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Origin of Life

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Origin of Life

"Origin of life" redirects here. For non-scientific views on the origins of life, see Creation myth.


Abiogenesis (/ˌb.ɵˈɛnɨsɪs/ [1]) or biopoiesis[2] is the hypothetical natural process by which life arises from simple organic compounds.[3][4][5][6] The earliest life on Earth existed at least 3.5 billion years ago,[7][8][9] during the Eoarchean Era when sufficient crust had solidified following the molten Hadean Eon.

Scientific hypotheses about the origins of life can be divided into a number of categories. Many approaches investigate how self-replicating molecules or their components came into existence. For example, the Miller–Urey experiment and similar experiments demonstrated that most amino acids, often called "the building blocks of life", can be racemically synthesized in conditions thought to be similar to those of the early Earth. Several mechanisms have been investigated, including lightning and radiation. Other approaches ("metabolism first" hypotheses) focus on understanding how catalysis in chemical systems in the early Earth might have provided the precursor molecules necessary for self-replication.

Conceptual history

Spontaneous generation

Belief in the present ongoing spontaneous generation of certain forms of life from non-living matter goes back to Aristotle and ancient Greek philosophy and continued to have support in Western scholarship until the 19th century; this was paired with the belief in heterogenesis, i.e. that one form of life derived from a different form (e.g. bees from flowers).[10] Classical notions of spontaneous generation, which can be considered under the modern term, abiogenesis, held that certain complex, living organisms are generated by decaying organic substances. According to Aristotle, it was a readily observable truth that aphids arise from the dew which falls on plants, flies from putrid matter, mice from dirty hay, crocodiles from rotting logs at the bottom of bodies of water, and so on.[11] In the 17th century, such assumptions started to be questioned. In 1646, Sir Thomas Browne published his Pseudodoxia Epidemica (subtitled Enquiries into Very many Received Tenets, and Commonly Presumed Truths), which was an attack on false beliefs and "vulgar errors." His conclusions were not widely accepted at the time. His contemporary, Alexander Ross wrote: "To question this (i.e., spontaneous generation) is to question reason, sense and experience. If he doubts of this let him go to Egypt, and there he will find the fields swarming with mice, begot of the mud of Nylus, to the great calamity of the inhabitants."[12]

In 1665, Robert Hooke published the first drawings of a microorganism. Hooke was followed in 1676 by Anton van Leeuwenhoek, who drew and described microorganisms that are now thought to have been protozoa and bacteria.[13] Many felt the existence of microorganisms was evidence in support of spontaneous generation, since microorganisms seemed too simplistic for sexual reproduction, and asexual reproduction through cell division had not yet been observed. Van Leeuwenhoek took issue with the ideas common at the time that fleas and lice could spontaneously result from putrefaction, and that frogs could likewise arise from slime. Using a broad range of experiments ranging from sealed and open meat incubation and the close study of insect reproduction, by the 1680s he became convinced that spontaneous generation was incorrect.[14]

The first experimental evidence against spontaneous generation came in 1668 when Francesco Redi showed that no maggots appeared in meat when flies were prevented from laying eggs. It was gradually shown that, at least in the case of all the higher and readily visible organisms, the previous sentiment regarding spontaneous generation was false. The alternative seemed to be biogenesis: that every living thing came from a pre-existing living thing (omne vivum ex ovo, Latin for "every living thing from an egg").

In 1768, Lazzaro Spallanzani demonstrated that microbes were present in the air, and could be killed by boiling. In 1861, Louis Pasteur performed a series of experiments that demonstrated that organisms such as bacteria and fungi do not spontaneously appear in sterile, nutrient-rich media, but only invade them from outside.

Pasteur and Darwin

By the middle of the 19th century, the theory of biogenesis had accumulated so much evidential support, due to the work of Louis Pasteur and others, that the alternative theory of spontaneous generation had been effectively disproven. Pasteur himself remarked, after a definitive finding in 1864, "Never will the doctrine of spontaneous generation recover from the mortal blow struck by this simple experiment."[15][16]

In a letter to Joseph Dalton Hooker on February 1, 1871,[17] Charles Darwin addressed the question, suggesting 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."[18] In other words, the presence of life itself makes the search for the origin of life dependent on the sterile conditions of the laboratory.

"Primordial soup" hypothesis

No new notable research or theory on the subject appeared until 1924, when Alexander Oparin reasoned that atmospheric oxygen prevents the synthesis of certain organic compounds that are necessary building blocks for the evolution of life. In his The Origin of Life,[19][20] Oparin proposed that the "spontaneous generation of life" that had been attacked by Louis Pasteur did in fact occur once, but was now impossible because the conditions found on the early Earth had changed, and preexisting organisms would immediately consume any spontaneously generated organism. Oparin argued that a "primeval soup" of organic molecules could be created in an oxygenless atmosphere through the action of sunlight. These would combine in ever more complex ways until they formed coacervate droplets. 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, and those that do not become extinct. Many modern theories of the origin of life still take Oparin's ideas as a starting point.

Around the same time, J. B. S. Haldane suggested that the Earth's prebiotic oceans—different from their modern counterparts—would have formed a "hot dilute soup" in which organic compounds could have formed. J.D. Bernal, a pioneer in x-ray crystallography, called this idea biopoiesis or biopoesis, the process of living matter evolving from self-replicating but nonliving molecules,[21][22] and proposed that biopoiesis passes through a number of intermediate stages.

One of the most important pieces of experimental support for the "soup" theory came in 1952. A graduate student, Stanley Miller, and his professor, Harold Urey, performed an experiment that demonstrated how organic molecules could have spontaneously formed from inorganic precursors, under conditions like those posited by the Oparin-Haldane Hypothesis. 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.[23] This provided direct experimental support for the second point of the "soup" theory, and it is around the remaining two points of the theory that much of the debate now centers. In the Miller–Urey experiment, a mixture of water, hydrogen, methane, and ammonia was cycled through an apparatus that delivered electrical sparks to the mixture. After one week, it was found that about 10% to 15% of the carbon in the system was now in the form of a racemic mixture of organic compounds, including amino acids, which are the building blocks of proteins.

The underlying hypothesis held by Oparin, Haldane, Bernal, Miller and Urey was that conditions on the primeval Earth favored chemical reactions that synthesized complex organic compounds from simple precursors. A recent reanalysis of the saved vials containing the original extracts that resulted from the Miller and Urey experiments, using current and more advanced analytical equipment and technology, has uncovered more biochemicals than originally discovered in the 1950s. One of the more important findings was 23 amino acids, far more than the five originally discovered.[24]

Complex biological molecules and protocells

Microspheres or protenoids

In trying to uncover the intermediate stages of abiogenesis mentioned by Bernal, Sidney W. Fox in the 1950s and 1960s, studied the spontaneous formation of peptide structures under conditions that might plausibly have existed early in Earth's history. He demonstrated that amino acids could spontaneously form small peptides. In one of his experiments, he allowed amino acids to dry out as if puddled in a warm, dry spot in prebiotic conditions. He found that, as they dried, the amino acids formed long, often cross-linked, thread-like, submicroscopic polypeptide molecules now named "proteinoids microspheres", which show many of the basic characteristics of 'life'.[25]

