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A Sourcebook for the Worldwide Discovery of a Creative Organic Universe
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V. Life's Corporeal Evolution Develops, Encodes and Organizes Itself: An EarthWinian Genesis Synthesis

4. Multicellular Fauna and Flora Organisms in Transition

Ratcliff, William, et al. Experimental Evolution of Multicellularity. Proceedings of the National Academy of Sciences. 109/1595, 2012. University of Minnesota, Evolution and Behavior and BioTechnology Institute, ecologists start their project by situating it within the popular “major transitions” scale and sequence from proteins to people. By this view, life’s emergent passage from eukaryotic cells to cellular assemblies was a natural next stage of “sophisticated, higher-level functionality via cooperation among component cells with complementary behaviors.” See herein concurrent work by Iaroslav Ispolatov, et al, Carl Simpson below, and Cooperative Alliances in the Emergent Individuality section.

Multicellularity was one of the most significant innovations in the history of life, but its initial evolution remains poorly understood. Using experimental evolution, we show that key steps in this transition could have occurred quickly. We subjected the unicellular yeast Saccharomyces cerevisiae to an environment in which we expected multicellularity to be adaptive. We observed the rapid evolution of clustering genotypes that display a novel multicellular life history characterized by reproduction via multicellular propagules, a juvenile phase, and determinate growth. The multicellular clusters are uniclonal, minimizing within-cluster genetic conflicts of interest. Simple among-cell division of labor rapidly evolved. Early multicellular strains were composed of physiologically similar cells, but these subsequently evolved higher rates of programmed cell death (apoptosis), an adaptation that increases propagule production. These results show that key aspects ofmulticellular complexity, a subject of central importance to biology, can readily evolve from unicellular eukaryotes. (Abstract, 1595)

Ravasz, E., et al. Hierarchical Organization of Modularity in Metabolic Networks. Science. 297/1551, 2002. The universal properties of scale-free systems apply in developing cellular organization which forms many functional modules arranged in interconnected webs with a nested hierarchical iteration.

Here, we show that the metabolic networks of 43 distinct organism are organized into many small, highly connected topologic modules that combine in a hierarchical manner into larger, less cohesive units, with their number and degree of clustering following a power law. (1551)

Reinhard, Christopher, et al. Earth’s Oxygen Cycle and the Evolution of Animal Life. Proceedings of the National Academy of Sciences. 113/8933, 2016. A senior team of geoscientists including Douglas Erwin, and edited by Andrew Knoll, quantify life’s billion year span before the Cambrian profusion some 600 million years ago. Our rare planetary surface balance of both sea and land is seen as a crucial reason that Earth life has developed beyond an anoxic marine phase which is not conducive.

Earth is currently the only planet known to harbor complex life. Understanding whether terrestrial biotic complexity is a unique phenomenon or can be expected to be widespread in the universe depends on a mechanistic understanding of the factors that led to the emergence of complex life on Earth. Here, we use geochemical constraints and quantitative models to suggest that marine environments may have been unfavorable for the emergence and large-scale proliferation of motile multicellular life for most of Earth’s history. Further, we argue that a holistic evaluation of environmental variability, organismal life history, and spatial ecological dynamics is essential for a full accounting of the factors that have allowed for the emergence of biological complexity on Earth. (Significance)

Retallack, Gregory. Great Moments in Plant Evolution. PNAS. 118/17, 2021. A University of Oregon paleontologist reviews notable work, as below, with regard to our whole scale reconstruction of how life proceeded through all the fauna and flora to arrive at a global speciesphere whom is able to do this. Our take, as the Abstract cites, is to muse that these past creaturely occasions appear as if they are intended, indeed expected, stages along the ascendant way.

Just as dinosaurs can be regarded as protobirds (1) and synapsids as protomammals (2), understanding the evolution of plants depends on extinct fossil groups, such as those linking spore and seed plants. A profound paleobotanical surprise in 1904 was discovery of pteridosperms, commonly called seed ferns, because of their unexpected combination of seeds and fern-like leaves. Another paleobotanical surprise in 1960 was discovery of progymnosperms, spore plants with woody anatomy comparable to modern conifer trees. Now, Wang et al. in PNAS (118/11) report a Chinese Permian fossil plant, Paratinga wuhaia. These permineralized fossils convincingly expand the progymnosperm clade to include the enigmatic Noeggerathiales, an extinct order of vascular plants.

