Origin Of Life Scenarios Research Articles

Three Biopolymers and Origin of Life Scenarios.

To track down the possible roots of life, various models for the initial living system composed of different combinations of the three extant biopolymers, RNA, DNA, and proteins, are presented. The suitability of each molecular set is assessed according to its ability to emerge autonomously, sustain, and evolve continuously towards life as we know it. The analysis incorporates current biological knowledge gained from high-resolution structural data and large sequence datasets, together with experimental results concerned with RNA replication and with the activity demonstrated by standalone constructs of the ribosomal Peptidyl Transferase Center region. The scrutiny excludes the DNA-protein combination and assigns negligible likelihood to the existence of an RNA-DNA world, as well as to an RNA world that contained a replicase made of RNA. It points to the precedence of an RNA-protein system, whose model of emergence suggests specific processes whereby a coded proto-ribosome ribozyme, specifically aminoacylated proto-tRNAs and a proto-polymerase enzyme, could have autonomously emerged, cross-catalyzing the formation of each other. This molecular set constitutes a feasible starting point for a continuous evolutionary path, proceeding via natural processes from the inanimate matter towards life as we know it.

Electron transport chains as a window into the earliest stages of evolution

The origin and early evolution of life is generally studied under two different paradigms: bottom up and top down. Prebiotic chemistry and early Earth geochemistry allow researchers to explore possible origin of life scenarios. But for these “bottom–up” approaches, even successful experiments only amount to a proof of principle. On the other hand, “top–down” research on early evolutionary history is able to provide a historical account about ancient organisms, but is unable to investigate stages that occurred during and just after the origin of life. Here, we consider ancient electron transport chains (ETCs) as a potential bridge between early evolutionary history and a protocellular stage that preceded it. Current phylogenetic evidence suggests that ancestors of several extant ETC components were present at least as late as the last universal common ancestor of life. In addition, recent experiments have shown that some aspects of modern ETCs can be replicated by minerals, protocells, or organic cofactors in the absence of biological proteins. Here, we discuss the diversity of ETCs and other forms of chemiosmotic energy conservation, describe current work on the early evolution of membrane bioenergetics, and advocate for several lines of research to enhance this understanding by pairing top–down and bottom–up approaches.

Submarine hydrothermal vent systems: the relevance of dynamic systems in chemical evolution and prebiotic chemistry experiments

AbstractSince their discovery, submarine hydrothermal vent systems have been pointed out as important places where chemical evolution on Earth could have occurred; and their role in the process has been highlighted. Similarly, some hypotheses have considered these systems in origin of life scenarios. In this way, many experiments have been developed, and the knowledge about these systems has increased. Due to their complexity, many experimental simulations have only included a few of the geochemical variables present in these environments, pressure and temperature. Other main variables have hardly been included, such as mineralogy, thermal and pH gradients, dissolved ions and/or redox reactions. As it has been understood, the dynamism and heterogeneity of these environments are huge, and it comprises different scales, from single vents to full hydrothermal fields. However, the vast majority of experiments focus on a specific part of these systems and do not include salinity, mineralogy and pH gradients. For this reason, in this paper, we pointed out some considerations about how this dynamism can be interpreted, and included in some models, as well their importance in prebiotic chemistry experiments and their extrapolations regarding the hypothesis about the origins of life.

Prebiotic Protocell Model Based on Dynamic Protein Membranes Accommodating Anabolic Reactions.

The nature of the first prebiotic compartments and their possible minimal molecular composition is of great importance in the origin of life scenarios. Current protocell model membranes are proposed to be lipid-based. This paradigm has several shortcomings such as limited membrane stability of monoacyl lipid-based membranes (e.g., fatty acids), missing pathways to synthesize protocell membrane components (e.g., phospholipids) under early earth conditions, and the requirement for different classes of molecules for the formation of compartments and the catalysis of reactions. Amino acids on the other hand are known to arise and persist with remarkable abundance under early earth conditions since the fundamental Miller-Urey experiments. They were also postulated early to form protocellular structures, for example, proteinoid capsules. Here, we present a protocell model constituted by membranes assembled from amphiphilic proteins based on prebiotic amino acids. Self-assembled dynamic protein membrane-based compartments (PMBCs) are impressively stable and compatible with prevalent cellular membrane constituents forming protein-only or protein-lipid hybrid membranes. They can embed processes essential for extant living cells, such as enclosure of molecules, membrane fusion, phase separation, and complex biosynthetic elements from modern cells demonstrating "upward" compatibility. Our findings suggest that prebiotic PMBCs represent a new type of protocell as a possible ancestor of current lipid-based cells. The presented prebiotic PMBC model can be used to design artificial cells, important for the study of structural, catalytic, and evolutionary pathways related to the emergence of life.