In another experiment using a similar method to set suitable conditions for life to form, Fox collected volcanic material from a cinder cone in Hawaii. He discovered that the temperature was over 100 °C (212 °F) just 4 inches (100 mm) beneath the surface of the cinder cone, and suggested that this might have been the environment in which life was created—molecules could have formed and then been washed through the loose volcanic ash and into the sea. He placed lumps of lava over amino acids derived from methane, ammonia and water, sterilized all materials, and baked the lava over the amino acids for a few hours in a glass oven. A brown, sticky substance formed over the surface and when the lava was drenched in sterilized water a thick, brown liquid leached out. It turned out that the amino acids had combined to form proteinoids, and the proteinoids had combined to form small, cell-like spheres. Fox called these "microspheres", a name that subsequently was displaced by the more informative term protobionts. His protobionts were not cells, although they formed clumps and chains reminiscent of cyanobacteria. They contained no functional nucleic acids, but split asexually and formed within double membranes that had some attributes suggestive of cell membranes. Based upon such experiments, Professor Colin S. Pittendrigh stated in December 1967 that "laboratories will be creating a living cell within ten years," a remark that reflected the typical contemporary levels of innocence of the complexity of cell structures.[26]

Jeewanu

About the same time as Fox's microspheres, synthesis of very similar microscopic particles called Jeewanu (a Sanskrit for the "particles of life") was reported by Krishna Bahadur and his team from Allahabad University in India. The first synthesis was in 1963, and followed by a series of reports in 1964. Their experiment involved a photochemical reaction of a mixture of inorganic nitrogenous compounds (such as ammonium phosphate and ammonium molybdate), organic compounds (such as formaldehyde), and minerals commonly found in living cells, in a sterile environment. Under the sunlight, the reaction produced particles surrounded by semipermeable membranes. Unlike the microspheres, the particles contained a complex mixture of organic compounds, including amino acids, phospholipids and carbohydrates. They also showed metabolic activities and an ability to divide.[27][28][29]

Early conditions

Main article: Timeline of evolution

The Hadean Earth is thought to have had a secondary atmosphere, formed through degassing of the rocks that accumulated from planetesimal impactors. At first it was thought by scientists like Harold Urey that the earth's atmosphere was made up of hydrides—methane, ammonia and water vapour, and that life began under such reducing conditions, conducive to the formation of organic molecules. However, it is now thought that the early atmosphere, based on today's volcanic evidence, would have contained 60% hydrogen, 20% oxygen (mostly in the form of water vapour), 10% carbon dioxide, 5 to 7% hydrogen sulfide, and smaller amounts of nitrogen, carbon monoxide, free hydrogen, methane and inert gases. As Earth lacked the gravity to hold any molecular hydrogen, this component of the atmosphere was rapidly lost during the Hadean period. Solution of the carbon dioxide in water is thought to have made the seas slightly acidic, with a pH of about 5.5.[30]

Morse and MacKenzie have suggested that oceans may have appeared first in the Hadean eon, as soon as two hundred million years (200 Ma) after the Earth was formed, in a hot 100 °C (212 °F) reducing environment, and that the pH of about 5.8 rose rapidly towards neutral.[31] This has been supported by Wilde[32] who has pushed the date of the zircon crystals found in the metamorphosed quartzite of Mount Narryer in Western Australia, previously thought to be 4.1–4.2 Ga, to 4.404 Ga. This means that oceans and continental crust existed within 150 Ma of Earth's formation. Rosing et al.,[33] suggest that between 4.4 and 4.3 Ga, the Earth was a water world, with little if any continental crust, with an extremely turbulent atmosphere and a hydrosphere subject to high UV, from a T Tauri sun and cosmic radiation and continued bolide impact.

As a result, the Hadean environment was one highly hazardous to modern life. Frequent collisions with large objects, up to 500 kilometres (310 mi) in diameter, would have been sufficient to sterilise the planet and vaporise the ocean within a few months of impact, with hot steam mixed with rock vapour leading to high altitude clouds completely covering the planet. After a few months the height of these clouds would have begun to decrease but the cloud base would still have been elevated for about the next thousand years. After that, it would have begun to rain at low altitude. For another two thousand years rains would slowly have drawn down the height of the clouds, returning the oceans to their original depth only 3,000 years after the impact event.[34]

Between 3.8 and 4.1 Ga, changes in the orbits of the gaseous giant planets may have caused a late heavy bombardment[35] that pockmarked the Moon and the other inner planets (Mercury, Mars, and presumably Earth and Venus). This would likely have repeatedly sterilized the planet, had life appeared before that time. Geologically the Hadean Earth would have been far more active than at any other time in its history. Studies of meteorites suggests that radioactive isotopes such as aluminium-26 with a half-life of 7.17×105 years, and potassium-40 with a half-life of 1.250×109 years, isotopes mainly produced in supernovae, were much more common, with the result that.[36] Coupled with internal heating as a result of gravitational sorting between the core and the mantle generated a great deal of mantle convection, with the probable result that there would have been many more smaller very active tectonic plates, than in modern times.

By examining the time interval between such devastating environmental events, the time interval when life might first have come into existence can be found for different early environments. The study by Maher and Stevenson shows that if the deep marine hydrothermal setting provides a suitable site for the origin of life, abiogenesis could have happened as early as 4.0 to 4.2 Ga, whereas if it occurred at the surface of the Earth abiogenesis could only have occurred between 3.7 and 4.0 Ga.[37]

Other research suggests a colder start to life. Work by Leslie Orgel and colleagues on the synthesis of purines has shown that freezing temperatures are advantageous, due to the concentrating effect for key precursors such as hydrogen cyanide.[38] Research by Stanley Miller and colleagues suggested that while adenine and guanine require freezing conditions for synthesis, cytosine and uracil may require boiling temperatures.[39] Research by the Miller group notes the formation of seven different amino acids and 11 types of nucleobases in ice when ammonia and cyanide were left in a freezer from 1972 to 1997.[40][41] This article also describes research by Christof Biebricher showing the formation of RNA molecules 400 bases long under freezing conditions using an RNA template, a single-strand chain of RNA that guides the formation of a new strand of RNA. As that new RNA strand grows, it adheres to the template.[42] The explanation given for the unusual speed of these reactions at such a low temperature is eutectic freezing. As an ice crystal forms, it stays pure: only molecules of water join the growing crystal, while impurities like salt or cyanide are excluded. These impurities become crowded in microscopic pockets of liquid within the ice, and this crowding causes the molecules to collide more often.

Evidence of the early appearance of life comes from the Isua supercrustal belt in Western Greenland and from similar formations in the nearby Akilia Island. Carbon entering into rock formations has a ratio of carbon-13 (13C) to carbon-12 (12C) of about −5.5 (in units of δ13C), where because of a preferential biotic uptake of 12C, biomass has a δ13C of between −20 and −30. These isotopic fingerprints are preserved in the sediments, and Mojzis has used this technique to suggest that life existed on the planet already by 3.85 billion years ago.[43] Lazcano and Miller (1994) suggest that the rapidity of the evolution of life is dictated by the rate of recirculating water through mid-ocean submarine vents. Complete recirculation takes 10 million years, thus any organic compounds produced by then would be altered or destroyed by temperatures exceeding 300 °C (572 °F). They estimate that the development of a 100 kilobase genome of a DNA/protein primitive heterotroph into a 7000 gene filamentous cyanobacterium would have required only 7 Ma.[44] Chemist Christian de Duve argues that the determination of chemistry means that "life has to emerge quickly ... Chemical reactions happen quickly or not at all; if any reaction takes a millennium to complete then the chances are all the reagents will simply dissipate or break down in the meantime, unless they are replenished by other faster reactions".[45][46]