Rokas, Antonis. The Origins of Multicellularity and the Early History of the Genetic Toolkit for Animal Development. Annual Review of Genetics. 42/235, 2008. A Vanderbilt University biologist reports the latest findings for the persistent, multiple appearance of cellular organisms from biomolecular substrates, which is then seen to convey group survival advantages. But as usual mechanical terms prevail, along with no inkling of a deeper source which could drive such increasing complexity. A icrobiology blog on the article takes Rokas to task for using the phrase “reasons” for multicellularity – to wit, don’t you know there are no reasons for anything. Not to be on his case, since this is the prevailing paradigm, but a 2012 blog citation on Rokas' website is "The Global Proteome Machine."

Multicellularity appeared early and repeatedly in life's history; its instantiations presumably required the confluence of environmental, ecological, and genetic factors. Comparisons of several independently evolved pairs of multicellular and unicellular relatives indicate that transitions to multicellularity are typically associated with increases in the numbers of genes involved in cell differentiation, cell-cell communication, and adhesion. Further examination of the DNA record suggests that these increases in gene complexity are the product of evolutionary innovation, tinkering, and expansion of genetic material. (Abstract, 235)

Ruiz-Trillo, Inaki and Aurora Nedelcu, eds. Evolutionary Transitions to Multicellular Life. Berlin: Springer, 2015. As an example of how much the major transitions model is now accepted and availed to structure and explain life’s episodic ascent, the many chapters herein explore its presence and effect across all manner of microbial forms such as biofilms and algae. The main sections are Multicellularity in the Tree of Life, Model Systems, Theoretical Approaches, Genomic Insights, and Molecular Mechanisms. See also a companion Springer volume Multicellularity: Origins and Evolution edited by Karl Niklas and Stuart Newman (2016).

The book integrates our understanding of the factors and processes underlying the evolution of multicellularity by providing several complementary perspectives (both theoretical and experimental) and using examples from various lineages in which multicellularity evolved. Recent years marked an increased interest in understanding how and why these transitions occurred, and data from various fields are providing new insights into the forces driving the several independent transitions to multicellular life as well as into the genetic and molecular basis for the evolution of this phenotype. The ultimate goal of this book is to facilitate the identification of general and unifying principles and mechanisms. (Publisher)

Ruiz-Trillo, Iraki, et al. The Origins of Multicellularity: a Multi-taxon Genome Initiative. Trends in Genetics. 23/3, 2007. A multi-author paper that lays out the project introduced in the abstract below.

The emergence of multicellular organisms from single-celled ancestors – which occurred several times, independently in different branches of the eukaryotic tree – is one of the most profound evolutionary transitions in the history of life. These events not only radically changed the course of life on Earth but also created new challenges, including the need for cooperation and communication between cells, and the division of labor among different cell types. However, the genetic changes that accompanied the several origins of multicellularity remain elusive. Recently, the National Human Genome Research Institute endorsed a multi-taxon genome-sequencing initiative that aims to gain insights into how multicellularity first evolved. This initiative (which we have termed UNICORN) will generate extensive genomic data from some of the closest extant unicellular relatives of both animals and fungi. (113)

Sachs, Joel and James Bull. Experimental Evolution of Conflict Mediation Between Genomes. Proceedings of the National Academy of Sciences. 102/390, 2005. Transitions to a new level of biological complexity requires cooperation among members, but a selection for individual advantage would seem to thwart that. In this study, an evolutionary life cycle for two bacteriophages inherently produced a salutary balance of interaction and independence. These findings, along with evidence from other stages, infer an innate propensity in emergent nature for cooperative behavior.

Specifically, the two phages evolved to copackage their genomes into one protein coat, ensuring cotransmission with each other and virtually eliminating conflict. (390) Our results parallel a variety of conflict mediation mechanisms existing in nature: evolution of reduced genomes in symbionts, cotransmission of partners, and obligate coexistence between cooperating species. (390)

Schopf, William. Cradle of Life. Princeton: Princeton University Press, 1999. A chatty exposition about Precambrian cellular organisms as “the discovery of earth’s earliest fossils,” with paleontologist Schopf as a major player.

