Pettigrew J D, Wallman J, Wildsoet C F
Vision, Touch and Hearing Research Centre, University of Queensland, St. Lucia 4067, Australia.
Nature. 1990 Jan 25;343(6256):362-3. doi: 10.1038/343362a0.
THE evolution of the eye is constrained by two conflicting requirements--good vascular perfusion of the retina, and an optical path through the retina that is unobstructed by blood vessels. Birds are interesting in that they have higher metabolic rates and thicker retinas than mammals, but have no retinal blood vessels. Nutrients and oxygen must thus reach the neurons of the inner retina either from the choroid through 300 micron of metabolically very active retina, or from the pecten, a pleated vascular structure protruding from the head of the optic nerve into the vitreous chamber, and more than a centimetre away from some retinal neurons. Despite the diffusional distance involved, several lines of evidence indicate that the pecten is the primary source of nutrients for the inner retina: the presence of an oxygen gradient from pecten to retina, the large surface area produced by macroscopic folds and by microscopic infoldings of the luminal and external surfaces of the capillary endothelium, extrusion of circulating fluorescein, high content of carbonic anhydrase and alkaline phosphatase, and retinal impairments after pecten ablation. Another peculiarity of birds, their saccadic oscillations, occur with a large cyclotor-sional component during every saccadic eye movement. In different species, saccades, which occur at intervals of 0.5-40 s, have up to 13 oscillations with frequencies of 15-30 Hz and ampliá-tudes of about 10 degrees. Therefore, as much as 12% of some birds' total viewing time may be subject to the image instability caused by the oscillations. Using fluorescein angiography, we show here that during every saccade, the pecten acts as an agitator which propels perfusate towards the central retina much more effectively than is observed during intersaccadic intervals.
眼睛的进化受到两个相互矛盾的需求的限制——视网膜要有良好的血管灌注,以及穿过视网膜的光路不能被血管阻挡。鸟类很有趣,因为它们的代谢率比哺乳动物高,视网膜也更厚,但却没有视网膜血管。因此,营养物质和氧气必须从脉络膜穿过300微米代谢非常活跃的视网膜,或者从栉膜到达视网膜内层神经元,栉膜是一种从视神经头部伸入玻璃体腔的褶皱血管结构,距离一些视网膜神经元超过一厘米。尽管存在扩散距离,但有几条证据表明栉膜是视网膜内层营养物质的主要来源:从栉膜到视网膜存在氧梯度,毛细血管内皮细胞腔面和外表面的宏观褶皱和微观内褶产生的大表面积,循环荧光素的挤出,碳酸酐酶和碱性磷酸酶的高含量,以及栉膜切除后的视网膜损伤。鸟类的另一个特点是它们的扫视振荡,在每次眼球扫视运动中都有很大的旋转扭转成分。在不同物种中,扫视间隔为0.5 - 40秒,有多达13次振荡,频率为15 - 30赫兹,振幅约为10度。因此,一些鸟类的总观察时间中多达12%可能会受到振荡引起的图像不稳定的影响。我们在此通过荧光素血管造影显示,在每次扫视期间,栉膜起到搅拌器的作用,比在扫视间隔期间更有效地将灌注液推向视网膜中央。