Current models

There is still no "standard model" of the origin of life. Most currently accepted models draw at least some elements from the framework laid out by the Oparin-Haldane hypothesis. Under that umbrella, however, are a wide array of disparate discoveries and conjectures such as the following, listed in a rough order of postulated emergence:

  • The Oparin-Haldane hypothesis suggests that the atmosphere of the early Earth may have been chemically reducing in nature, composed primarily of methane (CH4), ammonia (NH3), water (H2O), hydrogen sulfide (H2S), carbon dioxide (CO2) or carbon monoxide (CO), and phosphate (PO43-), with molecular oxygen (O2) and ozone (O3) either rare or absent.
  • In such a reducing atmosphere, electrical activity can catalyze 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 reported in 1953.
  • Phospholipids (of an appropriate length) can form lipid bilayers, a basic component of the cell membrane.
  • A fundamental question is about the nature of the first self-replicating molecule. Since replication is accomplished in modern cells through the cooperative action of proteins and nucleic acids, the major schools of thought about how the process originated can be broadly classified as "proteins first" and "nucleic acids first".
  • The principal thrust of the "nucleic acids first" argument is as follows:
    1. The polymerization of nucleotides into random RNA molecules might have resulted in self-replicating ribozymes (RNA world hypothesis)
    2. Selection pressures for catalytic efficiency and diversity might have resulted in ribozymes which catalyse peptidyl transfer (hence formation of small proteins), since oligopeptides complex with RNA to form better catalysts. The first ribosome might have been created by such a process, resulting in more prevalent protein synthesis.
    3. Synthesized proteins might then outcompete ribozymes in catalytic ability, and therefore become the dominant biopolymer, relegating nucleic acids to their modern use, predominantly as a carrier of genomic information.

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 focused on chemosynthesis of polymers. 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, successfully 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.[47][48] The biologist John Desmond Bernal coined the term biopoiesis for this process,[49] 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 evolution commenced between Stage 1 and 2.[50] Such in-vitro evolution of pre-biological polymer catalysts, has been demonstrated by Hiroaki Suga of the University of Buffalo, and supports, he suggests, the RNA World model of abiogenesis (see below).[51]

Origin of organic molecules

There are two possible sources of organic molecules on the early Earth:

  1. Terrestrial origins – organic synthesis driven by impact shocks or by other energy sources (such as ultraviolet light or electrical discharges) (e.g. Miller's experiments)
  2. Extraterrestrial origins – delivery by objects (e.g. carbonaceous chondrites) or gravitational attraction of organic molecules or primitive life-forms from space (see Panspermia)

Recently, estimates of these sources suggest that the heavy bombardment before 3.5 Ga within the early atmosphere made available quantities of organics comparable to those produced by other energy sources.[52][53]

"Soup" theory

Biochemist Robert Shapiro has summarized the "primordial soup" theory of Oparin and Haldane in its "mature form" as follows:[54]

  1. The early Earth had a chemically reducing atmosphere.
  2. This atmosphere, exposed to energy in various forms, produced simple organic compounds ("monomers").
  3. These compounds accumulated in a "soup", which may have been concentrated at various locations (shorelines, oceanic vents etc.).
  4. By further transformation, more complex organic polymers – and ultimately life – developed in the soup.

While steps 1-3 have been basically observed experimentally, step 4 has been criticised as simplistic - a stage of "then magic happens".

Reducing atmosphere

At the time of the Miller–Urey experiment, scientific consensus was that the early earth had a reducing atmosphere with compounds relatively rich in hydrogen and poor in oxygen (e.g., and as opposed to and ). However, current scientific consensus describes the primitive atmosphere as either weakly reducing or neutral[55][56] (see also Oxygen catastrophe). Such an atmosphere would diminish both the amount and variety of amino acids that could be produced. This change in opinion has focused scientific research on two other potential reducing environments: outer space and deep-sea thermal vents.[57][58][59]

Monomer formation

Apart from the Miller–Urey experiment, the next most important step in research on prebiotic organic synthesis was the demonstration by Joan Oró that the nucleic acid purine base, adenine, was formed by heating aqueous ammonium cyanide solutions.[60] In support of abiogenesis in eutectic ice, more recent work demonstrated the formation of s-triazines (alternative nucleobases), pyrimidines (including cytosine and uracil), and adenine from urea solutions subjected to freeze-thaw cycles under a reductive atmosphere (with spark discharges as an energy source).[61]

Monomer accumulation

The "soup" theory relies on the assumption proposed by Darwin that in an environment with no pre-existing life, organic molecules may have accumulated and provided an environment for chemical evolution.

Further transformation

The spontaneous formation of complex polymers from abiotically generated monomers under the conditions posited by the "soup" theory 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 Miller–Urey and Oró experiments.[62] The Miller experiment, for example, produces many substances that would undergo cross-reactions with the amino acids or terminate the peptide chain.[63]

More fundamentally, 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.

Eigen's hypothesis

In the early 1970s, the problem of the origin of life was approached by Manfred Eigen and Peter Schuster of the Max Planck Institute for Biophysical Chemistry. They examined the transient stages between the molecular chaos and a self-replicating hypercycle in a prebiotic soup.[64]

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, capable of catalyzing their own chemical reactions.[65] However, these reactions are limited to self-excisions (in which a longer RNA molecule becomes shorter), and much rarer small additions that are incapable of coding for any useful protein. The hypercycle theory is further degraded since the hypothetical RNA would require the existence of complex biochemicals such as nucleotides which are not formed under the conditions proposed by the Miller–Urey experiment.

Sol Spiegelman aimed to find the simplest life form by taking advantage of evolution's natural selection process. His new life form, Spiegelman's Monster, had a genome with just 218 bases. Manfred Eigen built on Spiegelman's work and produced a life form with just 48 or 54 nucleotides.[66] M. Sumper and R. Luce of Eigen's laboratory accidentally discovered that a mixture containing no RNA at all but only RNA bases and Q-Beta Replicase can, under the right conditions, spontaneously generate self-replicating RNA which evolves into a form similar to Spiegelman's Monster.[67]

Hoffmann's contributions

Geoffrey W. Hoffmann, a student of Eigen, contributed to the concept of life involving both replication and metabolism emerging from catalytic noise. His contributions included showing that an early sloppy translation machinery can be stable against an error catastrophe of the type that had been envisaged as problematical by Leslie Orgel ("Orgel's paradox")[68][69] and calculations regarding the occurrence of a set of required catalytic activities together with the exclusion of catalytic activities that would be disruptive. This is called the stochastic theory of the origin of life.[70]

Wächtershäuser's hypothesis

Another possible answer to this polymerization conundrum was provided in the 1980s by the German chemist Günter Wächtershäuser, encouraged and supported by Karl R. Popper,[71][72][73] 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 ultraviolet 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 produced a relatively small yield of dipeptides (0.4% to 12.4%) and a smaller yield of tripeptides (0.10%) but the authors also noted that: "under these same conditions dipeptides hydrolysed rapidly."[74]

Zn-World hypothesis

The Zn-World (zinc world) theory of Armen Mulkidjanian [75] is an extension of Wächtershäuser's pyrite hypothesis. Wächtershäuser based his theory of the initial chemical processes leading to informational molecules (i.e. RNA, peptides) on a regular mesh of electric charges at the surface of pyrite that may have made the primeval polymerization thermodynamically more favourable by attracting reactants and arranging them appropriately relative to each other.[76] The Zn-World theory specifies and differentiates further.[75][77] Hydrothermal fluids rich in H2S interacting with cold primordial ocean (or "Darwin pond") water leads to the precipitation of metal sulfide particles. Oceanic vent systems and other hydrothermal systems have a zonal structure reflected in ancient volcanogenic massive sulfide deposits (VMS) of hydrothermal origin. They reach many kilometers in diameter and date back to the Archean eon. Most abundant are pyrite (FeS2), chalcopyrite (CuFeS2), and sphalerite (ZnS), with additions of galena (PbS) and alabandite (MnS). ZnS and MnS have a unique ability to store radiation energy, e.g. provided by UV light. Since during the relevant time window of the origins of replicating molecules the primordial atmospheric pressure was high enough (> 100 bar) to precipitate ZnS near the earth's surface and UV irradiation was 10 to 100 times more intense than now, the unique photosynthetic properties mediated by ZnS provided just the right energy conditions to energize the synthesis of informational and metabolic molecules and the selection of photostable nucleobases.

