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The origin of life is the study of life on Earth by Charles Darwin.

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Origin of life, is the study of how life on Earth might have evolved from non-life sometime between 3.9 and 4.1 billion years ago. Origin of life topic also includes theories and ideas regarding possible extra-planetary or extra-terrestrial origins of life, thought to have possibly occurred over the last 13.7 billion years in the evolution of the known universe since the Big Bang.

Origin of life.
Pre-Cambrian stromatolites in the Siyeh Formation, Glacier National Park. In 2002, William Schopf of UCLA published a controversial paper in the scientific journal Nature arguing that geological formations such as this possess 3.5 billion year old fossilized algae microbes. If true, they would be the earliest known life on earth.

Origin of life studies is a limited field of research despite its profound impact on biology and human understanding of the natural world. Progress in this field is generally slow and sporadic, though it still draws the attention of many due to the eminence of the question being investigated. One plausible reason for the slow rate of progress is that it is difficult to obtain funding for research in this area, since practical commercial applications for the research are difficult to foresee. A few facts give insight into the conditions in which life may have emerged, but the mechanisms by which non-life became life are still elusive.

Given that the origin of life is proposed to have proceeded by spontaneous chemical reaction, the chemistry concerned should presumably be rather robust and therefore relatively easy to repeat. The fact that it is proving very difficult to do so is a major challenge for researchers in this field, suggesting they may be focusing on the wrong locations, and on the wrong initial chemistry.

History of the concept in science of the Origin of life.

In a letter to Joseph Dalton Hooker on February 1, 1871, Charles Darwin made the suggestion that the original spark of life may have begun in a "warm little pond, with all sorts of ammonia and phosphoric salts, lights, heat, electricity, etc. present, [so] that a protein compound was chemically formed ready to undergo still more complex changes". He went on to explain that "at the present day such matter would be instantly devoured or absorbed, which would not have been the case before living creatures were formed." In other words, the presence of life itself prevents the spontaneous generation of simple organic compounds from occurring on Earth today - a circumstance which makes the search for the origin of life dependent on the sterile conditions of the laboratory.

Aleksandr Oparin (right) at the laboratory.

An experimental approach to the question was beyond the scope of laboratory science in Darwin's day, and no real progress was made until 1924 when Aleksandr Ivanovich Oparin demonstrated that it was the presence of atmospheric oxygen and other more sophisticated life-forms that prevented the chain of events that would lead to the evolution of life. In his The Origin of Life on Earth, Oparin argued that a "primeval soup" of organic molecules could be created in an oxygen-less atmosphere through the action of sunlight. These would combine in ever-more complex fashion until they dissolved into a coacervate droplet. These droplets would "grow" by fusion with other droplets, and "reproduce" through fission into daughter droplets, and so have a primitive metabolism in which those factors which promote "cell integrity" survive, those that don't become extinct. Many modern theories of the origin of life still take Oparin's ideas as a starting point.

Current models on the Origin of life.

There is no truly "standard model" of the origin of life. But most currently accepted models build in one way or another upon a number of discoveries about the origin of molecular and cellular components for life, which are listed in a rough order of postulated emergence:

  1. Plausible pre-biotic conditions result in the creation of certain basic small molecules (monomers) of life, such as amino acids. This was demonstrated in the Miller-Urey experiment by Stanley L. Miller and Harold C. Urey in 1953.
  2. Phospholipids (of an appropriate length) can spontaneously form lipid bilayers, a basic component of the cell membrane.
  3. The polymerization of nucleotides into random RNA molecules might have resulted in self-replicating ribozymes (RNA world hypothesis).
  4. Selection pressures for catalytic efficiency and diversity result in ribozymes which catalyse peptidyl transfer (hence formation of small proteins), since oligopeptides complex with RNA to form better catalysts. Thus the first ribosome is born, and protein synthesis becomes more prevalent.
  5. Proteins outcompete ribozymes in catalytic ability, and therefore become the dominant biopolymer. Nucleic acids are restricted to predominantly genomic use.

The origin of the basic biomolecules, while not settled, is less controversial than the significance and order of steps 2 and 3. The basic chemicals from which life was thought to have formed are:

  • methane (CH4),.
  • ammonia (NH3),.
  • water (H2O),.
  • Hydrogen sulfide (H2S),.
  • carbon dioxide (CO2) or Carbon monoxide (CO), and.
  • phosphate (PO43-).