Autocatalytic confusion clarified

There is frequent confusion about the terms autocatalytic reaction, autocatalytic cycle, and autocatalytic set. As the use of the same adjective implies, these three systems do indeed share common properties, in particular their potential for exponential growth. However, the ways in which they achieve this potential are different, giving rise to different internal network structures and dynamics. Therefore, care should be taken which term is used in which context. Here, we explain and discuss the similarities and differences between the three systems in detail, in an effort to avoid any further confusion. We then also discuss the relevance of these autocatalytic systems for possible origin of life scenarios, with an emphasis on how autocatalytic sets may have played an important role in this.

Vesicle aggregates as a model for primitive cellular assemblies.

Primitive cell models help to understand the role that compartmentalization plays in origin of life scenarios. Here we present a combined experimental and modeling approach towards the construction of simple model systems for primitive cellular assemblies. Charged lipid vesicles aggregate in the presence of oppositely charged biopolymers, such as nucleic acids or polypeptides. Based on zeta potential measurements, dynamic light scattering and cryo-transmission electron-microscopy, we have characterized the behavior of empty and ferritin-filled large unilamellar POPC vesicles, doped with different amounts of cationic (DDAB, CTAB) and anionic (sodium oleate) surfactants, and their aggregation upon the addition of anionic (tRNA, poly-l-glutamic acid) and cationic (poly-l-arginine) biopolymers, respectively. The experimental results are rationalized by a phenomenological modeling approach that predicts the average size of the vesicle aggregates as function of the amount of added biopolymers. In addition, we discuss the mechanism of vesicle aggregation induced by oppositely charged biopolymers. Our study complements previous reports about the formation of giant vesicle clusters and thus provides a general vista on primitive cell systems, based on the association of vesicles into compartmentalized aggregates.

What roles for viruses in origin of life scenarios?

Important roles in origin of life (OL) scenarios have been and still are attributed to viruses. Yet the strict dependence of viruses on cells for their multiplication has been widely acknowledged since the first decades of the 20th century. How could viruses play critical roles in the OL if life relies on cellular organization and if viruses are defined as parasites of cells? In other words, how could viruses play a role in the emergence of cellular life if the existence of cells is a prerequisite for the existence of viruses? This paper investigates this issue and describes past and current OL scenarios conferring viruses with important roles, thereby completing the work of historian of science and physician Scott Podolsky who identified three major roles of viruses in past OL scenarios. Some objections raised by present OL scenarios conferring viruses with an important role are discussed. I argue that disagreements concerning the roles of viruses in OL scenarios stem from the different concepts of life and of virus scientists defend. Investigating the roles of viruses in OL scenarios not only helps identifying different ways to define life in the context of OL theorizing. It also offers the opportunity to better understand how viruses could be conceptualized. The relevance of the replication-first versus metabolism-first dichotomy in OL theorizing is briefly discussed.

The natural history of transfer RNA and its interactions with the ribosome.