Sebe-Pedros, Arnau, et al. The Origin of Metazoa: A Unicellular Perspective. Nature Reviews Genetics. 18/8, 2017. Not only has multicellularity evolved independently multiple times in eukaryotes, but each of these transitions also occurred at different time in the history of life. (499) Akin to van Gestel and Tarnita 2017, Weizmann Institute of Science, University of Queensland, and University of Barcelona researchers provide a wide and deep study of life’s persistent tendency to evolve into more complex unitary organisms. With an emphasis on a “unicellular urmetazoan genome,” a common cyclical process can be identified that occurs each many times. Figure 5 depicts “the origin of multicellularity as a transition from temporal to spatiotemporal cell differentiation.” Phylogenetic relationships of unicellular Holozoa and animals are then traced from fungi to mammals and we human beings whom collectively (another MET) in retrospect are able to learn all this. Some 150 references are cited in support.

The first animals evolved from an unknown single-celled ancestor in the Precambrian period. Recently, the identification and characterization of the genomic and cellular traits of the protists most closely related to animals have shed light on the origin of animals. Comparisons of animals with these unicellular relatives allow us to reconstruct the first evolutionary steps towards animal multicellularity. Here, we review the results of these investigations and discuss their implications for understanding the earliest stages of animal evolution, including the origin of metazoan genes and genome function. (Abstract)

Simpson, Carl. Adaptation to a Viscous Snowball Earth Ocean as a Path to Complex Multicellularity. American Naturalist. 198.5, 2021. As our collaborative studies proceed to reconstruct past planetary environs from which we all arose, the University of Colorado Museum of Natural History geobiologist (search) presents a thorough scenario of colder, glacial and milder, conducive phases that have affected complex organisms. It again amazes that our sapient issue can turn and retrace such a circuitous, stressful course, which reveals how chancy and perilous it was. See also Reproductive Innovations and Pulsed Rise in Plant Complexity by A. Leslie, C. Simpson and L. Mander in Science (373/1308, 2021).

Animals, fungi, and algae with complex multicellular bodies all evolved independently from unicellular ancestors. The early history of these major eukaryotic multicellular clades co-occur with an extreme phase of global glaciations known as the Snowball Earth. Here, I propose that the long-term loss of low-viscosity environments due to several rounds global glaciation drove the multiple origins of complex multicellularity in eukaryotes and the subsequent radiation of complex multicellular groups into previously unoccupied niches. In this scenario, life adapts to Snowball Earth oceans by evolving larger bodies and faster speeds for high-viscosity seawater. Warm, low-viscosity seawater returned with the melting of the glaciers which gave rise to new vital complexities. (Abstract)

Simpson, Carl. The Evolutionary History of Division of Labour. Proceedings of the Royal Society B. 279/116, 2012. A Duke University biologist contributes to the recognition that living systems at any and all scales evolve and emerge in complexity and cognizance by virtue of a reciprocal diversification of functional roles, as they proceed with a further formation of a bounded, individuated whole.

Functional specialization, or division of labour (DOL), of parts within organisms and colonies is common in most multi-cellular, colonial and social organisms, but it is far from ubiquitous. Several mechanisms have been proposed to explain the evolutionary origins of DOL; the basic feature common to all of them is that functional differences can arise easily. These mechanisms cannot explain the many groups of colonial and social animals that exhibit no DOL despite up to 500 million years of evolution. Here, I propose a new hypothesis, based on a multi-level selection theory, which predicts that a reproductive DOL is required to evolve prior to subsequent functional specialization. I test this hypothesis using a dataset consisting of the type of DOL for living and extinct colonial and social animals. The frequency distribution of DOL and the sequence of its acquisition confirm that reproductive specialization evolves prior to functional specialization. A corollary of this hypothesis is observed in colonial, social and also within multi-cellular organisms; those species without a reproductive DOL have a smaller range of internal variation, in terms of the number of polymorphs or cell types, than species with a reproductive DOL. (Abstract, 116)

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