Mills Margaret G, Nuckels Richard J, Parichy David M
Department of Biology, Institute for Stem Cell and Regenerative Medicine, University of Washington, Box 351800, Seattle WA 98195-1800, USA.
Development. 2007 Mar;134(6):1081-90. doi: 10.1242/dev.02799. Epub 2007 Feb 7.
The cellular bases for evolutionary changes in adult form remain largely unknown. Pigment patterns of Danio fishes are a convenient system for studying these issues because of their diversity and accessibility and because one species, the zebrafish D. rerio, is a model organism for biomedical research. Previous studies have shown that in zebrafish, stripes form by migration and differentiation of distinct populations of melanophores: early metamorphic (EM) melanophores arise widely dispersed and then migrate into stripes, whereas late metamorphic (LM) melanophores arise already within stripes. EM melanophores require the kit receptor tyrosine kinase, as kit mutants lack these cells but retain LM melanophores, which form a residual stripe pattern. To see if similar cell populations and genetic requirements are present in other species, we examined D. albolineatus, which has relatively few, nearly uniform melanophores. We isolated a D. albolineatus kit mutant and asked whether residual, LM melanophores develop in this species, as in D. rerio. We found that kit mutant D. albolineatus lack EM melanophores, yet retain LM melanophores. Histological analyses further show that kit functions during a late step in metamorphic melanophore development in both species. Interestingly, kit mutant D. albolineatus develop a striped melanophore pattern similar to kit mutant D. rerio, revealing latent stripe-forming potential in this species, despite its normally uniform pattern. Comparisons of wild types and kit mutants of the two species further show that species differences in pigment pattern reflect: (1) changes in the behavior of kit-dependent EM melanophores that arise in a dispersed pattern and then migrate into stripes in D. rerio, but fail to migrate in D. albolineatus; and (2) a change in the number of kit-independent LM melanophores that arise already in stripes and are numerous in D. rerio, but few in D. albolineatus. Our results show how genetic analyses of a species closely related to a biomedical model organism can reveal both conservatism and innovation in developmental mechanisms underlying evolutionary changes in adult form.
成体形态进化变化的细胞基础在很大程度上仍然未知。由于其多样性和易获取性,以及斑马鱼D. rerio这一物种是生物医学研究的模式生物,因此斑马鱼的色素模式是研究这些问题的一个便捷系统。先前的研究表明,在斑马鱼中,条纹是由不同种群的黑素细胞迁移和分化形成的:早期变态(EM)黑素细胞广泛分散产生,然后迁移到条纹中,而晚期变态(LM)黑素细胞则在条纹内已经产生。EM黑素细胞需要kit受体酪氨酸激酶,因为kit突变体缺乏这些细胞,但保留了LM黑素细胞,它们形成了残余的条纹模式。为了了解其他物种中是否存在类似的细胞群体和遗传需求,我们研究了拥有相对较少且近乎均匀黑素细胞的白线丹鱼(D. albolineatus)。我们分离出了一个白线丹鱼kit突变体,并询问在该物种中是否像在斑马鱼中一样会发育出残余的LM黑素细胞。我们发现,kit突变体白线丹鱼缺乏EM黑素细胞,但保留了LM黑素细胞。组织学分析进一步表明,在这两个物种中,kit在变态黑素细胞发育的后期发挥作用。有趣的是,kit突变体白线丹鱼发育出了与kit突变体斑马鱼相似的条纹状黑素细胞模式,这揭示了该物种潜在的条纹形成潜力,尽管其正常模式是均匀的。对这两个物种的野生型和kit突变体的比较进一步表明,色素模式的物种差异反映了:(1)依赖kit的EM黑素细胞行为的变化,这些细胞在斑马鱼中以分散模式出现,然后迁移到条纹中,但在白线丹鱼中未能迁移;(2)不依赖kit的LM黑素细胞数量的变化,这些细胞在条纹中已经出现,在斑马鱼中数量众多,但在白线丹鱼中很少。我们的结果表明,对与生物医学模式生物密切相关的物种进行遗传分析,如何能够揭示成体形态进化变化背后发育机制中的保守性和创新性。