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While modern physics and biology satisfactorily explain the passage from the Big Bang to the formation of Earth and the first cells to present-day life, respectively, the origins of biochemical life still remain an open question. Since life, as we know it, requires extremely long genetic polymers, any answer to the question must explain how an evolving system of polymers of ever-increasing length could come about on a planet that otherwise consisted only of small molecular building blocks. In this work we show that, under realistic constraints, an abstract polymer model can exhibit dynamics such that attractors in the polymer population space with a higher average polymer length are also more probable. We generalize from the model and formalize the notions of and for chemical reaction networks with multiple attractors. The complexity of a species is defined as the minimum number of reactions needed to produce it from a set of building blocks, which in turn is used to define a measure of complexity for an attractor. A transition between attractors is considered to be a if the attractor with the higher probability also has a higher complexity. In an environment where only monomers are readily available, the attractor with a higher average polymer length is more complex. Thus, by this criterion, our abstract polymer model can exhibit progressive evolution for a range of thermodynamically plausible rate constants. We also formalize criteria for and evolution and explain the role of autocatalysis in obtaining them. Our work provides a basis for searching for prebiotically plausible scenarios in which long polymers can emerge and yield populations with even longer polymers. Additionally, the existence of features like history dependence and open endedness support the view that the path from chemistry to biology was one of gradual complexification rather than an instantaneous origin of life. Published by the American Physical Society2025 

Developing a mathematical understanding of autocatalysis in reaction networks has both theoretical and practical implications. We review definitions of autocatalytic networks and prove some properties for minimal autocatalytic subnetworks (MASs). We show that it is possible to classify MASs in equivalence classes, and develop mathematical results about their behavior. We also provide linear-programming algorithms to exhaustively enumerate them and a scheme to visualize their polyhedral geometry and combinatorics. We then define cluster chemical reaction networks, a framework for coarse-graining real chemical reactions with positive integer conservation laws. We find that the size of the list of minimal autocatalytic subnetworks in a maximally connected cluster chemical reaction network with one conservation law grows exponentially in the number of species. We end our discussion with open questions concerning an ecosystem of autocatalytic subnetworks and multidisciplinary opportunities for future investigation. 

The vesicles of short chain amphiphiles have been demonstrated to grow and divide. Here, we explored whether vesicle populations show evidence of heritability. We prepared 1:1 decanoic acid:decylamine vesicles with or without a detergent and in either water or prebiotic soup, a mixture of compounds that might have been present on early Earth. The mixtures were subjected to transfer with dilution, where, after 24 h of incubation (one generation), we transferred 10% of the mix into a 90% volume of a fresh vesicle-containing solution. This was continued for 30 generations. Samples with a history of transfers were compared to no-transfer controls (NTCs), initiated each generation using the same solutions but without 10% of the prior generation. We compared the vesicle size distribution and chemical composition of the transfer samples and NTCs and compared their fluorescence signals in the presence of Nile Red dye. We observe changes in the vesicle size but did not detect differences in the chemical composition. In the samples with detergent and soup, we observed irregular changes in the Nile Red fluorescence, with a tendency for parent and offspring samples to have correlated values, suggestive of heritability. This last result, combined with evidence of temporal autocorrelation across generations, suggests the possibility that vesicles could respond to selection. 

Prior research on evolutionary mechanisms during the origin of life has mainly assumed the existence of populations of discrete entities with information encoded in genetic polymers. Recent theoretical advances in autocatalytic chemical ecology establish a broader evolutionary framework that allows for adaptive complexification prior to the emergence of bounded individuals or genetic encoding. This framework establishes the formal equivalence of cells, ecosystems and certain localized chemical reaction systems as autocatalytic chemical ecosystems (ACEs): food-driven (open) systems that can grow due to the action of autocatalytic cycles (ACs). When ACEs are organized in meta-ecosystems, whether they be populations of cells or sets of chemically similar environmental patches, evolution, defined as change in AC frequency over time, can occur. In cases where ACs are enriched because they enhance ACE persistence or dispersal ability, evolution is adaptive and can build complexity. In particular, adaptive evolution can explain the emergence of self-bounded units (e.g. protocells) and genetic inheritance mechanisms. Recognizing the continuity between ecological and evolutionary change through the lens of autocatalytic chemical ecology suggests that the origin of life should be seen as a general and predictable outcome of driven chemical ecosystems rather than a phenomenon requiring specific, rare conditions. 