The Zn-World theory has been further filled out with experimental and theoretical evidence for the ionic constitution of the interior of the first proto-cells before Archea, Eubacteria and Proto-Eukarya evolved. Archibald Maccallum noted the resemblance of organismal fluids such as blood, lymph to seawater;[78] however, the inorganic composition of all cells differ from that of modern sea water, which led Mulkidjanian and colleagues to reconstruct the "hatcheries" of the first cells combining geochemical analysis with phylogenomic scrutiny of the inorganic ion requirements of universal components of modern cells. The authors conclude that ubiquitous, and by inference primordial, proteins and functional systems show affinity to and functional requirement for K+, Zn2+, Mn2+, and phosphate. Geochemical reconstruction shows that the ionic composition conducive to the origin of cells could not have existed in what we today call marine settings but is compatible with emissions of vapor-dominated zones of what we today call inland geothermal systems. Under the anoxic, CO2-dominated primordial atmosphere, the chemistry of water condensates and exhalations near geothermal fields would resemble the internal milieu of modern cells. Therefore, the precellular stages of evolution may have taken place in shallow "Darwin-ponds" lined with porous silicate minerals mixed with metal sulfides and enriched in K+, Zn2+, and phosphorus compounds.[79][80]

Radioactive beach hypothesis

Zachary Adam at the University of Washington, Seattle, claims that tidal processes that occurred during a time when the moon was much closer may have concentrated grains of uranium and other radioactive elements at the high-water mark on primordial beaches, where they may have been responsible for generating life's building blocks.[81] According to computer models reported in Astrobiology,[82] a deposit of such radioactive materials could show the same self-sustaining nuclear reaction as that found in the Oklo uranium ore seam in Gabon. Such radioactive beach sand might have provided sufficient energy to generate organic molecules, such as amino acids and sugars from acetonitrile in water. Radioactive monazite also have released soluble phosphate into the regions between sand-grains, making it biologically "accessible". Thus amino acids, sugars, and soluble phosphates might have been produced simultaneously, according to Adam. Radioactive actinides, left behind in some concentration by the reaction, might have formed part of organo-metallic complexes. These complexes could have been important early catalysts to living processes.

John Parnell of the University of Aberdeen has suggested that such a process could provide part of the "crucible of life" in the early stages of any early wet rocky planet, so long as the planet is large enough to have generated a system of plate tectonics which brings radioactive minerals to the surface. As the early Earth is believed to have had many smaller "platelets" it might have provided a suitable environment for such processes.[83]

Ultraviolet and temperature-assisted replication model

From a thermodynamic perspective of the origin of life springs the ultraviolet and temperature-assisted replication (UVTAR) model. Karo Michaelian of the National Autonomous University of Mexico points out that any model for the origin of life must take into account the fact that life is an irreversible thermodynamic process which arises and persists because it produces entropy. Entropy production is not incidental to the process of life, but rather the fundamental reason for its existence. Present day life augments the entropy production of Earth by catalysing the water cycle through evapotranspiration.[84][85] Michaelian argues that if the thermodynamic function of life today is to produce entropy through coupling with the water cycle, then this probably was its function at its very beginnings. It turns out that both RNA and DNA when in water solution are very strong absorbers and extremely rapid dissipaters of ultraviolet light within the 200–300 nm wavelength range, which is that part of the sun's spectrum that could have penetrated the dense prebiotic atmosphere. Cnossen et al.[86] have shown that the amount of ultraviolet (UV) light reaching the Earth's surface in the Archean eon could have been up to 31 orders of magnitude greater than it is today at 260 nm where RNA and DNA absorb most strongly. Absorption and dissipation of UV light by the organic molecules at the Archean ocean surface would have significantly increased the temperature of the surface skin layer and led to enhanced evaporation and thus to have augmented the primitive water cycle. Since absorption and dissipation of high energy photons is an entropy producing process, Michaelian argues that non-equilbrium abiogenic synthesis of RNA and DNA utilizing UV light[87] would have been thermodynamically favored.

A simple mechanism that could explain the replication of RNA and DNA without resort to the use of enzymes could also be provided within the same thermodynamic framework by assuming that life arose when the temperature of the primitive seas had cooled to somewhat below the denaturing temperature of RNA or DNA (based on the ratio of 18O/16O found in cherts of the Barberton greenstone belt of South Africa of about 3.5 to 3.2 Ga., surface temperatures are predicted to have been around 70±15 °C,[88] close to RNA or DNA denaturing temperatures). During the night, the surface water temperature would drop below the denaturing temperature and single strand RNA/DNA could act as a template for the formation of double strand RNA/DNA. During the daylight hours, RNA and DNA would absorb UV light and convert this directly to heat the ocean surface, thereby raising the local temperature enough to allow for denaturing of RNA and DNA. The copying process would have been repeated with each diurnal cycle.[89][90] Such a temperature assisted mechanism of replication bears similarity to polymerase chain reaction (PCR), a routine laboratory procedure employed to multiply DNA segments. Michaelian suggests that the traditional origin of life research, that expects to describe the emergence of life from near-equilibrium conditions, is erroneous and that non-equilibrium conditions must be considered, in particular, the importance of entropy production to the emergence of life.

Since denaturation would be most probable in the late afternoon when the Archean sea surface temperature would be highest, and since late afternoon submarine sunlight is somewhat circularly polarized, the homochirality of the organic molecules of life can also be explained within the proposed thermodynamic framework.[91][92]

Models to explain homochirality

Main article: Homochirality

Some process in chemical evolution must account for the origin of homochirality, i.e. all building blocks in living organisms having the same "handedness" (amino acids being left-handed, nucleic acid sugars (ribose and deoxyribose) being right-handed, and chiral phosphoglycerides). Chiral molecules can be synthesized, but in the absence of a chiral source or a chiral catalyst, they are formed in a 50/50 mixture of both enantiomers. This is called a racemic mixture. Clark has suggested that homochirality may have started in space, as the studies of the amino acids on the Murchison meteorite showed L-alanine to be more than twice as frequent as its D form, and L-glutamic acid was more than 3 times prevalent than its D counterpart. It is suggested that polarised light has the power to destroy one enantiomer within the proto-planetary disk. Noyes[93] showed that beta decay caused the breakdown of D-leucine, in a racemic mixture, and that the presence of 14C, present in larger amounts in organic chemicals in the early Earth environment, could have been the cause. Robert M. Hazen reports upon experiments conducted in which various chiral crystal surfaces act as sites for possible concentration and assembly of chiral monomer units into macromolecules.[94] Once established, chirality would be selected for.[95] Work with organic compounds found on meteorites tends to suggest that chirality is a characteristic of abiogenic synthesis, as amino acids show a left-handed bias, whereas sugars show a predominantly right-handed bias.[96]

Self-organization and replication

Main article: Self-organization

While features of self-organization and self-replication are often considered the hallmark of living systems, there are many instances of abiotic molecules exhibiting such characteristics under proper conditions. For example Martin and Russell show that physical compartmentation by cell membranes from the environment and self-organization of self-contained redox reactions are the most conserved attributes of living things, and they argue therefore that inorganic matter with such attributes would be life's most likely last common ancestor.[97] Later Palasek showed that self-assembly of RNA molecules can occur spontaneously due to physical factors in hydrothermal vents.[98]

Virus self-assembly within host cells has implications for the study of the origin of life,[99] as it lends further credence to the hypothesis that life could have started as self-assembling organic molecules.[100][101]

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.