Molecular Oxygen (O2) and Ozone (O3) were either rare or absent.

As of 2007, no one has yet synthesized a "protocell" using basic components which would have the necessary properties of life (the so-called "bottom-up-approach"). Without such a proof-of-principle, explanations have tended to be short on specifics. However, some researchers are working in this field, notably Steen Rasmussen at Los Alamos National Laboratory and Jack Szostak at Harvard University. Others have argued that a "top-down approach" is more feasible. One such approach, attempted by Craig Venter and others at The Institute for Genomic Research, involves engineering existing prokaryotic cells with progressively fewer genes, attempting to discern at which point the most minimal requirements for life were reached. The biologist John Desmond Bernal, coined the term Biopoesis for this process, and suggested that there were a number of clearly defined "stages" that could be recognised in explaining the origin of life.

Stage 1: The origin of biological monomers.
Stage 2: The origin of biological polymers.
Stage 3: The evolution from molecules to cell.

Bernal suggested that Darwinian evolution may have commenced early, some time between Stage 1 and 2.

Origin of organic molecules.

Origin of life.
The Miller-Urey experiment attempted to recreate the chemical conditions of the primitive Earth in the laboratory, and synthesized some of the building blocks of life.

Miller's experiments on the Origin of life.

In 1953 a graduate student, Stanley Miller, and his professor, Harold Urey, performed an experiment that proved organic molecules could have spontaneously formed on Early Earth from inorganic precursors. The now-famous "Miller-Urey experiment" used a highly reduced mixture of gases - methane, ammonia and Hydrogen - to form basic organic monomers, such as amino acids. Whether the mixture of gases used in the Miller-Urey experiment truly reflects the atmospheric content of Early Earth is a controversial topic. Other less reducing gases produce a lower yield and variety. It was once thought that appreciable amounts of molecular oxygen were present in the prebiotic atmosphere, which would have essentially prevented the formation of organic molecules; however, the current scientific consensus is that such was not the case.

In 2006 another experiment showed that a thick organic haze might have blanketed Early Earth. An organic haze can form over a wide range of methane and carbon dioxide concentrations, believed to be present in the atmosphere of Early Earth. After forming, these organic molecules would have floated down all over the Earth, allowing life to flourish globally.

Simple organic molecules are, of course, a long way from a fully functional self-replicating life form. But in an environment with no pre-existing life these molecules may have accumulated and provided a rich environment for chemical evolution ("soup theory"). On the other hand, the spontaneous formation of complex polymers from abiotically generated monomers under these conditions is not at all a straightforward process. Besides the necessary basic organic monomers, compounds that would have prohibited the formation of polymers were formed in high concentration during the experiments.

Other sources of complex molecules have been postulated, including sources of extra-terrestrial stellar or interstellar origin. For example, from spectral analyses, organic molecules are known to be present in comets and meteorites. In 2004, a team detected traces of polycyclic aromatic hydrocarbons (PAH's) in a nebula, the most complex molecule, to that date, found in space. The use of PAH's has also been proposed as a precursor to the RNA world in the PAH world hypothesis.

It can be argued that the most crucial challenge unanswered by this theory is how the relatively simple organic building blocks polymerise and form more complex structures, interacting in consistent ways to form a protocell. For example, in an aqueous environment hydrolysis of oligomers/polymers into their constituent monomers would be favored over the condensation of individual monomers into polymers. Also, the Miller experiment produces many substances that would undergo cross-reactions with the amino acids or terminate the peptide chain.

Recent Experiments mimicking Miller's experiments

More recent experiments by chemist Jeffrey Bada at Scripps Institution of Oceanography, La Jolla, Calif. were similar to those performed by Miller. However Bada noted that in current models of early Earth conditions carbon dioxide and nitrogen create nitrites, which destroy amino acids as fast as they form. However, the early Earth may have had significant amounts of iron and carbonate minerals able to neutralize the effects of the nitrites. When Bada performed the Miller-type experiment with the addition of iron and carbonate minerals, the products were rich in amino acids. This suggests the origin of significant amounts of amino acids may have occurred on Earth even with an atmosphere containing carbon dioxide and nitrogen.

Eigen's hypothesis on the Origin of life.

In the early 1970s a major attack on the problem of the origin of life was organized by a team of scientists gathered around Manfred Eigen of the Max Planck Institute. They tried to examine the transient stages between the molecular chaos in a prebiotic soup and the transient stages of a self replicating hypercycle, between the molecular chaos in a prebiotic soup and simple macromolecular self-reproducing systems.

