Abstract
This article reviews the production of different phenotypes from the same genotype in the same environment by stochastic cellular events, nonlinear mechanisms during patterning and morphogenesis, and probabilistic self-reinforcing circuitries in the adult life. These aspects of phenotypic variation are summarized under the term ‘stochastic developmental variation’ (SDV) in the following. In the past, SDV has been viewed primarily as a nuisance, impairing laboratory experiments, pharmaceutical testing, and true-to-type breeding. This article also emphasizes the positive biological effects of SDV and discusses implications for genotype-to-phenotype mapping, biological individuation, ecology, evolution, and applied biology. There is strong evidence from experiments with genetically identical organisms performed in narrowly standardized laboratory set-ups that SDV is a source of phenotypic variation in its own right aside from genetic variation and environmental variation. It is obviously mediated by molecular and higher-order epigenetic mechanisms. Comparison of SDV in animals, plants, fungi, protists, bacteria, archaeans, and viruses suggests that it is a ubiquitous and phylogenetically old phenomenon. In animals, it is usually smallest for morphometric traits and highest for life history traits and behaviour. SDV is thought to contribute to phenotypic diversity in all populations but is particularly relevant for asexually reproducing and genetically impoverished populations, where it generates individuality despite genetic uniformity. In each generation, SDV produces a range of phenotypes around a well-adapted target phenotype, which is interpreted as a bet-hedging strategy to cope with the unpredictability of dynamic environments. At least some manifestations of SDV are heritable, adaptable, selectable, and evolvable, and therefore, SDV may be seen as a hitherto overlooked evolution factor. SDV is also relevant for husbandry, agriculture, and medicine because most pathogens are asexuals that exploit this third source of phenotypic variation to modify infectivity and resistance to antibiotics. Since SDV affects all types of organisms and almost all aspects of life, it urgently requires more intense research and a better integration into biological thinking.
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Acknowledgements
The author is grateful to the following colleagues for providing pictures and information: Martin Ackermann (Zurich), Gregory S Archer (Davis), Gregory A Babitt (Rochester), Jürgen Berger and Ralf J Sommer (Tübingen), Brian Bagatto (Akron), John Daugman (Cambridge), Jan Dijksterhuis and Han Wösten (Utrecht), David Fenwick (Penzance), Theodore H Friend (College Station), Mikael Häggström (Uppsala), David S Haskell (Sewanee), John F Hasler (Bogart), Anil K Jain (East Lansing), Nikos C Kyrpides (Walnut Creek), Winfried Lampert (Plön), Eric C Lai (New York), David IK Martin (Oakland), Erskine L Palmer and Russell Regnery (Atlanta), Soo-Bong Park (Suwon), Jorge A Piedrahita (Raleigh), Pascal Radtke (Düsseldorf), George E Seidel Jr (Fort Collins), Michael R Strand (Athens), Stefano Tiozzo (Villefranche-sur-Mer), Rogier R van Vugt (Leiden), Larry Wadsworth and Mark Westhusin (College Station), and Alex Wild (Champaign-Urbana). Many thanks also to Vidyanand Nanjundiah (Bangalore) and an anonymous referee for constructive criticism and numerous suggestions that helped to improve the manuscript.
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[Vogt G 2015 Stochastic developmental variation, an epigenetic source of phenotypic diversity with far-reaching biological consequences. J. Biosci. 40 1–46] DOI 10.1007/s12038-015-9506-8
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Vogt, G. Stochastic developmental variation, an epigenetic source of phenotypic diversity with far-reaching biological consequences. J Biosci 40, 159–204 (2015). https://doi.org/10.1007/s12038-015-9506-8
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DOI: https://doi.org/10.1007/s12038-015-9506-8