Researcher Martin Hanczyc supports the idea of a gradient between life and non-life (i.e. there is no simple line between the two). He thinks that building simple protocells, in the lab, is one of the first steps towards understanding more complex cells, including those that may have later evolved into complex life. Hanczyc says that living cells often consist of somewhere around 1,000,000 types of molecules, whereas his labs are first aiming at creating lifelike systems using around 10 molecules. His protocells display behaviors even simpler than those displayed by things like viruses (e.g. only basic motion, dividing and combining cell walls, and so on).[102] These lifelike behaviors are visible in the short film Protocell Circus by Rachel Armstrong and Michael Simon Toon.[103][104][105]

Deep sea vent hypothesis

The deep sea vent, or alkaline hydrothermal vent, theory for the origin of life on Earth posits that life may have begun at submarine hydrothermal vents, where hydrogen-rich fluids emerge from below the sea floor, as a result of serpentization of ultra mafic olivine with sea water and a pH interface with carbon dioxide-rich ocean water. Sustained chemical energy in such systems is derived from redox reactions, in which electron donors, such as molecular hydrogen, react with electron acceptors, such as carbon dioxide (see iron-sulfur world theory).These are highly exothermic reactions.

Reaction 1a:
Fayalite + water → magnetite + aqueous silica + hydrogen

3Fe2SiO4 + 2H2O → 2Fe3O4 + 3SiO2 + 2H2


Reaction 1b:
Forsterite + aqueous silica → serpentine

3Mg2SiO4 + SiO2 + 4H2O → 2Mg3Si2O5(OH)4


Reaction 1c:
Forsterite + water → serpentine + brucite

2Mg2SiO4 + 3H2O → Mg3Si2O5(OH)4 + Mg(OH)2

Reaction 1c describes the hydration of olivine with water only to yield serpentine and Mg(OH)2 (brucite). Serpentine is stable at high pH in the presence of brucite like calcium silicate hydrate, (C-S-H) phases formed along with portlandite (Ca(OH)2) in hardened Portland cement paste after the hydration of belite (Ca2SiO4), the artificial calcium equivalent of forsterite.

Analogy of reaction 1c with belite hydration in ordinary Portland cement:
Belite + water → C-S-H phase + portlandite

2 Ca2SiO4 + 4 H2O → 3 CaO · 2 SiO2 · 3 H2O + Ca(OH)2

Mike Russell demonstrated that alkaline vents created an abiogenic proton-motive force chemiosmotic gradient,[106] in which conditions are ideal for an abiogenic hatchery for life. Their microscopic compartments "provide a natural means of concentrating organic molecules", composed of iron-sulfur minerals such as mackinawite, endowed these mineral cells with the catalytic properties envisaged by Günter Wächtershäuser.[107] This movement of ions across the membrane depends on a combination of two factors:

  1. Diffusion force caused by concentration gradient - all particles including ions tend to diffuse from higher concentration to lower.
  2. Electrostatic force caused by electrical potential gradient - cations like protons H+ tend to diffuse down the electrical potential, anions in the opposite direction.

These two gradients taken together can be expressed as an electrochemical gradient, providing energy for abiogenic synthesis. The proton-motive force (PMF) can be described as the measure of the potential energy stored as a combination of proton and voltage gradients across a membrane (differences in proton concentration and electrical potential). In 1978, for the discovery of the PMF, Peter Mitchell was awarded the Nobel Prize in Chemistry.[108]

Coenzyme world

Recent studies, applying the equivalent of Moore's Law to biological evolution and extrapolating backwards, propose that life began "9.7±2.5 billion years ago", billions of years before the Earth was formed.[109][110] In the case of evolution, empirical evidence suggested a doubling of complexity every 376 million years. As the age of trees can be measured by the number of rings, the hypothesis that the age of life could be measured by biological complexity (i.e., the length of functional non-redundant DNA in the genome) was studied.[109][110] If log-transformed complexity is plotted against the time of origin of large evolutionary lineages, then the points fit to a straight line (see figure). The exponential increase in complexity can be explained by a positive self-activating feed back loop.[110] The regression line hits zero (i.e., one nucleotide) at "9.7±2.5 billion years ago".[109] If this model is correct, and since our Solar System is 4.6 billion years old,[111] then life somehow arrived to Earth from older stellar systems. This hypothesis was criticized by Eugene Koonin who suggested that the rates of early biological evolution might have been much faster due to the absence of competition on early Earth.[112] Chris Adami argued that "it is inconceivable that life began with just a few nucleotides" (see discussion[110]). To answer this criticism, Sharov proposed a hypothetical abiogenesis scenario that starts from coenzyme-like molecules that are functionally equivalent to single nucleotides.[113][114]

RNA world

Main article: RNA world hypothesis

The RNA world hypothesis describes an early Earth with self-replicating and catalytic RNA but no DNA or proteins. This has spurred scientists to try to determine if RNA molecules could have spontaneously formed that were capable of catalyzing their own replication.[115][116][117] Evidences suggest chemical conditions (including the presence of boron, molybdenum and oxygen) for initially producing RNA molecules may have been better on the planet Mars than those on the planet Earth.[115][116] If so, life-suitable molecules, originating on Mars, may have later migrated to Earth via panspermia or similar process.[115][116]

A number of hypotheses of modes of formation have been put forward. Early cell membranes could have formed spontaneously from proteinoids, which are protein-like molecules produced when amino acid solutions are heated while in the correct concentration in aqueous solution. These are seen to form micro-spheres which are observed to behave similarly to membrane-enclosed compartments. Other possibilities include systems of chemical reactions that take 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 act both to store information and to catalyze 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 the conditions that approximated the early Earth. Relatively short RNA molecules have been artificially produced in labs, which are capable of replication.[118] Such replicase RNA, which functions as both code and catalyst provides its own template upon which copying can occur. Jack Szostak has shown that certain catalytic RNAs can, indeed, join smaller RNA sequences together, creating the potential, in the right conditions for self-replication. If these conditions were present, Darwinian selection would favour the proliferation of such self-catalysing structures, to which further functionalities could be added.[119] Lincoln and Joyce have identified an RNA enzyme capable of self-sustained replication.[120]

Researchers have pointed out difficulties for the abiotic synthesis of nucleotides from cytosine and uracil.[121] Cytosine has a half-life of 19 days at 100 °C (212 °F) and 17,000 years in freezing water.[122] Larralde et al., say that "the generally accepted prebiotic synthesis of ribose, the formose reaction, yields numerous sugars without any selectivity."[123] and they conclude that their "results suggest that the backbone of the first genetic material could not have contained ribose or other sugars because of their instability." The ester linkage of ribose and phosphoric acid in RNA is known to be prone to hydrolysis.[124]

