Lisney Thomas J, Collin Shaun P
Oceans Graduate School, The University of Western Australia, Crawley, Washington, Australia,
The Oceans Institute, The University of Western Australia, Crawley, Washington, Australia,
Brain Behav Evol. 2018;92(3-4):97-116. doi: 10.1159/000495285. Epub 2019 Jan 24.
Little is known about the visual systems of large baleen whales (Mysticeti: Cetacea). In this study, we investigate eye morphology and the topographic distribution of retinal ganglion cells (RGCs) in two species of mysticete, Bryde's whale (Balaenoptera edeni) and the humpback whale (Megaptera novaeanglia). Both species have large eyes characterised by a thickened cornea, a heavily thickened sclera, a highly vascularised fibro-adipose bundle surrounding the optic nerve at the back of the eye, and a reflective blue-green tapetum fibrosum. Using stereology and retinal whole mounts, we estimate a total of 274,268 and 161,371 RGCs in the Bryde's whale and humpback whale retinas, respectively. Both species have a similar retinal topography, consisting of nasal and temporal areas of high RGC density, suggesting that having higher visual acuity in the anterior and latero-caudal visual fields is particularly important in these animals. The temporal area is larger in both species and contains the peak RGC densities (160 cells mm-2 in the humpback whale and 200 cells mm-2 in Bryde's whale). In the Bryde's whale retina, the two high-density areas are connected by a weak centro-ventral visual streak, but such a specialisation is not evident in the humpback whale. Measurements of RGC soma area reveal that although the RGCs in both species vary substantially in size, RGC soma area is inversely proportional to RGC density, with cells in the nasal and temporal high-density areas being relatively more homogeneous in size compared to the RGCs in the central retina and the dorsal and ventral retinal periphery. Some of the RGCs were very large, with soma areas of over 2,000 µm2. Using peak RGC density and eye axial diameter (Bryde's whale: 63.5 mm; humpback whale: 48.5 mm), we estimated the peak anatomical spatial resolving power in water to be 4.8 cycles/degree and 3.3 cycles/degree in the Bryde's whale and the humpback whale, respectively. Overall, our findings for these two species are very similar to those reported for other species of cetaceans. This indicates that, irrespective of the significant differences in body size and shape, behavioural ecology and feeding strategy between mysticetes and odontocetes (toothed whales), cetacean eyes are adapted to vision in dim light and adhere to a common "bauplan" that evolved prior to the divergence of the two cetacean parvorders (Odontoceti and Mysticeti) over 30 million years ago.
对于大型须鲸(须鲸亚目:鲸目)的视觉系统,我们所知甚少。在本研究中,我们调查了两种须鲸——布氏鲸(Balaenoptera edeni)和座头鲸(Megaptera novaeanglia)的眼睛形态以及视网膜神经节细胞(RGC)的地形图分布。这两个物种的眼睛都很大,其特征包括角膜增厚、巩膜显著增厚、在眼球后部视神经周围有高度血管化的纤维脂肪束,以及一个反射蓝绿色的纤维性反光层。我们使用体视学和视网膜整装片,分别估计布氏鲸和座头鲸视网膜中RGC的总数为274,268个和161,371个。这两个物种都有相似的视网膜地形图,由RGC密度较高的鼻侧和颞侧区域组成,这表明在这些动物中,在前侧和后外侧视野具有更高的视敏度尤为重要。两个物种的颞侧区域都更大,且包含RGC密度峰值(座头鲸为160个细胞/mm²,布氏鲸为200个细胞/mm²)。在布氏鲸视网膜中,两个高密度区域由一条微弱的中央腹侧视觉条纹相连,但在座头鲸中这种特化并不明显。对RGC胞体面积的测量表明,尽管两个物种的RGC大小差异很大,但RGC胞体面积与RGC密度成反比,与中央视网膜以及背侧和腹侧视网膜周边的RGC相比,鼻侧和颞侧高密度区域的细胞大小相对更均匀。一些RGC非常大,胞体面积超过2000 µm²。利用RGC密度峰值和眼轴直径(布氏鲸:63.5毫米;座头鲸:48.5毫米),我们估计布氏鲸和座头鲸在水中的峰值解剖空间分辨能力分别为4.8周/度和3.3周/度。总体而言,我们对这两个物种的研究结果与其他鲸类物种的报道非常相似。这表明,尽管须鲸和齿鲸(有齿鲸类)在体型、形状、行为生态学和觅食策略上存在显著差异,但鲸类的眼睛适应于在弱光下视觉,并且遵循一种在3000多万年前两个鲸类小目(齿鲸亚目和须鲸亚目)分化之前就已演化出来的共同“基本结构”。