User:Brinaluvsrocks/Abiogenesis

Early universe with first stars[edit]
See also: Chronology of the universe

Soon after the Big Bang, which occurred roughly 14 Gya, the only chemical elements present in the universe were hydrogen, helium, and lithium, the three lightest atoms in the periodic table. '''These elements gradually accreted and began orbiting in disks of gas and dust. Gravitational accretion of material at the hot and dense centers of these protoplantary disks formed stars by the fusion of hydrogen. Early stars were massive and short-lived, producing all the heavier elements through stellar nucleosynthesis. Element formation through stellar nucleosynthesis proceeds to its most stable element Iron-56. Heavier elements were formed during supernovae at the end of a stars lifecycle.''' Carbon, currently the fourth most abundant chemical element in the universe (after hydrogen, helium, and oxygen), was formed mainly in white dwarf stars. As these stars reached the end of their lifecycles, they ejected these heavier elements, among them carbon and oxygen, throughout the universe. According to the nebular hypothesis, the formation and evolution of the Solar System began 4.6 Gya with the gravitational collapse of a small part of a giant molecular cloud. Most of the collapsing mass collected in the center, forming the Sun, while the rest flattened into a protoplanetary disk out of which the planets, moons, asteroids, and other small Solar System bodies formed.

Emergence of a Habitable Earth[edit]
See also: Geological history of Earth, Circumstellar habitable zone, and Prebiotic atmosphere

The age of the Earth is 4.54 Gya as found by radiometric dating of calcium-aluminium-rich inclusions in carbonaceous chrondrite meteorites, the oldest material in the Solar System. The Hadean Earth (formation to 4 Ga) was at first inhospitable to any living organisms. During its formation, the Earth lost a significant part of its initial mass, and consequentially lacked the gravity to hold molecular hydrogen and the bulk of the original inert gases. '''Soon after initial accretion of Earth at 4.48 Ga, its collision with Theia, a hypothesised impactor, is thought to have created the ejected debris that would eventually form the Moon. This impact would have removed the Earth's primary atmosphere, leaving behind clouds of viscous silicates and carbon dioxide. This unstable atmosphere was short-lived and condensed shortly after to form the bulk silicate Earth, leaving behind an atmosphere largely consisting of water vapor, nitrogen, and carbon dioxide, with smaller amounts of carbon monoxide, hydrogen, and sulfur compounds.'''

'''Condensation to form liquid oceans is theorised to have occurred as early as the Moon-forming impact. ''' The solution of carbon dioxide in water is thought to have made the seas slightly acidic, with a pH of about 5.5. 'This scenario has found support from the dating of 4.404 Gya zircon crystals with high δ''18O values from metamorphosed quartzite of Mount Narryer in Western Australia. ''' The Hadean atmosphere has been characterized as a "gigantic, productive outdoor chemical laboratory," similar to volcanic gases today which still support some abiotic chemistry. '''Despite the likely increased volcanism from early plate tectonics, the Earth may have been a predominantly water world between 4.4 and 4.3 Gya. It is debated whether or not crust was exposed above this ocean due to uncertainties of what early plate tectonics looked like. For early life to have developed, it is generally thought that a land setting is required, so this question is essential to determining when in Earth's history life evolved.''' The post-Moon-forming impact Earth likely existed with little if any continental crust, a turbulent atmosphere, and a hydrosphere subject to intense ultraviolet light from a T Tauri stage Sun, from cosmic radiation, and from continued asteroid and comet impacts. Despite all this, niche environments likely existed conducive to life on Earth in the Late-Hadean to Early-Archaean.

