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      Biphasic patterns of diversification and the emergence of modules

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          Abstract

          The intricate molecular and cellular structure of organisms converts energy to work, which builds and maintains structure. Evolving structure implements modules, in which parts are tightly linked. Each module performs characteristic functions. In this work we propose that a module can emerge through two phases of diversification of parts. Early in the first phase of this biphasic pattern, the parts have weak linkage—they interact weakly and associate variously. The parts diversify and compete. Under selection for performance, interactions among the parts increasingly constrain their structure and associations. As many variants are eliminated, parts self-organize into modules with tight linkage. Linkage may increase in response to exogenous stresses as well as endogenous processes. In the second phase of diversification, variants of the module and its functions evolve and become new parts for a new cycle of generation of higher-level modules. This linkage hypothesis can interpret biphasic patterns in the diversification of protein domain structure, RNA and protein shapes, and networks in metabolism, codes, and embryos, and can explain hierarchical levels of structural organization that are widespread in biology.

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          Most cited references45

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          Data growth and its impact on the SCOP database: new developments

          The Structural Classification of Proteins (SCOP) database is a comprehensive ordering of all proteins of known structure, according to their evolutionary and structural relationships. The SCOP hierarchy comprises the following levels: Species, Protein, Family, Superfamily, Fold and Class. While keeping the original classification scheme intact, we have changed the production of SCOP in order to cope with a rapid growth of new structural data and to facilitate the discovery of new protein relationships. We describe ongoing developments and new features implemented in SCOP. A new update protocol supports batch classification of new protein structures by their detected relationships at Family and Superfamily levels in contrast to our previous sequential handling of new structural data by release date. We introduce pre-SCOP, a preview of the SCOP developmental version that enables earlier access to the information on new relationships. We also discuss the impact of worldwide Structural Genomics initiatives, which are producing new protein structures at an increasing rate, on the rates of discovery and growth of protein families and superfamilies. SCOP can be accessed at http://scop.mrc-lmb.cam.ac.uk/scop.
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            Consciousness as integrated information: a provisional manifesto.

            The integrated information theory (IIT) starts from phenomenology and makes use of thought experiments to claim that consciousness is integrated information. Specifically: (i) the quantity of consciousness corresponds to the amount of integrated information generated by a complex of elements; (ii) the quality of experience is specified by the set of informational relationships generated within that complex. Integrated information (Phi) is defined as the amount of information generated by a complex of elements, above and beyond the information generated by its parts. Qualia space (Q) is a space where each axis represents a possible state of the complex, each point is a probability distribution of its states, and arrows between points represent the informational relationships among its elements generated by causal mechanisms (connections). Together, the set of informational relationships within a complex constitute a shape in Q that completely and univocally specifies a particular experience. Several observations concerning the neural substrate of consciousness fall naturally into place within the IIT framework. Among them are the association of consciousness with certain neural systems rather than with others; the fact that neural processes underlying consciousness can influence or be influenced by neural processes that remain unconscious; the reduction of consciousness during dreamless sleep and generalized seizures; and the distinct role of different cortical architectures in affecting the quality of experience. Equating consciousness with integrated information carries several implications for our view of nature.
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              Robustness and evolvability: a paradox resolved.

              Understanding the relationship between robustness and evolvability is key to understand how living things can withstand mutations, while producing ample variation that leads to evolutionary innovations. Mutational robustness and evolvability, a system's ability to produce heritable variation, harbour a paradoxical tension. On one hand, high robustness implies low production of heritable phenotypic variation. On the other hand, both experimental and computational analyses of neutral networks indicate that robustness enhances evolvability. I here resolve this tension using RNA genotypes and their secondary structure phenotypes as a study system. To resolve the tension, one must distinguish between robustness of a genotype and a phenotype. I confirm that genotype (sequence) robustness and evolvability share an antagonistic relationship. In stark contrast, phenotype (structure) robustness promotes structure evolvability. A consequence is that finite populations of sequences with a robust phenotype can access large amounts of phenotypic variation while spreading through a neutral network. Population-level processes and phenotypes rather than individual sequences are key to understand the relationship between robustness and evolvability. My observations may apply to other genetic systems where many connected genotypes produce the same phenotypes.
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                Author and article information

                Journal
                Front Genet
                Front Genet
                Front. Gene.
                Frontiers in Genetics
                Frontiers Media S.A.
                1664-8021
                03 July 2012
                07 August 2012
                2012
                : 3
                : 147
                Affiliations
                [1] 1simpleDepartment of Cell and Developmental Biology, University of Illinois Urbana-Champaign, IL, USA
                [2] 2simpleEvolutionary Bioinformatics Laboratory, Department of Crop Sciences, University of Illinois Urbana, IL, USA
                [3] 3simpleInstitute for Genomic Biology, University of Illinois Urbana-Champaign, IL, USA
                Author notes

                Edited by: Firas H. Kobeissy, University of Florida, USA

                Reviewed by: Nehme Hachem, AUB, Canada; Bilal Fadlallah, University of Florida, USA

                *Correspondence: Gustavo Caetano-Anollés, Evolutionary Bioinformatics Laboratory, Department of Crop Sciences, University of Illinois at Urbana-Champaign, 1101 W. Peabody Drive, Urbana, IL 61801, USA. e-mail: gca@ 123456illinois.edu

                This article was submitted to Frontiers in Systems Biology, a specialty of Frontiers in Genetics.

                Article
                10.3389/fgene.2012.00147
                3413098
                22891076
                c311e0cf-c268-443d-95b6-2223032eec0f
                Copyright © 2012 Mittenthal, Caetano-Anollés and Caetano-Anollés.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.

                History
                : 22 May 2012
                : 19 July 2012
                Page count
                Figures: 4, Tables: 0, Equations: 0, References: 70, Pages: 12, Words: 9976
                Categories
                Psychiatry
                Hypothesis and Theory Article

                Genetics
                biphasic hourglass,competitive optimization,module,linkage,diversification
                Genetics
                biphasic hourglass, competitive optimization, module, linkage, diversification

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