Sixty-five years after Turing first revealed the potential of systems with local activation and long-range inhibition to generate pattern, we have only recently begun to identify the biological elements that operate at many scales to generate periodic patterns in nature. In this Primer, we first review the theoretical framework provided by Turing, Meinhardt, and others that suggests how periodic patterns could self-organize in developing animals. This Primer was developed to provide context for recent studies that reveal how diverse molecular, cellular, and physical mechanisms contribute to the establishment of the periodic pattern of hair or feather buds in the developing skin. From an initial emphasis on trying to disambiguate which specific mechanism plays a primary role in hair or feather bud development, we are beginning to discover that multiple mechanisms may, in at least some contexts, operate together. While the emergence of the diverse mechanisms underlying pattern formation in specific biological contexts probably reflects the contingencies of evolutionary history, an intriguing possibility is that these mechanisms interact and reinforce each other, producing emergent systems that are more robust.

在图灵首次揭示具有局部激活和远距离抑制作用的系统产生模式的潜力后的65年，我们直到最近才开始鉴定可在许多规模上运行以在自然界中产生周期性模式的生物元素。在本入门手册中，我们首先回顾由Turing，Meinhardt等人提供的理论框架，该框架提出了周期性模式如何在发育中的动物中自我组织。开发该底漆可为最近的研究提供背景信息，这些研究揭示了各种分子，细胞和物理机制如何促进发育中的皮肤中毛发或羽毛芽周期性模式的建立。从最初试图消除歧义的特定机制在毛发或羽毛芽发育中起主要作用开始，我们开始发现，至少在某些情况下，多种机制可能会共同起作用。虽然在特定生物学背景下模式形成的各种机制的出现可能反映了进化史的偶然性，但一个引人入胜的可能性是，这些机制相互作用并相互加强，从而产生了更强大的涌现系统。