A slightly different version of the RNA-world hypothesis is that a different type of nucleic acid, such as PNA, TNA or GNA, was the first one to emerge as a self-reproducing molecule, to be replaced by RNA only later.[125][126] Pyrimidine ribonucleosides and their respective nucleotides have been prebiotically synthesised by a sequence of reactions which by-pass the free sugars, and are assembled in a stepwise fashion by going against the dogma that nitrogenous and oxygenous chemistries should be avoided. In a series of publications, The Sutherland Group at the School of Chemistry, University of Manchester have demonstrated high yielding routes to cytidine and uridine ribonucleotides built from small 2 and 3 carbon fragments such as glycolaldehyde, glyceraldehyde or glyceraldehyde-3-phosphate, cyanamide and cyanoacetylene. One of the steps in this sequence allows the isolation of enantiopure ribose aminooxazoline if the enantiomeric excess of glyceraldehyde is 60% or greater.[127] This can be viewed as a prebiotic purification step, where the said compound spontaneously crystallised out from a mixture of the other pentose aminooxazolines. Ribose aminooxazoline can then react with cyanoacetylene in a mild and highly efficient manner to give the alpha cytidine ribonucleotide. Photoanomerization with UV light allows for inversion about the 1' anomeric centre to give the correct beta stereochemistry.[128] In 2009 they showed that the same simple building blocks allow access, via phosphate controlled nucleobase elaboration, to 2',3'-cyclic pyrimidine nucleotides directly, which are known to be able to polymerise into RNA. This paper also highlights the possibility for the photo-sanitization of the pyrimidine-2',3'-cyclic phosphates.[87] James Ferris's studies have shown that clay minerals of montmorillonite will catalyze the formation of RNA in aqueous solution, by joining activated mono RNA nucleotides to join together to form longer chains.[129] Although these chains have random sequences, the possibility that one sequence began to non-randomly increase its frequency by increasing the speed of its catalysis is possible to "kick start" biochemical evolution.

"Life" can be considered to have emerged when RNA chains began to express the basic conditions necessary for natural selection to operate as conceived by Darwin: heritability, variation of type, and competition for limited resources. Fitness of an RNA replicator (its per capita rate of increase) would likely be a function of adaptive capacities that were intrinsic (in the sense that they were determined by the nucleotide sequence) and the availability of resources.[130][131] The three primary adaptive capacities may have been (1) the capacity to replicate with moderate fidelity (giving rise to both heritability and variation of type), (2) the capacity to avoid decay, and (3) the capacity to acquire and process resources.[130][131] These capacities would have been determined initially by the folded configurations of the RNA replicators that, in turn, would be encoded in their individual nucleotide sequences. Competitive success among different replicators would have depended on the relative values of these adaptive capacities.

"Metabolism first" models

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. The centrality of the Krebs cycle to energy production in aerobic organisms, and in drawing in carbon dioxide and hydrogen ions in biosynthesis of complex organic chemicals, including amino acids and nucleotides, suggests that it was one of the first parts of the metabolism to evolve.[132] Harold J. Morowitz concludes that given sufficient concentrations of ingredients the cycle will "spin" of its own, as the concentration of each intermediate rises, it tends to convert into the next intermediate spontaneously. It thus appears to be in origin, not a creation of the genes, but the product of thermodynamics and chemistry alone.[133] Somewhat in agreement with these notions, physicist Sean Carroll has proposed that "the purpose of life is to hydrogenate carbon dioxide" (as part of a "metabolism-first", rather than a "genetics-first", scenario).[134]

Iron-sulfur world

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 debated. In an article entitled "Self-Organizing Biochemical Cycles",[135] the late Leslie Orgel summarized 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 five recognised ways of carbon dioxide fixation in nature today) would be 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.

Thermosynthesis world

Today's bioenergetic process of fermentation is carried out by either the aforementioned citric acid cycle or the Acetyl-CoA pathway, both of which have been connected to the primordial iron-sulfur world. In a different approach, the Thermosynthesis hypothesis considers the bioenergetic process of chemiosmosis, which plays an essential role in cellular respiration and photosynthesis, more basal than fermentation: in Anthonie Muller's "thermosynthesis world" the ATP Synthase enzyme, which sustains chemiosmosis, is proposed as the currently extant enzyme most closely related to the first metabolic process.[136][137]

First life needed an energy source to bring about the condensation reaction that yielded the peptide bonds of proteins and the phosphodiester bonds of RNA. In a generalization and thermal variation of the binding change mechanism of today's ATP Synthase, the "First Protein" would have bound substrates (peptides, phosphate, nucleosides, RNA 'monomers') and condensed them to a reaction product that remained bound until after a temperature change it was released by thermal unfolding.

The energy source of the thermosynthesis world was thermal cycling, the result of suspension of the protocell in a convection current, as is plausible in a volcanic hot spring; the convection accounts for the self-organization and dissipative structure required in any origin of life model. The still ubiquitous role of thermal cycling in germination and cell division is considered a relic of primordial thermosynthesis.

By phosphorylating cell membrane lipids, this 'First Protein' gave a selective advantage to the lipid protocell that contained the protein. In the beginning this First Protein also synthesized a library of many proteins, of which only a minute fraction had thermosynthesis capabilities. Just as proposed by Dyson[138] for the first proteins, the First Protein propagated functionally: it made daughters with similar capabilities, but it did not copy itself. Functioning daughters consisted of different amino acid sequences.

It is assumed RNA sequences were selected among the randomly synthesized RNAs by the relative speed and efficiency increase of First Protein synthesis, for instance by the creation of RNA that functioned as messenger RNA,[139] Transfer RNA[140] and ribosomal RNA, or, even more generally, all the components of the RNA World were also generated and selected. The thermosynthesis world therefore in theory accounts for the origin of the genetic machinery.

Whereas the iron-sulfur world identifies a circular pathway as the most simple—and therefore assumes the existence of enzymes—the thermosynthesis world does not even invoke a pathway, and does not assume the existence of regular enzymes: ATP Synthase's binding change mechanism resembles a physical adsorption process that yields free energy,[141] rather than a regular enzyme's mechanism, which decreases the free energy. The RNA World also implies the existence of several enzymes. It has been claimed that even the emergence of a single enzyme by chance is implausible.[142]

Bubbles

Waves breaking on the shore create a delicate foam composed of bubbles. Winds sweeping across the ocean have a tendency to drive floating surface particles to shore. Possibly such shoreline sea foam and windblown organic particles could interact on the beach. Shallow coastal waters also tend to be warmer, possibly concentrating organic molecules through more rapid evaporation. While bubbles composed mostly of water burst quickly, water containing amphiphiles forms much more stable bubbles, allowing more time for crucial reactions to occur inside them.

Amphiphiles are oily compounds containing a hydrophilic head on one or both ends of a hydrophobic molecule. Some amphiphiles have the tendency to spontaneously form membranes in water. A spherically closed membrane contains water and is a hypothetical precursor to the modern cell membrane. If a protein would increase the integrity of its parent bubble, that bubble had an advantage, and was placed nearer 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.[143]

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).

A recent model by Fernando and Rowe[144] suggests that the enclosure of an autocatalytic non-enzymatic metabolism within protocells may have been one way of avoiding the side-reaction problem that is typical of metabolism first models.

