Stochastic developmental variation, an epigenetic source of phenotypic diversity with far-reaching biological consequences
Stochastic developmental variation, an epigenetic source of phenotypic diversity with far-reaching biological consequences
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随机发育变异，表型多样性的表观遗传来源，具有深远的生物学影响
作者：
G. Vogt
本文综述了相同基因型在相同环境下通过随机细胞事件、模式和形态发生过程中的非线性机制以及成人生活中的概率自我强化回路产生不同表型的研究进展。以下将表型变异的这些方面概括为“随机发育变异”（SDV）。在过去，SDV主要被视为一种讨厌的东西，损害实验室实验、药物测试和真正的育种。本文还强调了SDV的积极生物学效应，并讨论了SDV在基因型-表型定位、生物个体化、生态学、进化和应用生物学方面的意义。在严格标准化的实验室设置中对基因相同的生物体进行的实验有强有力的证据表明，除了遗传变异和环境变异外，SDV本身也是表型变异的一个来源。这显然是由分子和高阶表观遗传机制介导的。SDV在动物、植物、真菌、原生生物、细菌、古生菌和病毒中的比较表明，它是一种普遍存在的、在系统发育上古老的现象。在动物中，通常形态特征最小，生活史特征和行为最高。SDV被认为有助于所有人群的表型多样性，但与无性繁殖和遗传贫乏的人群特别相关，在这些人群中，尽管遗传一致性，但它产生了个性。在每一代中，SDV围绕一个适应良好的目标表型产生一系列表型，这被解释为一种下注对冲策略，以应对动态环境的不可预测性。至少SDV的一些表现是可遗传的、可适应的、可选择的和进化的，因此，SDV可能被视为一个迄今为止被忽视的进化因素。SDV也与畜牧业、农业和医药有关，因为大多数病原体是无性的，它们利用这第三种表型变异来源来改变传染性和对抗生素的耐药性。由于SDV影响所有类型的生物体和几乎生命的所有方面，因此迫切需要对其进行更深入的研究，并更好地融入生物学思维。
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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