In a hypercycle, the information storing system (possibly RNA) produces an enzyme, which catalyzes the formation of another information system, in sequence until the product of the last aids in the formation of the first information system. Mathematically treated, hypercycles could create quasispecies, which through natural selection entered into a form of Darwinian evolution. A boost to hypercycle theory was the discovery that RNA, in certain circumstances forms itself into ribozymes, a form of RNA enzyme.

Wächtershäuser's hypothesis on the Origin of life.

Another possible answer to this polymerization conundrum was provided in 1980s by Günter Wächtershäuser, in his iron-sulfur world theory. In this theory, he postulated the evolution of (bio)chemical pathways as fundamentals of the evolution of life. Moreover, he presented a consistent system of tracing today's biochemistry back to ancestral reactions that provide alternative pathways to the synthesis of organic building blocks from simple gaseous compounds.

In contrast to the classical Miller experiments, which depend on external sources of energy (such as simulated lightning or UV irradiation), "Wächtershäuser systems" come with a built-in source of energy, sulfides of iron and other minerals (e.g. pyrite). The energy released from redox reactions of these metal sulfides is not only available for the synthesis of organic molecules, but also for the formation of oligomers and polymers. It is therefore hypothesized that such systems may be able to evolve into autocatalytic sets of self-replicating, metabolically active entities that would predate the life forms known today.

The experiment as performed produced a relatively small yield of dipeptides (0.4% to 12.4%) and a smaller yield of tripeptides (0.10%) and the authors note that: "under these same conditions dipeptides hydrolysed rapidly." Another criticism of the result is that the experiment did not include any organomolecules that would most likely cross-react or chain-terminate (Huber and Wächtershäuser, 1998).

The latest modification of the iron-sulfur-hypothesis was provided by William Martin and Michael Russell in 2002. According to their scenario, the first cellular life forms may have evolved inside so-called black smokers at seafloor spreading zones in the deep sea. These structures consist of microscale caverns that are coated by thin membraneous metal sulfide walls. Therefore, these structures would solve several critical points of the "pure" Wächtershäuser systems at once:

  1. the micro-caverns provide a means of concentrating newly synthesised molecules, thereby increasing the chance of forming oligomers;.
  2. the steep temperature gradients inside a black smoker allow for establishing "optimum zones" of partial reactions in different regions of the black smoker (e.g. monomer synthesis in the hotter, oligomerisation in the colder parts);.
  3. the flow of hydrothermal water through the structure provides a constant source of building blocks and energy (freshly precipitated metal sulfides);.
  4. the model allows for a succession of different steps of cellular evolution (prebiotic chemistry, monomer and oligomer synthesis, peptide and protein synthesis, RNA world, ribonucleoprotein assembly and DNA world) in a single structure, facilitating exchange between all developmental stages;.
  5. synthesis of lipids as a means of "closing" the cells against the environment is not necessary, until basically all cellular functions are developed.

This model locates the "last universal common ancestor" (LUCA) inside a black smoker, rather than assuming the existence of a free-living form of LUCA. The last evolutionary step would be the synthesis of a lipid membrane that finally allows the organisms to leave the microcavern system of the black smokers and start their independent lives. This postulated late acquisition of lipids is consistent with the presence of completely different types of membrane lipids in archaebacteria and eubacteria (plus eukaryotes) with highly similar cellular physiology of all life forms in most other aspects.

Another unsolved issue in chemical evolution is the origin of homochirality, i.e. all monomers having the same "handedness" (amino acids being left handed, and nucleic acid sugars being right handed). Homochirality is essential for the formation of functional ribozymes (and probably proteins too). The origin of homochirality might simply be explained by an initial asymmetry by chance followed by common descent. Work performed in 2003 by scientists at Purdue identified the amino acid serine as being a probable root cause of organic molecules' homochirality. Serine forms particularly strong bonds with amino acids of the same chirality, resulting in a cluster of eight molecules that must be all right-handed or left-handed. This property stands in contrast with other amino acids which are able to form weak bonds with amino acids of opposite chirality. Although the mystery of why left-handed serine became dominant is still unsolved, this result suggests an answer to the question of chiral transmission: how organic molecules of one chirality maintain dominance once asymmetry is established.

