1.3
IS EVOLUTION AN ESSENTIAL FEATURE OF LIFE?

Many of the definitions of life include phrases like undergoes Darwinian evolution. The implication is that phenotypic changes and adaptation are necessary to exploit unstable environmental conditions and to function optimally in the environment. Evolutionary changes have even been suggested for the hypothesized “clay crystal life” of Cairns-Smith,5 referring to randomly occurring errors in crystal structure during crystal growth as analogous to mutations. Would a self-replicating chemical system capable of chemical transformations in the environment be considered life? If self-replicating chemical compounds are not life, replication by itself is not sufficient as a defining characteristic of life. Likewise, the ability to undergo Darwinian evolution, a process that results in heritable changes in a population, is also not sufficient to define life if we consider minerals that are capable of reproducing errors in their crystal structure to be equivalent to evolution. Although that property of clays may have been vital in the origin of life and particularly in the prebiotic synthesis of organic macromolecules and as catalysts for metabolic reactions, can the perpetuation of “mistakes” in crystal structure result in the selection of a “more fit” crystal structure? It is important to emphasize that evolution is not simply reproducing mutations (mistakes in clays), but also selecting variants that are functionally more fit.

The canonical characteristics of life are an inherent capacity to adapt to changing environmental conditions and to interact with other living organisms (and, at least on Earth, also with viruses).6 Natural selection is the key to evolution and the main reason that Darwinian evolution persists as a characteristic of many definitions of life. The only alternative to evolution for producing diversity would be to have environmental conditions that continuously create different life forms or similar life forms with random and frequent “mistakes” in the synthesis of chemical templates used for replication or metabolism. Such mistakes would be equivalent to mutations and could lead to traits that gave some selective advantage in an existing community or in exploiting new habitats. That random process could lead to life forms that undergo a form of evolution without a master information macromolecule, such as DNA or RNA. It is difficult to imagine such life forms as able to “evolve” into complex structures unless other mechanisms, such as symbiosis or cell-cell fusion, are available.

Evolution is the key mechanism of heritable changes in a population. However, although mutation and natural selection are important processes, they are not the only mechanisms for acquiring new genes. It is understood that lateral gene transfer is one of the most important and one of the earliest mechanisms for creating diversity and possibly for building genomes with the requisite information to result in free-living cells.7 Lateral gene transfer is also one of the mechanisms to align genes from different sources into complex functional activities, such as magnetotaxis and dissimilatory sulfate reduction.8 It is possible that this mechanism was important in the evolution of metabolic and biosynthetic pathways and other physiological traits that may have evolved only once even though they are present in a wide variety of organisms. Coevolution of two or more species is also a hallmark of evolution manifested in many ways, from insect-plant interactions to the involvement of hundreds of species of bacteria in the nutrition of ruminant animals. Organisms and the environment also coevolve, depending on the dominant characteristics of the environment and the availability of carbon and energy sources.

If the ability to undergo Darwinian evolution is a canonical trait of life no matter how different a life form is from Earth life, are there properties of evolving extraterrestrial organisms that would be detectable as positive signs of life? Evolution provides organisms the opportunity to exploit new and changing environments, and one piece of evidence for the cosmic ubiquity of evolution is that on Earth life occupies all available habitats and even creates new ones as a consequence of metabolism. Another hallmark of evolution is the ability of organisms to coevolve with other organisms and to form permanent and obligatory associations. It is highly probable that an inevitable consequence of evolution is the elimination of radically different biochemical lineages of life that may have formed during the earliest period of the evolution of life. Extant Earth life is the result of either selection of the most fit lineage or homogenization of some or all of the different lineages into a common ancestral community that developed into the current three major lineages (domains). All have a common biochemistry based on presumably the most “fit” molecular information strategies and energy-yielding pathways among a potpourri of possibilities.

Thus, one of the apparent generalizations that can be drawn from extant Earth life, and the explanation for the development of a “unity of biochemistry” in all organisms, is that lateral gene transfer is an ancient and efficient mechanism for rapidly creating diversity and complexity. Lateral gene transfer is also an efficient mechanism for



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