Valen Ragnhild, Eilertsen Mariann, Edvardsen Rolf Brudvik, Furmanek Tomasz, Rønnestad Ivar, van der Meeren Terje, Karlsen Ørjan, Nilsen Tom Ole, Helvik Jon Vidar
Department of Biology, University of Bergen, NO-5020 Bergen, Norway.
Institute of Marine Research, Nordnes, NO-5005 Bergen, Norway.
Dev Biol. 2016 Aug 15;416(2):389-401. doi: 10.1016/j.ydbio.2016.06.041. Epub 2016 Jun 30.
Unlike in mammals, persistent postembryonic retinal growth is a characteristic feature of fish, which includes major remodeling events that affect all cell types including photoreceptors. Consequently, visual capabilities change during development, where retinal sensitivity to different wavelengths of light (photopic vision), -and to limited photons (scotopic vision) are central capabilities for survival. Differently from well-established model fish, Atlantic cod has a prolonged larval stage where only cone photoreceptors are present. Rods do not appear until juvenile transition (metamorphosis), a hallmark of indirect developing species. Previously we showed that whole gene families of lws (red-sensitive) and sws1 (UV-sensitive) opsins have been lost in cod, while rh2a (green-sensitive) and sws2 (blue-sensitive) genes have tandem duplicated. Here, we provide a comprehensive characterization of a two-step developing duplex retina in Atlantic cod. The study focuses on cone subtype dynamics and delayed rod neurogenesis and differentiation in all cod life stages. Using transcriptomic and histological approaches we show that different opsins disappear in a topographic manner during development where central to peripheral retina is a key axis of expressional change. Early cone differentiation was initiated in dorso-temporal retina different from previously described in fish. Rods first appeared during initiation of metamorphosis and expression of the nuclear receptor transcription factor nr2e3-1, suggest involvement in rod specification. The indirect developmental strategy thus allows for separate studies of cones and rods development, which in nature correlates with visual changes linked to habitat shifts. The clustering of key retinal genes according to life stage, suggests that Atlantic cod with its sequenced genome may be an important resource for identification of underlying factors required for development and function of photopic and scotopic vision.
与哺乳动物不同,胚胎后视网膜持续生长是鱼类的一个特征,其中包括影响所有细胞类型(包括光感受器)的主要重塑事件。因此,视觉能力在发育过程中会发生变化,视网膜对不同波长光(明视觉)以及对有限光子(暗视觉)的敏感度是生存的核心能力。与成熟的模式鱼类不同,大西洋鳕鱼有一个延长的幼体阶段,在此阶段仅存在视锥光感受器。直到幼鱼过渡(变态)时才出现视杆细胞,这是间接发育物种的一个标志。此前我们表明,鳕鱼中lws(对红色敏感)和sws1(对紫外线敏感)视蛋白的整个基因家族已经丢失,而rh2a(对绿色敏感)和sws2(对蓝色敏感)基因发生了串联重复。在此,我们对大西洋鳕鱼两步发育的双重复合视网膜进行了全面表征。该研究聚焦于鳕鱼所有生命阶段视锥亚型动态以及视杆细胞神经发生和分化延迟的情况。通过转录组学和组织学方法,我们表明不同的视蛋白在发育过程中以一种拓扑方式消失,其中视网膜中央到周边是表达变化的关键轴。早期视锥细胞分化始于背颞侧视网膜,这与之前在鱼类中描述的情况不同。视杆细胞首先在变态开始时出现,核受体转录因子nr2e3-1的表达表明其参与视杆细胞的特化。这种间接发育策略因此允许对视锥细胞和视杆细胞的发育进行单独研究,这在自然界中与与栖息地转变相关的视觉变化相关。关键视网膜基因根据生命阶段的聚类表明,具有已测序基因组的大西洋鳕鱼可能是鉴定明视觉和暗视觉发育及功能所需潜在因素的重要资源。