Reuter T E, White R H, Wald G
J Gen Physiol. 1971 Oct;58(4):351-71. doi: 10.1085/jgp.58.4.351.
Though it had been supposed earlier that the bullfrog undergoes a virtually complete metamorphosis of visual systems from vitamin A(2) and porphyropsin in the tadpole to vitamin A(1) and rhodopsin in the adult, the present observations show that the retina of the adult frog may contain as much as 30-40% porphyropsin, all of it segregated in the dorsal zone. The most dorsal quarter of the adult retina may contain 81-89% porphyropsin mixed with a minor amount of rhodopsin; the ventral half contains only rhodopsin. Further, the dorsal zone contains a two to three times higher concentration of visual pigments than the ventral retina. The pigment epithelium underlying the retina contains a corresponding distribution of vitamins A(1) and A(2), predominantly vitamin A(2) in the dorsal pigment epithelium, exclusively vitamin A(1) in the ventral zone. The retina accepts whatever vitamin A the pigment epithelium provides it with, and turns it into the corresponding visual pigment. Thus, a piece of light-adapted dorsal retina laid back on ventral pigment epithelium regenerates rhodopsin, whereas a piece of light-adapted ventral retina laid back on dorsal pigment epithelium regenerates predominantly porphyropsin. Vitamin A(2) must be made from vitamin A(1), by dehydrogenation at the 3,4-bond in the ring. This conversion must occur in the pigment epithelium, presumably through the action of a vitamin A-3,4-dehydrogenase. The essential change at metamorphosis is to make much less of this dehydrogenase, and to sequester it in the dorsal pigment epithelium. Some adult bullfrogs, perhaps characteristically taken in the summer, contain very little porphyropsin-only perhaps 5%-still sequestered in the dorsal retina. The gradient of light over the retinal surface has little if any effect on this distribution. The greater density of visual pigments in the dorsal retina, and perhaps also-although this is less clear-the presence of porphyropsin in this zone, has some ecological importance in increasing the retinal sensitivity to the dimmer and, on occasion, redder light received from below.
尽管早期人们认为牛蛙的视觉系统经历了从蝌蚪期的维生素A(2)和视紫蓝质到成年期的维生素A(1)和视紫红质的几乎完全变态,但目前的观察表明,成年青蛙的视网膜可能含有高达30 - 40%的视紫蓝质,所有这些都集中在背侧区域。成年视网膜最靠背部的四分之一可能含有81 - 89%的视紫蓝质并混有少量视紫红质;腹侧半部只含有视紫红质。此外,背侧区域的视觉色素浓度比腹侧视网膜高三到两倍。视网膜下方的色素上皮含有维生素A(1)和A(2)的相应分布,背侧色素上皮中主要是维生素A(2),腹侧区域则只有维生素A(1)。视网膜接受色素上皮提供给它的任何维生素A,并将其转化为相应的视觉色素。因此,一块适应光照的背侧视网膜放在腹侧色素上皮上会再生视紫红质,而一块适应光照的腹侧视网膜放在背侧色素上皮上则主要再生视紫蓝质。维生素A(2)必须由维生素A(1)通过环中3,4键的脱氢作用合成。这种转化一定发生在色素上皮中,大概是通过维生素A - 3,4 - 脱氢酶的作用。变态时的关键变化是使这种脱氢酶的产量大大减少,并将其隔离在背侧色素上皮中。一些成年牛蛙,可能典型地是在夏季捕获的,含有很少的视紫蓝质——可能只有5%——仍然集中在背侧视网膜中。视网膜表面的光照梯度对这种分布几乎没有影响。背侧视网膜中视觉色素的密度更高,而且也许——尽管这一点不太清楚——该区域视紫蓝质的存在,在提高视网膜对从下方接收到的较暗且有时较红的光的敏感度方面具有一定的生态重要性。