From organic molecules to protocells

The question "How do simple organic molecules form a protocell?" is largely unanswered but there are many hypotheses. Some of these postulate the early appearance of nucleic acids ("genes-first") whereas others postulate the evolution of biochemical reactions and pathways first ("metabolism-first"). Recently, trends are emerging to create hybrid models that combine aspects of both.

"Genes first" models: the RNA world.

The RNA world hypothesis suggests that relatively short RNA molecules could have spontaneously formed that were capable of catalyzing their own continuing replication. It is difficult to gauge the probability of this formation. A number of theories of modes of formation have been put forward. Early cell membranes could have formed spontaneously from proteinoids, protein-like molecules that are produced when amino acid solutions are heated - when present at the correct concentration in aqueous solution, these form microspheres which are observed to behave similarly to membrane-enclosed compartments. Other possibilities include systems of chemical reactions taking place within clay substrates or on the surface of pyrite rocks. Factors supportive of an important role for RNA in early life include its ability to replicate (see Spiegelman Monster); its ability to act both to store information and catalyse chemical reactions (as a ribozyme); its many important roles as an intermediate in the expression and maintenance of the genetic information (in the form of DNA) in modern organisms; and the ease of chemical synthesis of at least the components of the molecule under conditions approximating the early Earth.

A number of problems with the RNA world hypothesis remain, particularly the instability of RNA when exposed to ultraviolet light, the difficulty of activating and ligating nucleotides and the lack of available phosphate in solution required to constitute the backbone, and the instability of the base cytosine (which is prone to hydrolysis). Recent experiments also suggest that the original estimates of the size of an RNA molecule capable of self-replication were most probably vast underestimates. More-modern forms of the RNA World theory propose that a simpler molecule was capable of self-replication (that other "World" then evolved over time to produce the RNA World). At this time however, the various hypotheses have incomplete evidence supporting them. Many of them can be simulated and tested in the lab, but a lack of undisturbed sedimentary rock from that early in Earth's history leaves few opportunities to test this hypothesis robustly.

"Metabolism first" models: iron-sulfur world and others

Several models reject the idea of the self-replication of a "naked-gene" and postulate the emergence of a primitive metabolism which could provide an environment for the later emergence of RNA replication.

One of the earliest incarnations of this idea was put forward in 1924 with Alexander Oparin's notion of primitive self-replicating vesicles which predated the discovery of the structure of DNA. More recent variants in the 1980s and 1990s include Günter Wächtershäuser's iron-sulfur world theory and models introduced by Christian de Duve based on the chemistry of thioesters. More abstract and theoretical arguments for the plausibility of the emergence of metabolism without the presence of genes include a mathematical model introduced by Freeman Dyson in the early 1980s and Stuart Kauffman's notion of collectively autocatalytic sets, discussed later in that decade.

However, the idea that a closed metabolic cycle, such as the reductive citric acid cycle, could form spontaneously (proposed by Günter Wächtershäuser) remains unsupported. According to Leslie Orgel, a leader in origin-of-life studies for the past several decades, there is reason to believe the assertion will remain so. In an article entitled "Self-Organizing Biochemical Cycles" (PNAS, vol. 97, no. 23, November 7, 2000, p12503-12507), Orgel summarizes his analysis of the proposal by stating, "There is at present no reason to expect that multistep cycles such as the reductive citric acid cycle will self-organize on the surface of FeS/FeS2 or some other mineral." It is possible that another type of metabolic pathway was used at the beginning of life. For example, instead of the reductive citric acid cycle, the "open" acetyl-CoA pathway (another one of the four recognised ways of carbon dioxide fixation in nature today) would be even more compatible with the idea of self-organisation on a metal sulfide surface. The key enzyme of this pathway, carbon monoxide dehydrogenase/acetyl-CoA synthase harbours mixed nickel-iron-sulfur clusters in its reaction centers and catalyses the formation of acetyl-CoA (which may be regarded as a modern form of acetyl-thiol) in a single step.

Bubble Theory on the Origin of life.

Waves breaking on the shore create a delicate foam composed of bubbles. Winds sweeping across the ocean have a tendency to drive things to shore, much like driftwood collecting on the beach. It is possible that organic molecules were concentrated on the shorelines in much the same way. Shallow coastal waters also tend to be warmer, further concentrating the molecules through evaporation. While bubbles comprised mostly of water burst quickly, oily bubbles happen to be much more stable, lending more time to the particular bubble to perform these crucial experiments.

