes, suggesting a selective advantage for the evolution of cross-linked fatty acids and even the phospholipids of today.[3] This micro-encapsulation allowed for metabolism within the membrane, exchange of small molecules and prevention of passage of large substances across it.[8] The main advantages of encapsulation include increased solubility of the cargo and creating energy in the form of chemical gradient. Energy is thus often said to be stored by cells in the structures of molecules of substances such as carbohydrates (including sugars), lipids, and proteins, which release energy when chemically combined with oxygen during cellular respiration.[9][10]


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