Pumice rafts

An alternative (or perhaps adjunct) theory, to the formation of bubbles via waves breaking on the shore creating delicate foam, is the hypothetical creation of bubbles formed within pores of a pumice raft. Like the windblown foam, the pumice rafts would also have made landfall, and this is observed in modern times. Paleontological evidence of pumice rafts associated with Archean life have been discovered in Australia.[145]

Although the windblown concentration of organic molecules may have been a key part of the abiogenesis puzzle, even with amphiphilic stabilization, exposure to the elements may have rendered the fragile foam too unstable to be an abiogenesis precursor and/or its ongoing natural selection actor.

A possibly more probable bubble formation environment for the 'cradle of life' to occur (due to its greater stability-longer 'lifetime') and optimum size (micron) range would have been the protected environment within the pores of the pumice. The crucial reaction time necessary could have been greatly extended in this protected environment. Relatively rapid selection pressure could have been applied if the pumice raft landed on an active geothermal outgassing site (acting something like an airstone in an aquarium) producing various types of bubbles which interacted and evolved.

Other models

Autocatalysis

In 1993 Stuart Kauffman proposed that life initially arose as autocatalytic chemical networks.[146]

Autocatalysts are substances that catalyze the production of themselves, and therefore have the property of being a simple molecular replicator. British ethologist Richard Dawkins wrote about autocatalysis as a potential explanation for the origin of life in his 2004 book The Ancestor's Tale. 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 esters 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 hypothesis

A model for the origin of life based on clay was forwarded by A. Graham Cairns-Smith of the University of Glasgow in 1985 and explored as a plausible illustration by several scientists.[147] The Clay hypothesis postulates that complex organic molecules arose gradually on a pre-existing, non-organic replication platform of silicate crystals in solution.

Cairns-Smith is a trenchant critic of other models of chemical evolution.[148] However, he admits that like many models of the origin of life, his own also has its shortcomings.

In 2007, Kahr and colleagues reported their experiments that tested the idea that crystals can act as a source of transferable information, using crystals of potassium hydrogen phthalate. "Mother" crystals with imperfections were cleaved and used as seeds to grow "daughter" crystals from solution. They then examined the distribution of imperfections in the new crystals and found that the imperfections in the mother crystals were reproduced in the daughters, but the daughter crystals also had many additional imperfections. For gene-like behavior to be observed, the quantity of inheritance of these imperfections should have exceeded that of the mutations in the successive generations, but it did not. Thus Kahr concluded that the crystals, "were not faithful enough to store and transfer information from one generation to the next".[149][150]

Gold's "deep-hot biosphere" model

In the 1970s, Thomas Gold proposed the theory that life first developed not on the surface of the Earth, but several kilometers below the surface. The discovery in the late 1990s of nanobes (filamental structures that are smaller than bacteria, but that may contain DNA) in deep rocks[151] might be seen as lending support to Gold's theory.

It is now reasonably well established that microbial life is plentiful at shallow depths in the Earth, up to 5 kilometres (3.1 mi) below the surface,[151] in the form of extremophile archaea, rather than the better-known eubacteria (which live in more accessible conditions). 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. Thomas Gold also asserted that a trickle of food from a deep, unreachable, source is needed for survival because life arising in a puddle of organic material is likely to consume all of its food and become extinct. Gold's theory is that the flow of such food is due to out-gassing of primordial methane from the Earth's mantle; more conventional explanations of the food supply of deep microbes (away from sedimentary carbon compounds) is that the organisms subsist on hydrogen released by an interaction between water and (reduced) iron compounds in rocks.

"Primitive" extraterrestrial life

Main article: Panspermia

An alternative to Earthly abiogenesis is the hypothesis that primitive life may have originally formed extraterrestrially (Extraterrestrial life),[109][110] either in space, on Mars or elsewhere. (Note that exogenesis is related to, but not the same as, the notion of panspermia). A supporter of this theory was Francis Crick. Also related, NIH scientists reported studies, based on extrapolating "genetic complexity of organisms to earlier times", suggesting life began "9.7±2.5 billion years ago", billions of years before the Earth was formed. The scientists noted that Life may have started "from systems with single heritable elements that are functionally equivalent to a nucleotide".[109][110] (see Abiogenesis#Coenzyme world and Panspermia#Complexity)

Organic compounds are relatively common in space, especially in the outer solar system where volatiles are not evaporated by solar heating.[152] 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.[153]

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 existence of life; 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 hypothesis is scant, but it finds support in studies of Martian meteorites found in Antarctica and in studies of extremophile microbes.[154] Additional support comes from a recent discovery of a bacterial ecosystem whose energy source is radioactivity.[155]

A 2001 experiment led by Jason Dworkin[156] subjected a frozen mixture of water, methanol, ammonia and carbon monoxide to UV radiation, mimicking conditions found in an extraterrestrial environment. This combination yielded large amounts of organic material that self-organised to form bubbles or micelles when immersed in water. Dworkin considered these bubbles to resemble cell membranes that enclose and concentrate the chemistry of life, separating their interior from the outside world.

The bubbles produced in these experiments were between 10 to 40 micrometres (0.00039 to 0.00157 in), or about the size of red blood cells. Remarkably, the bubbles 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 bubbles 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.[96]

Extraterrestrial organic molecules

Another idea is that amino acids which were formed extraterrestrially arrived on Earth via comets. In 2009 it was announced by NASA that scientists had identified one of the fundamental chemical building blocks of life in a comet for the first time: glycine, an amino acid, was detected in the material ejected from Comet Wild-2 in 2004 and grabbed by NASA's Stardust probe. Tiny grains, just a few thousandths of a millimeter in size, were collected from the comet and returned to Earth in 2006 in a sealed capsule, and distributed among the world's leading astro-biology labs. NASA said in a statement that it took some time for the investigating team, led by Dr Jamie Elsila, to convince itself that the glycine signature found in Stardust's sample bay was genuine and not just Earthly contamination. Glycine has been detected in meteorites before and there are also observations in interstellar gas clouds claimed for telescopes, but the Stardust find is described as a first in cometary material. Isotope analysis indicates that the Late Heavy Bombardment included cometary impacts after the Earth coalesced but before life evolved.[157] Dr. Carl Pilcher, who leads NASA's Astrobiology Institute commented that "The discovery of glycine in a comet supports the idea that the fundamental building blocks of life are prevalent in space, and strengthens the argument that life in the Universe may be common rather than rare."[158]

Based on computer model studies, the complex organic molecules necessary for life may have formed in the protoplanetary disk of dust grains surrounding the Sun before the formation of the Earth.[159] According to the computer studies, this same process may also occur around other stars that acquire planets.[159] (Also see Cosmic dust/Earth.)