The Late Heavy Bombardment hypothesis posits that a period of intense impact occurred at ~3.9 Ga during the Hadean. ''' A cataclysmic impact event would have had the potential to sterilise all life on Earth by volatilising liquid oceans and blocking the Sun needed for photosynthesising primary producers, pushing back the earliest possible emergence of life to after Late Heavy Bombardment. Recent research questions both the intensity of the Late Heavy Bombardment as well as its potential for sterilisation. Uncertainties as to whether Late Heavy Bombardment was one giant impact or a period of greater impact rates greatly changed the implication of its destructive power.  The 3.9 Ga date arises from dating of Apollo return mission samples collected mostly near the Imbrium Basin, biasing the age of recorded impacts. Impact modelling of the lunar surface reveals that rather than a cataclysmic event at 3.9 Ga, multiple small-scale, short-lived periods of bombardment likely occurred. Terrestrial data backs this idea by showing multiple periods of ejecta in the rock record both before and after the 3.9 Ga marker, suggesting that the early Earth was subject to continuous impacts that would not have had as great an impact on extinction as previously thought. If the Late Heavy Bombardment didn't take place, this allows for the emergence of life to have taken place far before 3.9 Ga.'''

Earliest evidence of life[edit]
Main article: Earliest known life forms

The exact timing at which life emerged on Earth is unknown. '''Minimum age estimates are based on evidence from the geologic rock record. The earliest physical evidence of life so far found consists of microbialites in the Nuvvuagittuq Greenstone Belt of Northern Quebec, in banded iron formation rocks at least 3.77 and possibly as old as 4.32 Gya'''. The micro-organisms lived within hydrothermal vent precipitates, soon after the 4.4 Gya formation of oceans during the Hadean. The microbes resembled modern hydrothermal vent bacteria, supporting the view that abiogenesis began in such an environment.

Biogenic graphite has been found in 3.7 Gya metasedimentary rocks from southwestern Greenland and in microbial mat fossils from 3.49 Gya sandstones in the Pilbara region of Western Australian. Evidence of early life in rocks from Akilia Island, near the Isua supracrustal belt in southwestern Greenland, dating to 3.7 Gya, have shown biogenic carbon isotopes. In other parts of the Isua supracrustal belt, graphite inclusions trapped within garnet crystals are connected to the other elements of life: oxygen, nitrogen, and possibly phosphorus in the form of phosphate, providing further evidence for life 3.7 Gya. In the Pilbara region, compelling evidence of early life was found in pyrite-bearing sandstone in a fossilized beach, with rounded tubular cells that oxidized sulfur by photosynthesis in the absence of oxygen. '''Carbon isotope ratios on graphite inclusions from the Jack Hills zircons suggest that life could have existed on Earth from 4.1 Gya. '''

The Pilbara region of Western Australia contains the Dresser Formation with rocks 3.48 Gya, including layered structures called stromatolites. Their modern counterparts are created by photosynthetic micro-organisms including cyanobacteria. These lie within undeformed hydrothermal-sedimentary strata; their texture indicates a biogenic origin. Parts of the Dresser formation preserve hot springs on land, but other regions seem to have been shallow seas. A molecular clock analysis suggests the LUCA emerged prior to the Late Heavy Bombardment (3.9 Gya).

Suitable geological environments[edit]
Further information: Alternative abiogenesis scenarios

'''A variety of geologic and environmental settings have been proposed for an origin of life. These theories are often in competition with one another as there are many differing views of prebiotic compound availability, geophysical setting, and early life characteristics. The first organism on Earth likely looked different from LUCA. Between the first appearance of life and where all modern phylogenies began branching, an unknown amount of time passed. Within this time, unknown gene transfer, extinctions, and evolutions may have occurred, creating adaptations for a more diverse amount of environmental niches. One notable shifts is the one from an RNA-based to an RNA-DNA-protein world. There is more known about LUCA than the first organisms due to genomic reconstructions based on modern phylogenies. '''

'''The most popular hypotheses for settings in which an origin of life are possible are deep sea hydrothermal vents and surface bodies of water. Surface waters can be broken down into hot springs, moderate temperature lakes and ponds, and cold settings.'''