Transfer RNA (tRNA) is undoubtedly the most central and one of the oldest molecules of the cell. Without it genetics and coded protein synthesis are impossible. The crucial specificities responsible for the genetic code and accurate translation are by far entrusted to interactions between tRNA and translation proteins, fundamentally aminoacyl-tRNA synthetase (aaRS) enzymes and elongation factor (EF) switches (Yadavalli and Ibba, 2012). Discrimination mediated by aaRSs and EFs against misincorporated tRNA and amino acids is at least 20 times more stringent than ribosomal recognition, editing, and other proofreading mechanisms (Reynolds et al., 2010). The fact that crucial genetic code specificities in highly selective interactions with protein enzymes do not involve the ribosomal ribonucleoprotein biosynthetic machinery challenges the “replicators first” origin of life scenario of an ancient RNA world (Caetano-Anolles and Seufferheld, 2013). It also highlights the central functional, mechanistic, and evolutionary roles of tRNA and its recognition determinants, which enable coevolution between nucleic acids and proteins. These coevolutionary relationships are compatible with a late origin of the ribosome in its mechanism and not in protein biosynthesis, which was inferred from the computational analysis of thousands of RNAs and proteomes (Harish and Caetano-Anolles, 2012). These analyses showed tight coevolution of ribosomal RNA (rRNA) and ribosomal proteins (r-proteins). While these relationships delimit molecular makeup when organisms use translation to negotiate growth and viability amidst environmental change, coevolution also constrains recruitment of the canonical L-shaped structure of the tRNA molecule into a multiplicity of modern functions. These new functions include the synthesis of antibiotics, bacterial cell wall peptidoglycans and tetrapyrroles, modification of bacterial membrane lipids, protein turnover, and the synthesis of other aminoacyl-tRNA molecules (Francklyn and Minajigi, 2010). Here we unfold coevolutionary relationships between tRNA substructures and translation proteins that embody crucial protein-nucleic acid interactions. We focus on a series of computational biology analyses of the structure and conformational diversity of tRNAs and their interacting proteins that provide information about the history of structural accretion of this “adaptor” molecule. Using this information, we place tRNA history within the framework of an evolutionary timeline of protein domain innovation, uncovering the natural history of tRNA within the context of the geological record.

Recent Theoretical Approaches to Minimal Artificial Cells

Minimal artificial cells (MACs) are self-assembled chemical systems able to mimic the behavior of living cells at a minimal level, i.e. to exhibit self-maintenance, self-reproduction and the capability of evolution. The bottom-up approach to the construction of MACs is mainly based on the encapsulation of chemical reacting systems inside lipid vesicles, i.e. chemical systems enclosed (compartmentalized) by a double-layered lipid membrane. Several researchers are currently interested in synthesizing such simple cellular models for biotechnological purposes or for investigating origin of life scenarios. Within this context, the properties of lipid vesicles (e.g., their stability, permeability, growth dynamics, potential to host reactions or undergo division processes…) play a central role, in combination with the dynamics of the encapsulated chemical or biochemical networks. Thus, from a theoretical standpoint, it is very important to develop kinetic equations in order to explore first—and specify later—the conditions that allow the robust implementation of these complex chemically reacting systems, as well as their controlled reproduction. Due to being compartmentalized in small volumes, the population of reacting molecules can be very low in terms of the number of molecules and therefore their behavior becomes highly affected by stochastic effects both in the time course of reactions and in occupancy distribution among the vesicle population. In this short review we report our mathematical approaches to model artificial cell systems in this complex scenario by giving a summary of three recent simulations studies on the topic of primitive cell (protocell) systems.

The Origin of Cellular Life and Biosemiotics

Recent successes of systems biology clarified that biological functionality is multilevel. We point out that this fact makes it necessary to revise popular views about macromolecular functions and distinguish between local, physico-chemical and global, biological functions. Our analysis shows that physico-chemical functions are merely tools of biological functionality. This result sheds new light on the origin of cellular life, indicating that in evolutionary history, assignment of biological functions to cellular ingredients plays a crucial role. In this wider picture, even if aggregation of chance mutations of replicator molecules and spontaneously self-assembled proteins led to the formation of a system identical with a living cell in all physical respects but devoid of biological functions, it would remain an inanimate physical system, a pseudo-cell or a zombie-cell but not a viable cell. In the origin of life scenarios, a fundamental circularity arises, since if cells are the minimal units of life, it is apparent that assignments of cellular functions require the presence of cells and vice versa. Resolution of this dilemma requires distinguishing between physico-chemical and biological symbols as well as between physico-chemical and biological information. Our analysis of the concepts of symbol, rule and code suggests that they all rely implicitly on biological laws or principles. We show that the problem is how to establish physico-chemically arbitrary rules assigning biological functions without the presence of living organisms. We propose a solution to that problem with the help of a generalized action principle and biological harnessing of quantum uncertainties. By our proposal, biology is an autonomous science having its own fundamental principle. The biological principle ought not to be regarded as an emergent phenomenon. It can guide chemical evolution towards the biological one, progressively assigning greater complexity and functionality to macromolecules and systems of macromolecules at all levels of organization. This solution explains some perplexing facts and posits a new context for thinking about the problems of the origin of life and mind.