The phospholipid is a good example of an oily compound believed to have been prevalent in the prebiotic seas. Because phospholipids contain a hydrophilic head on one end, and a hydrophobic tail on the other, they have the spontaneous tendency to form lipid membranes in water. A lipid monolayer bubble can contain only oil, and, therefore, is not conducive to harbouring water-soluble organic molecules. On the other hand, a lipid bilayer bubble can contain water, and is a likely precursor to the modern cell membrane. If a protein came along that increased the integrity of its parent bubble, then that bubble had an advantage, and was placed at the top of the natural selection waiting list. Primitive reproduction can be envisioned when the bubbles burst, releasing the results of the experiment into the surrounding medium. Once enough of the 'right stuff' was released into the medium, the development of the first prokaryotes, eukaryotes, and multicellular organisms could be achieved. This theory is expanded upon in the book, "The Cell: Evolution of the First Organism" by Joseph Panno Ph.D.

Similarly, bubbles formed entirely out of protein-like molecules, called microspheres, will form spontaneously under the right conditions. But they are not a likely precursor to the modern cell membrane, as cell membranes are composed primarily of lipid compounds rather than amino-acid compounds (for types of membrane spheres associated with abiogenesis, see protobionts, micelle, coacervate).

Other models on the Origin of life.

British ethologist Richard Dawkins wrote about autocatalysis as a potential explanation for the origin of life in his 2004 book The Ancestor's Tale. Autocatalysts are substances which catalyze the production of themselves, and therefore have the property of being a simple molecular replicator. In his book, Dawkins cites experiments performed by Julius Rebek and his colleagues at the Scripps Research Institute in California in which they combined amino adenosine and pentafluorophenyl ester with the autocatalyst amino adenosine triacid ester (AATE). One system from the experiment contained variants of AATE which catalysed the synthesis of themselves. This experiment demonstrated the possibility that autocatalysts could exhibit competition within a population of entities with heredity, which could be interpreted as a rudimentary form of natural selection.

Clay theory on the Origin of life.

A theory for the origin of life based on clay was forwarded by Dr A. Graham Cairns-Smith of the University of Glasgow in 1985 and adopted as a plausible illustration by just a handful of other scientists (including Richard Dawkins). Clay theory postulates that complex organic molecules arose gradually on a pre-existing, non-organic replication platform -- silicate crystals in solution. Complexity in companion molecules developed as a function of selection pressures on types of clay crystal is then exapted to serve the replication of organic molecules independently of their silicate "launch stage". It is, truly, "life from a rock."

Cairns-Smith is a staunch critic of other models of chemical evolution (see Genetic Takeover: And the Mineral Origins of Life ISBN 0-521-23312-7). However, he admits, that like many models of the origin of life, his own also has its shortcomings (Horgan 1991).

Peggy Rigou of the National Institute of Agronomic Research (INRA), in Jouy-en-Josas, France reports in the February 11, 2006 edition of Science News that Prions are capable of binding to clay particles and migrate off the particles when the clay becomes negatively charged. While no reference is made in the report to implications for origin-of-life theories, this research may suggest prions as a likely pathway to early reproducing molecules.

"Deep-hot biosphere" model of Gold.

The discovery of nanobes (filamental structures smaller than bacteria containing DNA) in deep rocks, led to a controversial theory put forward by Thomas Gold in the 1990s that life first developed not on the surface of the Earth, but several kilometers below the surface. It is now known that microbial life is plentiful up to five kilometers below the earth's surface in the form of archaea, which are generally considered to have originated either before or around the same time as eubacteria, most of which live on the surface including the oceans. It is claimed that discovery of microbial life below the surface of another body in our Solar System would lend significant credence to this theory. He also noted that a trickle of food from a deep, unreachable, source promotes survival because life arising in a puddle of organic material is likely to consume all of its food and become extinct.

"Primitive" extraterrestrial life.

An alternative to Earthly abiogenesis is the hypothesis that primitive life may have originally formed extraterrestrially, either in space or on a nearby planet (Mars). (Note that exogenesis is related to, but not the same as, the notion of panspermia).

Organic compounds are relatively common in space, especially in the outer solar system where volatiles are not evaporated by solar heating. Comets are encrusted by outer layers of dark material, thought to be a tar-like substance composed of complex organic material formed from simple carbon compounds after reactions initiated mostly by irradiation by ultraviolet light. It is supposed that a rain of material from comets could have brought significant quantities of such complex organic molecules to Earth.