Deep sea hydrothermal vents[edit]
Further information: Hydrothermal vent and Chemiosmosis § Emergence of chemiosmosis

Early micro-fossils may have come from a hot world of gases such as methane, ammonia, carbon dioxide, and hydrogen sulfide, toxic to much current life. Analysis of the tree of life places thermophilic and hyperthermophilic bacteria and archaea closest to the root, suggesting that life may have evolved in a hot environment. The deep sea or alkaline hydrothermal vent theory posits that life began at submarine hydrothermal vents. William Martin and Michael Russell have suggested "that life evolved in structured iron monosulphide precipitates in a seepage site hydrothermal mound at a redox, pH, and temperature gradient between sulphide-rich hydrothermal fluid and iron(II)-containing waters of the Hadean ocean floor. The naturally arising, three-dimensional compartmentation observed within fossilized seepage-site metal sulphide precipitates indicates that these inorganic compartments were the precursors of cell walls and membranes found in free-living prokaryotes. The known capability of FeS and NiS to catalyze the synthesis of the acetyl-methylsulphide from carbon monoxide and methylsulphide, constituents of hydrothermal fluid, indicates that pre-biotic syntheses occurred at the inner surfaces of these metal-sulphide-walled compartments".

These form where hydrogen-rich fluids emerge from below the sea floor, as a result of serpentinization of ultra-mafic olivine with seawater and a pH interface with carbon dioxide-rich ocean water. The vents form a sustained chemical energy source derived from redox reactions, in which electron donors (molecular hydrogen) react with electron acceptors (carbon dioxide); see iron–sulfur world theory. These are exothermic reactions.

Russell demonstrated that alkaline vents created an abiogenic proton motive force chemiosmotic gradient, ideal for abiogenesis. 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. 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 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).

The surfaces of mineral particles inside deep-ocean hydrothermal vents have catalytic properties similar to those of enzymes and can create simple organic molecules, such as methanol (CH3OH) and formic, acetic, and pyruvic acids out of the dissolved CO2 in the water, if driven by an applied voltage or by reaction with H2 or H2S.

The research reported by Martin in 2016 supports the thesis that life arose at hydrothermal vents, that spontaneous chemistry in the Earth's crust driven by rock–water interactions at disequilibrium thermodynamically underpinned life's origin and that the founding lineages of the archaea and bacteria were H2-dependent autotrophs that used CO2 as their terminal acceptor in energy metabolism. Martin suggests, based upon this evidence, that the LUCA "may have depended heavily on the geothermal energy of the vent to survive". Pores at deep sea hydrothermal vents are suggested to have been occupied by membrane-bound compartments which promoted biochemical reactions. Metabolic intermediates in the Krebs cycle, gluconeogenesis, amino acid bio-synthetic pathways, glycolysis, the pentose phosphate pathway, and including sugars like ribose, and lipid precursors can occur non-enzymatically at conditions relevant to deep-sea alkaline hydrothermal vents.

'''If the deep marine hydrothermal setting was the site for the origin of life, then abiogenesis could have happened as early as 4.0-4.2 Gya. If life evolved in the ocean at depths of more than ten meters, it would have been shielded both from impacts and the then high levels of ultraviolet radiation from the sun. The available energy in hydrothermal vents is maximized at 100–150 °C, the temperatures at which hyperthermophilic bacteria and thermoacidophilic archaea live. Arguments against a hydrothermal origin of life state that hyperthermophily was a result of convergent evolution in bacteria and archaea, and that a mesophilic environment would have been more likely.  This hypothesis, suggested in 1999 by Galtier, was proposed one year before the discovery of the Lost City Hydrothermal Field, where white-smoker hydrothermal vents average ~45-90°C. Moderate temperatures and alkaline seawater at Lost City are now the favoured hydrothermal vent setting in contrast to acidic, high temperature (~350°C) black-smokers.'''

'''Production of prebiotic organic compounds at hydrothermal vents is estimated to be 1x108 kg yr-1.undefined While a large amounts important prebiotic compounds, such as methane, are found at vents, they are still in lower concentrations as estimates of a Miller-Urey Experiment environment. In the case of methane, the production rate at vents is around 2-4 orders of magnitude lower than predicted amounts in a Miller-Urey Experiment surface atmosphere. '''

'''Other arguments against an oceanic vent setting for the origin of life include the inability to concentrate prebiotic materials due to strong dilution from seawater. This open-system cycles compounds through minerals that make up vents, leaving little residence time to accumulate. All modern cells rely on phosphates and potassium for nucleotide backbone and protein formation respectively, making it likely that the first life forms also shared these functions. These elements were not available in high quantities in the Archaean oceans as both primarily come from the weathering of continental rocks on land, far from vent settings. Submarine hydrothermal vents are not conducive to condensation reactions that are needed for polymerisation to form macromolecules. Wet-dry cycling that allows for condensation reactions occur in terrestrial settings.'''