Formamide in non-life/life transition

We would like to thank all commentators for their insightful commentaries [1–7]. Here we briefly discuss some ofthe important observations raised in the commentaries.Kaempfer [1] raised an important point about the role of RNA in the origin of life scenario. The plasticity of RNAis illustrated not only by catalytic properties, by its role in performing the functions of the ribosome in translation ofmRNA, by its riboswitch responsiveness: it is also shown by its function of active sensor in stress signaling [8].Thisproperty has possibly endowed RNA with the ability to evolve to what is the extant protective immune response [9].All these properties, as stressed by the author, lend support to the scenario that puts RNA in the key position tokick-start life.Burton [2] underscores the relevance, in the assessment of the possible prebiotic role of formamide-based syntheticreactions, of the physical chemical conditions in which they have been observed. Nucleic bases, carboxylic acidsand aminoacids are typically obtained in the presence of catalysts by warming at 140–160

A case for landing on the moon's farside to test nitrogen abundances

AbstractA high research priority in astrobiology is the search and eventual identification of biomarkers in the Solar System. In spite of numerous steps forward, lunar science remains largely disjoint from the main stream of astrobiology, but in recent years the Moon has begun to emerge as a novel target for astrobiologists. We discuss an overlap between lunar geology and terrestrial geomicrobiology that arises from analysis of lunar soils and some uncertainties in chemical evolution and the origin of life scenarios. Unexpected isotopic heterogeneity of nitrogen (N) was found to be remarkable in samples from Apollo and the Luna programme. Both the stable isotope geochemical data of the biogenic elements, as well as the noble gases trapped in lunar soils added valuable new and relevant data. These discoveries are potential sources of information on early Earth evolution. The elusive ratio of N's two stable isotopes 15N/14N has played a fundamental role in this research. The analysis of individual grains of ilmenite suggests that 90% of all the trapped N does not originate from solar wind (SW). We discuss the significance of these stable isotopes from the point of view of astrobiology in the light of the next generation of lunar exploration. We underline the high priority of testing the origin of non-solar N source trapped in the regolith of the lunar farside. In the proposals of new lunar missions, the characterization of the geochemistry at several lunar sites is a major objective. Some arguments are presented in favour of using novel space technologies in a search for biomarkers in geographical distinct lunar landing sites. We restrict our attention to one aspect of the science requirements for the forthcoming missions by focusing on a very limited objective: to take a closer look at the geochemical characterization of the chemical element N on the soils of the lunar farside.

Protective effect of clay minerals on adsorbed nucleic acid against UV radiation: possible role in the origin of life

The effect of UV radiation on solutions of free and clay-adsorbed DNA has been investigated. It turns out that clay (montmorillonite/kaolinite) adsorbed nucleic acid undergoes less radiation damage than free nucleic acid. Our laboratory experiments have an astronomical counterpart in terms of solar irradiance on the Earth. An origin of life scenario is proposed where ubiquitous clay minerals lead the surface chemistry of the molecules relevant to the biological evolution and at the same time protect them from the deadly rainfall of UV photons.

Archaeal genome organization and stress responses: implications for the origin and evolution of cellular life.

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.

The Molecular Roots of Compositional Inheritance

Non-covalent compositional assemblies, made of monomeric mutually catalytic molecules, constitute an alternative to alphabet-based informational biopolymers as a mechanism of primordial inheritance. Such assemblies appear implicitly in many “Metabolism First” origin of life scenarios, and more explicitly in the Graded Autocatalysis Replication Domain (GARD) model [Segréet al. (2000). Proc. Natl Acad. Sci. U.S.A.97, 4112–4117]. In the present work, we provide a detailed analysis of the quantitative molecular roots of such behavior. It is demonstrated that the fidelity of reproduction provided by a newly defined heritability measure η*s, strongly depends on the values of molecular recognition parameters and on assembly size. We find that if the catalytic rate acceleration coefficients are distributed normally, transfer of compositional information becomes impossible, due to frequent “compositional error catastrophes”. In contrast, if the catalytic acceleration rates obey a lognormal distribution, as actually predicted by a statistical formalism for molecular repertoires, high reproduction fidelity is obtained. There is also a clear dependence on assembly size N, whereby maximal η is seen in a narrow range around N∼3.5NG /λ, where NGis the size of the primordial molecular repertoire and λ is a molecular interaction statistical parameter. Such relationships help define the physicochemical conditions that could underlie the early steps in pre-biotic evolution.