Special essay. The seven pillars of life.
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For DNA to be used as an informational molecule it must exist in the cell on the edge of stability because all genomic processes require local controlled melting. This presents mechanistic opportunities and problems for genomic DNA from hyperthermophilic organisms, whose unpackaged DNA could melt at optimal temperatures for growth. Hyperthermophiles are suggested to employ the novel positively supercoiling topoisomerase enzyme reverse gyrase (RG) to form positively supercoiled DNA that is intrinsically resistant to thermal denaturation. RG is presently the only archaeal gene that is uniquely found in hyperthermophiles and therefore is central to hypotheses suggesting a hypothermophilic origin of life. However, the suggestion that RG has evolved by the fusion of two pre-existing enzymes has led to hypotheses for a lower temperature for the origin of life. In addition to the action of topoisomerases, DNA packaging and the intracellular ionic environment can also manipulate DNA topology significantly. In the Euryarchaeota, nucleosomes containing minimal histones can adopt two alternate DNA topologies in a salt-dependent manner. From this we hypothesize that since internal salt concentrations are increased following an increase in temperature, the genomic effects of temperature fluctuations could also be accommodated by changes in nucleosome organization. In addition, stress-induced changes in the nucleoid proteins could also play a role in maintaining the genome in the optimal topological state in changing environments. The function of these systems could therefore be central to temperature adaptation and thus be implicated in origin of life scenarios involving hyperthermophiles.
This review addresses the question of the origin of life, with emphasis on plausible boundary structures that may have initially provided cellular compartmentation. Some form of compartmentation is a necessary prerequisite for maintaining the integrity of interdependent molecular systems that are associated with metabolism, and for permitting variations required for speciation. The fact that lipid-bilayer membranes define boundaries of all contemporary living cells suggests that protocellular compartments were likely to have required similar, self-assembled boundaries. Amphiphiles such as short-chain fatty acids, which were presumably available on the early Earth, can self-assemble into stable vesicles that encapsulate hydrophilic solutes with catalytic activity. Their suspensions in aqueous media have therefore been used to investigate nutrient uptake across simple membranes and encapsulated catalyzed reactions, both of which would be essential processes in protocellular life forms.
The origin of life at its abiotic evolutionary stage, requires a combination of constituents and environmental conditions that enable the synthesis of complex replicating macromolecules from simpler monomeric molecules. It is very likely that the early stages of this evolutionary process have been spontaneous, rapid and widespread on the surface of the primitive Earth, resulting in the formation of quite sophisticated living organisms within less than a billion years. To what extent did such conditions prevail on Mars? Two companion-papers (Life on Mars? I and II) will review and discuss the available information related to the chemical, physical and environmental conditions on Mars and assess it from the perspective of potential exobiological evolution.
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The question of the chemical origins of life is engraved in the European scientific patrimony as it can be traced back to the pioneer ideas of Charles Darwin, Louis Pasteur, and more recently to Alexander Oparin. During the last decades, the European community of origin of life scientists has organized seven out of the twelve International Conferences on the Origins of Life held since 1957. This community contributed also to enlarge the field of research to the study of life in extreme environments and to the search for extraterrestrial life, i.e. exobiology in its classical definition or astrobiology if one uses a more NASA-inspired terminology. The present paper aims to describe the European science background in exo/astrobiology as well as the project of a European Network of Exo/Astrobiology.
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The origin and diversification of RNA secondary structure were traced using cladistic methods. Structural components were coded as polarized and ordered multi-state characters, following a model of character state transformation outlined by considerations in statistical mechanics. Several classes of functional RNA were analyzed, including ribosomal RNA (rRNA). Considerable phylogenetic signal was present in their secondary structure. The intrinsically rooted phylogenies reconstructed from evolved RNA structure depicted those derived from nucleic acid sequence at all taxonomical levels, and grouped organisms in concordance with traditional classification, especially in the archaeal and eukaryal domains. Natural selection appears therefore to operate early in the information flow that originates in sequence and ends in an adapted phenotype. When examining the hierarchical classification of the living world, phylogenetic analysis of secondary structure of the small and large rRNA subunits reconstructed a universal tree of life that branched in three monophyletic groups corresponding to Eucarya, Archaea, and Bacteria, and was rooted in the eukaryotic branch. Ribosomal characters involved in the translational cycle could be easily traced and showed that transfer RNA (tRNA) binding domains in the large rRNA subunit evolved concurrently with the rest of the rRNA molecule. Results suggest it is equally parsimonious to consider that ancestral unicellular eukaryotes or prokaryotes gave rise to all extant life forms and provide a rare insight into the early evolution of nucleic acid and protein biosynthesis.
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Intertwined cycles of matters are proposed to be one of the essential characteristics of life. The increase in the amount of recycled matters within biological groups and the complication of the cycling network are considered to be the early development of life soon after its appearance on the Earth. From this point of view, the origin of a cell alone can not sufficiently be regarded as the origin of life. The origin of a cell must be followed by a formation of material cycling among cells, if the cells are to stay on the Earth long enough so that they could be called life. This biogeochemical viewpoint leads to a quantitative analysis of life activity. A formula is presented to exemplify the potential utility of the viewpoint. It is applied to analyses of possible early developments of life and of an impact of present human activity on the global carbon cycle. Further application of the viewpoint to a variety of biogeochemical, sociogeochemical, ecological, and environmental problems should eventually show whether there is some utility in this view.
In 1959, Miller and Urey (Science 130, 245) published their classic compilation of energy sources for indigenous prebiotic organic synthesis on the early Earth. Much contemporary origins of life research continues to employ their original estimates for terrestrial energy dissipation by lightning and coronal discharges, 2 x 10(19) J yr-1 and 6 x 10(19) J yr-1, respectively. However, more recent work in terrestrial lightning and point discharge research suggests that these values are overestimates by factors of about 20 and 120, respectively. Calculated concentrations of amino acids (or other prebiotic organic products) in the early terrestrial oceans due to electrical discharge sources may therefore have been equally overestimated. A review of efficiencies for those experiments that provide good analogues to naturally-occurring lightning and coronal discharges suggests that lightning energy yields for organic synthesis (nmole J-1) are about one order of magnitude higher than those for coronal discharge. Therefore organic production by lightning may be expected to have dominated that due to coronae on early Earth. Limited data available for production of nitric oxide in clouds suggests that coronal emission within clouds, a source of energy heretofore too uncertain to be included in the total coronal energy inventory, is insufficient to change this conclusion. Our recommended values for lightning and coronal discharge dissipation rates on the early Earth are, respectively, 1 x 10(18) J yr-1 and 5 x 10(17) J yr-1.
The origin of the channels on Mars has been a subject of intense interest since they were first recognized on early Mariner 9 images (Driscoll, 1972; Masursky, 1973). Their presence on the planet, and their striking resemblance to terrestrial flood channels related to glacial outbursts or to dendritic river systems has suggested to most investigators (Baker, 1974, 1977; Nummedal, 1978; Carr, 1979; Masursky et al., 1977) that they were formed by running water. Because life as we know it is dependent on water, the discovery by the 'Mariner cameras', of watercut channels and volcanoes as a source for water, and water ice in the residual north polar cap by Viking, has reaffirmed the choice of Mars as the best target for the search for extraterrestrial life.