Iron–sulfur world[edit]
Main article: Iron–sulfur world hypothesis

In the 1980s, Wächtershäuser and Karl Popper postulated the iron–sulfur world hypothesis for the evolution of pre-biotic chemical pathways. It traces today's biochemistry to primordial reactions which synthesize organic building blocks from gases. Wächtershäuser systems have a built-in source of energy: iron sulfides such as pyrite. The energy released by oxidising these metal sulfides can support synthesis of organic molecules. Such systems may have evolved into autocatalytic sets constituting self-replicating, metabolically active entities predating modern life forms. Experiments with sulfides in an aqueous environment at 100 °C produced a small yield of dipeptides (0.4% to 12.4%) and a smaller yield of tripeptides (0.10%). However, under the same conditions, dipeptides were quickly broken down.

Several models postulate a primitive metabolism, allowing RNA replication to emerge later. The centrality of the Krebs cycle (citric acid cycle) to energy production in aerobic organisms, and in drawing in carbon dioxide and hydrogen ions in biosynthesis of complex organic chemicals, suggests that it was one of the first parts of the metabolism to evolve. Concordantly, geochemists Szostak and Kate Adamala demonstrated that non-enzymatic RNA replication in primitive protocells is only possible in the presence of weak cation chelators like citric acid. This provides further evidence for the central role of citric acid in primordial metabolism. Russell has proposed that "the purpose of life is to hydrogenate carbon dioxide" (as part of a "metabolism-first", rather than a "genetics-first", scenario). The physicist Jeremy England has argued from general thermodynamic considerations that life was inevitable. An early version of this idea was Oparin's 1924 proposal for self-replicating vesicles. In the 1980s and 1990s came Wächtershäuser's iron–sulfur world theory and Christian de Duve's thioester models. More abstract and theoretical arguments for metabolism without genes include Freeman Dyson's mathematical model and Stuart Kauffman's collectively autocatalytic sets in the 1980s. Kauffman's work has been criticized for ignoring the role of energy in driving biochemical reactions in cells.

A multistep biochemical pathway like the Krebs cycle did not just self-organize on the surface of a mineral; it must have been preceded by simpler pathways. The Wood–Ljungdahl pathway is compatible with self-organization on a metal sulfide surface. Its key enzyme unit, carbon monoxide dehydrogenase/acetyl-CoA synthase, contains mixed nickel-iron-sulfur clusters in its reaction centers and catalyzes the formation of acetyl-CoA. However, prebiotic thiolated and thioester compounds are thermodynamically and kinetically unlikely to accumulate in the presumed prebiotic conditions of hydrothermal vents. One possibility is that cysteine and homocysteine may have reacted with nitriles from the Strecker reaction, forming catalytic thiol-rich polypeptides.

It has been suggested that the iron-sulfur world hypothesis and RNA world hypothesis are not mutually exclusive as modern cellular processes do involve both metabolites and genetic molecules.

Zinc world[edit]
Armen Mulkidjanian's zinc world (Zn-world) hypothesis extends Wächtershäuser's pyrite hypothesis. The Zn-world theory proposes that hydrothermal fluids rich in H2S interacting with cold primordial ocean (or Darwin's "warm little pond") water precipitated metal sulfide particles. Oceanic hydrothermal systems have a zonal structure reflected in ancient volcanogenic massive sulfide ore deposits. They reach many kilometers in diameter and date back to the Archean. 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. from ultraviolet light. When replicating molecules were originating, the primordial atmospheric pressure was high enough (>100 bar) to precipitate near the Earth's surface, and ultraviolet irradiation was 10 to 100 times more intense than now; hence the photosynthetic properties mediated by ZnS provided the right energy conditions for the synthesis of informational and metabolic molecules and the selection of photostable nucleobases.