The problem of identifying conditions that enable major evolutionary transitions, in which distinct units come together to form a new higher level unit, is a complex and difficult topic spanning many disciplines. Here, we approach this problem from the perspective of the origin of life, which allows us to make the simplifying assumption that the lower-level units are not also evolving. This assumption lets us focus on identifying environmental factors that promote egalitarian major transitions in general and the origin of life specifically. To study this question, we build a simple artificial ecology model. We quantify major-transition-like dynamics using a maximum likelihood approach and a set of null models predicting the behavior of our system under various dynamics. Ultimately, we find that, even in a maximally simple artificial ecology model, we are able to observe evidence of community-level selection and thus the beginnings of a major evolutionary transition. The regions of parameter space that promote community-level selection vary based on species interactions but we observe consistent trends. 

Identifying conditions that promote egalitarian major transitions, where unlike replicating units unite to form a higher-level unit, is an open problem with far-reaching implications. We propose that egalitarian major transitions can only begin in ecological communities that are conducive to them. To formalize this idea, we introduce the concept of “transition-ability”, which describes the extent to which a community is poised to undergo an egalitarian major transition. We hypothesize that transitionability is a property of ecological interaction networks, which represent the set of pairwise interactions among members of a community. Using a digital artificial ecology that simulates interactions between species based on a static interaction network, we test the transition-ability of interaction networks created by a range of graph-generation techniques, as well as some real-world ecological networks. To measure the extent to which a community is moving towards a major transition, we quantify the increase in community-level fitness relative to individual-level fitness across five different fitness proxies. We find that some network generation protocols produce more transitionable networks than others. In particular, interaction strengths (i.e. edge weights) have a substantial impact on transitionability, despite receiving low attention in the literature. 

“Prebiotic soup” often features in discussions of origins of life research, both as a theoretical concept when discussing abiological pathways to modern biochemical building blocks and, more recently, as a feedstock in prebiotic chemistry experiments focused on discovering emergent, systems-level processes such as polymerization, encapsulation, and evolution. However, until now, little systematic analysis has gone into the design of well-justified prebiotic mixtures, which are needed to facilitate experimental replicability and comparison among researchers. This paper explores principles that should be considered in choosing chemical mixtures for prebiotic chemistry experiments by reviewing the natural environmental conditions that might have created such mixtures and then suggests reasonable guidelines for designing recipes. We discuss both “assembled” mixtures, which are made by mixing reagent grade chemicals, and “synthesized” mixtures, which are generated directly from diversity-generating primary prebiotic syntheses. We discuss different practical concerns including how to navigate the tremendous uncertainty in the chemistry of the early Earth and how to balance the desire for using prebiotically realistic mixtures with experimental tractability and replicability. Examples of two assembled mixtures, one based on materials likely delivered by carbonaceous meteorites and one based on spark discharge synthesis, are presented to illustrate these challenges. We explore alternative procedures for making synthesized mixtures using recursive chemical reaction systems whose outputs attempt to mimic atmospheric and geochemical synthesis. Other experimental conditions such as pH and ionic strength are also considered. We argue that developing a handful of standardized prebiotic recipes may facilitate coordination among researchers and enable the identification of the most promising mechanisms by which complex prebiotic mixtures were “tamed” during the origin of life to give rise to key living processes such as self-propagation, information processing, and adaptive evolution. We end by advocating for the development of a public prebiotic chemistry database containing experimental methods (including soup recipes), results, and analytical pipelines for analyzing complex prebiotic mixtures.