Recent observations suggests that the majority of organic compounds introduced on Earth by interstellar dust particles are considered principal agents in the formation of complex molecules, thanks to their peculiar surface-catalytic activities.[160][161] Studies reported in 2008, based on 12C/13C isotopic ratios of organic compounds found in the Murchison meteorite, suggested that the RNA component uracil and related molecules, including xanthine, were formed extraterrestrially.[162][163] On August 8, 2011, a report, based on NASA studies with meteorites found on Earth, was published suggesting DNA components (adenine, guanine and related organic molecules) were made in outer space.[160][164][165][166] More recently, scientists found that the cosmic dust permeating the universe contains complex organic matter ("amorphous organic solids with a mixed aromatic-aliphatic structure") that could be created naturally, and rapidly, by stars.[167][168][169] As one of the scientists noted, "Coal and kerogen are products of life and it took a long time for them to form ... How do stars make such complicated organics under seemingly unfavorable conditions and [do] it so rapidly?"[167] Further, the scientist suggested that these compounds may have been related to the development of life on earth and said that, "If this is the case, life on Earth may have had an easier time getting started as these organics can serve as basic ingredients for life."[167]

In 2000, Jes Jørgensen and Jan M. Hollis from the Copenhagen University, reported that in the star-forming region near the center of our galaxy they found glycolaldehyde – the first evidence of an interstellar sugar molecule.[170] Then, in August 29, 2012, the same team reported the detection of glycolaldehyde in a distant star system. The molecule was found around the protostellar binary IRAS 16293-2422, which is located 400 light years from Earth.[171][172] Glycolaldehyde is needed to form ribonucleic acid, or RNA, which is similar in function to DNA. These findings suggest that complex organic molecules may form in stellar systems prior to the formation of planets, eventually arriving on young planets early in their formation.[173] Because sugars are associated with both metabolism and the genetic code, two of the most basic aspects of life, it is rationalized the discovery of any sugar in space would increase the likelihood that life may exist elsewhere in our galaxy.[170]

On April 3, 2013, NASA reported that complex organic chemicals could arise on Titan, a moon of Saturn, based on studies simulating the atmosphere of Titan.[174]

Lipid world

Main article: Gard model

The lipid world theory postulates that the first self-replicating object was lipid-like.[175][176] It is known that phospholipids form lipid bilayers in water while under agitation – the same structure as in cell membranes. These molecules were not present on early Earth, but other amphiphilic long chain molecules also form membranes. Furthermore, these bodies may expand (by insertion of additional lipids), and under excessive expansion may undergo spontaneous splitting which preserves the same size and 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. Studies on vesicles from potentially prebiotic amphiphiles have so far been limited to systems containing one or two types of amphiphiles. This in contrast to the output of simulated prebiotic chemical reactions, which typically produce very heterogeneous mixtures of compounds.[177] Within the hypothesis of a lipid bilayer membrane composed of a mixture of various distinct amphiphilic compounds there is the opportunity of a huge number of theoretically possible combinations in the arrangements of these amphiphiles in the membrane. Among all these potential combinations, a specific local arrangement of the membrane would have favored the constitution of an hypercycle,[178][179] according to the terminology by Manfred Eigen, actually a positive feedback composed of two mutual catalysts represented by a membrane site and a specific compound trapped in the vesicle. Such site/compound pairs are transmissible to the daughter vesicles leading to the emergence of distinct lineages of vesicles which would have allowed Darwinian natural selection.[180]

Polyphosphates

A problem in 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.[181][182] Polyphosphates are formed by polymerization of ordinary monophosphate ions PO4−3. Several mechanisms for such polymerization have been suggested. Polyphosphates cause polymerization of amino acids into peptides. They are also logical precursors in the synthesis of such key biochemical compounds as ATP. A key issue seems to be that calcium reacts with soluble phosphate to form insoluble calcium phosphate (apatite), so some plausible mechanism must be found to keep calcium ions from causing precipitation of phosphate. There has been much work on this topic over the years, but an interesting new idea is that meteorites may have introduced reactive phosphorus species on the early Earth.[183]

PAH world hypothesis

Main article: PAH world hypothesis

Other sources of complex molecules have been postulated, including extraterrestrial 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 (PAHs) in a nebula.[184] More recently, in 2010, another team also detected PAHs, along with fullerenes (or "buckyballs"), in nebulae.[185] PAHs are the most complex molecules so far found in space. The use of PAHs has also been proposed as a precursor to the RNA world in the PAH world hypothesis. The Spitzer Space Telescope has recently detected a star, HH 46-IR, which is forming by a process similar to that by which the sun formed. In the disk of material surrounding the star, there is a very large range of molecules, including cyanide compounds, hydrocarbons, and carbon monoxide. In September 2012, NASA scientists reported that PAHs, subjected to interstellar medium (ISM) conditions, are transformed, through hydrogenation, oxygenation and hydroxylation, to more complex organics - "a step along the path toward amino acids and nucleotides, the raw materials of proteins and DNA, respectively".[186][187] Further, as a result of these transformations, the PAHs lose their spectroscopic signature which could be one of the reasons "for the lack of PAH detection in interstellar ice grains, particularly the outer regions of cold, dense clouds or the upper molecular layers of protoplanetary disks."[186][187]

Multiple genesis

Different forms of life may have appeared quasi-simultaneously in the early history of Earth.[188] The other forms may be extinct, leaving distinctive fossils through their different biochemistry (e.g., using arsenic instead of phosphorus), survive as extremophiles, or simply be unnoticed through their being analogous to organisms of the current life tree. Hartman[189] for example combines a number of theories together, by proposing that:

The first organisms were self-replicating iron-rich clays which fixed carbon dioxide into oxalic and other dicarboxylic acids. This system of replicating clays and their metabolic phenotype then evolved into the sulfide rich region of the hotspring acquiring the ability to fix nitrogen. Finally phosphate was incorporated into the evolving system which allowed the synthesis of nucleotides and phospholipids. If biosynthesis recapitulates biopoiesis, then the synthesis of amino acids preceded the synthesis of the purine and pyrimidine bases. Furthermore the polymerization of the amino acid thioesters into polypeptides preceded the directed polymerization of amino acid esters by polynucleotides.

Lynn Margulis's endosymbiotic theory suggests that multiple forms of archea entered into symbiotic relationship to form the eukaryotic cell. The horizontal transfer of genetic material between archea promotes such symbiotic relationships, and thus many separate organisms may have contributed to building what has been recognised as the Last Universal Common Ancestor (LUCA) of modern organisms.

See also

References

Further reading

  • Buehler, Lukas K. (2000–2005) The physico-chemical basis of life, http://www.whatislife.com/about.html accessed 27 October 2005.
  • (Cited on p. 108).
  • (Cited on p. 108).
  • Morowitz, Harold J. (1992) "Beginnings of Cellular Life: Metabolism Recapitulates Biogenesis". Yale University Press. ISBN 0-300-05483-1
  • NASA Astrobiology Institute: Earth's Early Environment and Life
  • NASA Specialized Center of Research and Training in Exobiology: Gustaf O. Arrhenius
  • on Major Steps in Cell Evolution freely available.
  • on the Emergence of Life on the Early Earth freely available.

External links

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  • John Maynard Smith
  • "Harvard Team Creates the World's 1st Synthesized Cells"
  • Martin A. Nowak and Hisashi Ohtsuki. Prevolutionary dynamics and the origin of evolution. Proceedings of the National Academy of Sciences 2008
  • "Exploring Life's Origins: a Virtual Exhibit"
  • Stuart A. Kauffman (web archive version as original page no longer accessible)
  • Origins of Life website including papers, resources, by Dr. Michael Russell at the U. of Glasgow
  • Possible Connections Between Interstellar Chemistry and the Origin of Life on the Earth
  • Scientists Find Clues That Life Began in Deep Space—NASA Astrobiology Institute
  • Self-organizing biochemical cycles—by Leslie Orgel
  • How Life Began: New Research Suggests Simple Approach
  • Primordial Soup's On: Scientists Repeat Evolution's Most Famous Experiment–an article in Scientific American. March 28, 2007
  • (textbook)
  • An abiogenesis primer for laymen
  • Evolution before genes. Vera Vasas et al., Biology Direct, 2012, 7:1.
  • Debate bubbles over the origin of life
  • How life began on Earth
  • Early Archean Serpentine mud Volcanoes at Isua, Greenland, as a Niche for Early Life Marie-Laure Pons et al. PNAS
  • The Origins of Life Smithsonian magazine

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