The Zn-world theory has been filled out with evidence for the ionic constitution of the interior of the first protocells. In 1926, the Canadian biochemist Archibald Macallum noted the resemblance of body fluids such as blood and lymph to seawater; however, the inorganic composition of all cells differ from that of modern seawater, 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 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 [PO4]3−. Geochemical reconstruction shows that this ionic composition could not have existed in the ocean but is compatible with inland geothermal systems. In the oxygen-depleted, CO2-dominated primordial atmosphere, the chemistry of water condensates near geothermal fields would resemble the internal milieu of modern cells. Therefore, precellular 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.

Surface bodies of water
'''Surface bodies of water are an appealing hypothesis for an origin of life as they provide environments able to dry out and be rewetted. These continued wet-dry cycles allow for concentration of prebiotic compounds and condensation reactions to polymerise macromolecules. Moreover, lake and ponds on land allow for detrital input from the weathering of continental rocks which contain apatite, the most common source of phosphates needed for nucleotide backbones. The amount of exposed continental crust in the Hadean is an unanswered question, but strong evidence based on models considering early ocean depths and rates of ocean island and continental crust growth make it very plausible that there was exposed land. Another line of evidence for a surface start to life, as opposed to a deep sea setting, is the requirement of UV for organism function. UV is necessary for the formation of the U+C nucleotide base pair by partial hydrolysis and nucleobase loss. Simultaneously, UV can be harmful and sterilising to life, especially for simple early lifeforms with few adaptations to repair radiation damage. Irradiation levels from a young Sun were likely greater, and with no ozone layer harmful shortwave UV rays would reach the surface of Earth. For life to begin, a shielded environment with influx from UV-exposed sources is necessary to both benefit and protect from UV. Shielding under ice, liquid water, mineral surfaces (e.g. clay) or regolith is possible in a range of surface water settings. While deep sea vents may have input from raining down of surface exposed materials, the likelihood of concentration is lessened by how open a system the ocean is.'''

Hot springs[edit]
'''Most branching phylogenies are thermophilic or hyperthermophilic, making it possible that the Last Universal Common Ancestor (LUCA) and preceding lifeforms were too. Hot springs are formed from the heating of groundwater by geothermal activity. This unique intersection allows for influxes of material from deep penetrating waters and from surface runoff that transports eroded continental sediments. Interconnected groundwater systems create a mechanism for distribution of life to wider area.'''

Mulkidjanian and co-authors think that marine environments did not provide the ionic balance and composition universally found in cells, or the ions required by essential proteins and ribozymes, especially with respect to high K+/Na+ ratio, Mn2+, Zn2+ and phosphate concentrations. They argue that the only environments that mimic the needed conditions on Earth are hot springs similar to ones at Kamchatka. Mineral deposits in these environments under an anoxic atmosphere would have suitable pH (while current pools in an oxygenated atmosphere would not), contain precipitates of photocatalytic sulfide minerals that absorb harmful ultraviolet radiation, have wet-dry cycles that concentrate substrate solutions to concentrations amenable to spontaneous formation of biopolymers created both by chemical reactions in the hydrothermal environment, and by exposure to UV light during transport from vents to adjacent pools that would promote the formation of biomolecules. The hypothesized pre-biotic environments are similar to hydrothermal vents, with additional components that help explain peculiarities of the LUCA.

A phylogenomic and geochemical analysis of proteins plausibly traced to the LUCA shows that the ionic composition of its intracellular fluid is identical to hot springs. The LUCA likely was dependent upon synthesized organic matter for its growth. Experiments show that RNA-like polymers can be synthesized in wet-dry cycling and UV light exposure. These polymers were encapsulated in vesicles after condensation, which would not happen in saltwater because of the high concentrations of ionic solutes. Potential sources of organics at hot springs might have been transport by interplanetary dust particles, extraterrestrial projectiles, or atmospheric or geochemical synthesis. Hot springs could have been abundant at volcanic landmasses during the Hadean.

