功能约束下的鲁棒性：果蝇神经发生中时间表达的遗传网络
Robustness under functional constraint: The genetic network for temporal expression in Drosophila neurogenesis
摘要：
基因表达的精确时间协调对于许多发育过程是至关重要的。发育生物学的一个中心问题是如何有力地控制这种协调的表达模式。在果蝇中枢神经系统的胚胎发育过程中，神经干细胞（称为神经母细胞）按一定的顺序表达一组基因，从而产生细胞类型的多样性。通过产生这些基因的所有可能的调控网络，并检查它们的表达动力学数值，我们确定必要的法规和预测一个未知的因素，以重现已知的表达谱所造成的功能丧失或过度表达的基因在体内，以及在野生型。然后，我们评估的稳定性，实际的果蝇网络的顺序表达。该网络显示出最高的鲁棒性，对参数变化和基因表达波动之间的可能的网络，再现的表达谱。我们提出了一个由三种规则组成的调节模块，负责精确的顺序表达。本研究提出了一个基本原则，生物系统如何鲁棒设计功能约束下。
英文摘要：
Precise temporal coordination of gene expression is crucial for many developmental processes. One central question in developmental biology is how such coordinated expression patterns are robustly controlled. During embryonic development of the Drosophila central nervous system, neural stem cells called neuroblasts sequentially express a group of genes in a definite order, which generates the diversity of cell types. By producing all possible regulatory networks of these genes and examining their expression dynamics numerically, we identify requisite regulations and predict an unknown factor to reproduce known expression profiles caused by loss-of-function or overexpression of the genes in vivo, as well as in the wild type. We then evaluate the stability of the actual Drosophila network for sequential expression. This network shows the highest robustness against parameter variations and gene expression fluctuations among the possible networks that reproduce the expression profiles. We propose a regulatory module composed of three kinds of regulations which is responsible for precise sequential expression. The present study suggests an underlying principle on how biological systems are robustly designed under functional constraint.
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