An alternative but related hypothesis, proposed to explain the presence of life on Earth so soon after the planet had cooled down, with apparently very little time for prebiotic evolution, is that life formed first on early Mars. Due to its smaller size Mars cooled before Earth (a difference of hundreds of millions of years), allowing prebiotic processes there while Earth was still too hot. Life was then transported to the cooled Earth when crustal material was blasted off Mars by asteroid and comet impacts. Mars continued to cool faster and eventually became hostile to the continued evolution or even existence of life (it lost its atmosphere due to low volcanism), Earth is following the same fate as Mars, but at a slower rate.

Neither hypothesis actually answers the question of how life first originated, but merely shifts it to another planet or a comet. However, the advantage of an extraterrestrial origin of primitive life is that life is not required to have evolved on each planet it occurs on, but rather in a single location, and then spread about the galaxy to other star systems via cometary and/or meteorite impact. Evidence to support the plausibility of the concept is scant, but it finds support in recent study of Martian meteorites found in Antarctica and in studies of extremophile microbes. Additional support comes from a recent discovery of a bacterial ecosytem whose energy source is radioactivity.

The Lipid World.

There is a theory that ascribes the first self-replicating object to be lipid-like. It is known that phospholipids spontaneously form bilayers in water - the same structure as in cell membranes. Furthermore, these bodies may expand (by insertion of additional phospholipids), and under excessive expansion may undergo spontaneous splitting which preserves the same composition of lipids in the two progenies. The main idea in this theory is that the molecular composition of the lipid bodies is the preliminary way for information storage, and evolution led to the appearance of polymer entities such as RNA or DNA that may store information favorably.

The Polyphosphate Model

The problem with most scenarios of abiogenesis is that the Thermodynamic equilibrium of amino acid versus peptides is in the direction of separate amino acids. What has been missing is some force that drives polymerization. The resolution of this problem may well be in the properties of polyphosphates. Polyphosphates are formed by polymerization of ordinary monophosphate ions PO4-3 by ultraviolet light. Polyphosphates cause polymerization of amino acids into peptides, driving the process against the direction of equilibrium. Ample ultraviolet light must have existed in the early oceans. The key issue seems to be that calcium reacts with soluble phosphate to form insoluble calcium phosphate (apatitie), so some plausible mechanism must be found to keep free calcium ions from solution. Possibly, the answer may be in some stable, non-reactive complex such as calcium citrate.

The Ecopoesis model on the Origin of life.

The Ecopoesis model proposes that the geochemical cycles of biogenic elements, driven by an early oxygen-rich atmosphere, were the basis of a planetary metabolism that preceded and conditioned the gradual evolution of organismal life.

Relevant fields.

  • Astrobiology is a field that may shed light on the nature of life in general, instead of just life as we know it on Earth, and may give clues as to how life originates.
  • complex systems.

References on the Origin of life.

  • Brooks, J; Shaw, G. (1973). Origins and Development of Living Systems.. Academic Press, 359. ISBN 0-12-135740-6.
  • De Duve, Christian (Jan 1996). Vital Dust: The Origin and Evolution of Life on Earth. Basic Books. ISBN 0-465-09045-1.
  • Horgan, J (1991). "In the beginning". Scientific American 264: 100-109. (Cited on p. 108).
  • Huber, C. and Wächterhäuser, G., (1998). "Peptides by activation of amino acids with CO on (Ni,Fe)S surfaces: implications for the origin of life". Science 281: 670-672. (Cited on p. 108).
  • Martin, W. and Russell M.J. (2002). "On the origins of cells: a hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cells". Philosophical Transactions of the Royal Society: Biological sciences 358: 59-85.
  • Russell MJ, Hall AJ, Cairns-Smith AG, Braterman PS (1988). "Submarine hot springs and the origin of life". Nature 336: 117.
  • JW Schopf et al. (2002). "Laser-Raman imagery of Earth's earliest fossils.". Nature 416: 73-76.
  • Maynard Smith, John; Szathmary, Eors (2000-03-16). The Origins of Life: From the Birth of Life to the Origin of Language. Oxford Paperbacks. ISBN 0-19-286209-X.
  • Hazen, Robert M. (Dec 2005). Genesis: The Scientific Quest for Life's Origins. Joseph Henry Press. ISBN 0-309-09432-1.

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