Temperate surface bodies of water
'''A mesophilic start in surface bodies of waters hypothesis has evolved from Darwin's concept of a 'warm little pond' and the Oparin-Haldane hypothesis. Freshwater bodies under temperate climates can accumulate prebiotic materials while providing suitable environmental conditions conducive to simply life forms. The climate during the Archaean is still a highly debated topic, as there is uncertainty about what continents, oceans, and the atmosphere looked like then. Atmospheric reconstructions of the Archaean from geochemical proxies and models state that sufficient greenhouse gases were present to maintain surface temperatures between 0-40°C. Under this assumption, there are a greater abundance of moderate temperature niches in which life could begin.'''

'''Strong lines of evidence for mesophily from biomolecular studies include Galtier's G+C nucleotide thermometer. G+C are more abundant in thermophiles due to the added stability of an additional hydrogen bond not present between A+T nucleotides. rRNA sequencing on a diverse range of modern lifeforms show that LUCA's reconstructed G+C content was likely representative of moderate temperatures.'''

Although most modern phylogenies are thermophilic or hyperthermophilic, it is possible that their widespread diversity today is a product of convergent evolution and horizontal gene transfer rather than an inherited trait from LUCA. '''The reverse gyrase topoisomerase is found exclusive in thermophiles and hyperthermophiles as it allows for coiling of DNA. The reverse gyrase enzyme requires ATP to function, both of which are complex biomolecules. If an origin of life is hypothesised to be a simple organism that had not yet evolved a membrane let alone ATP, this would make the existence of reverse gyrase improbable. Moreover, phylogenetic studies has revealed that reverse gyrase has an archaeal origin that was transferred to bacterial organisms via horizontal gene transfer. This disputes the idea that reverse gyrase was present in LUCA.'''

Clay[edit]
The clay hypothesis was proposed by Graham Cairns-Smith in 1985. It postulates that complex organic molecules arose gradually on pre-existing, non-organic replication surfaces of silicate crystals in contact with an aqueous solution. The clay mineral montmorillonite has been shown to catalyze the polymerization of RNA in aqueous solution from nucleotide monomers, and the formation of membranes from lipids. In 1998, Hyman Hartman proposed that "the first organisms were self-replicating iron-rich clays which fixed carbon dioxide into oxalic acid and other dicarboxylic acids. This system of replicating clays and their metabolic phenotype then evolved into the sulfide rich region of the hot spring acquiring the ability to fix nitrogen. Finally phosphate was incorporated into the evolving system which allowed the synthesis of nucleotides and phospholipids."

Icy surface bodies of water
'''Cold-start origin of life theories stem from the idea there may have been cold enough regions on the early Earth that large ice cover could be found. Stellar evolution models predict that the Sun's luminosity was ~25% weaker than it is today. Fuelner states that although this significant decrease in solar energy would have formed an icy planet, there is strong evidence for liquid water to be present, possibly driven by a greenhouse effect. This would create an early Earth with both liquid oceans and icy poles.'''

'''Ice melts that form from ice sheets or glaciers melts create freshwater pools, another niche capable of experiencing wet-dry cycles. While these pools that exist on the surface would be exposed to intense UV radiation, bodies of water within and under ice are sufficiently shielded while remaining connected to UV exposed areas through ice cracks. Suggestions of impact melting of ice allow freshwater paired with meteoritic input, a popular vessel for prebiotic components. Near-seawater levels of sodium chloride are found to unstabilize fatty acid membrane self-assembly, making freshwater settings appealing for early membranous life. '''

'''Icy environments would trade the faster reaction rates that occur in warm environments for increased stability and accumulation of larger polymers. Experiments simulating Europa-like conditions of ~20°C have synthesised amino acids and adenine, showing that Miller-Urey type syntheses can still occur at cold temperatures.  In an RNA-world, the ribozyme would have had even more functions than in a later DNA-RNA-protein-world. For RNA to function they must be able to fold, a process that is hindered temperatures >30°C. While RNA folding in psychrophilic organisms is slower, there is a greater success of the process due to slower rates of hydrolysis. Shorter nucleotides would also not suffer from higher temperatures.'''