Adams Daniel L, Horton Jonathan C
Beckman Vision Center, University of California, San Francisco, San Francisco, California 94143-0730, USA.
J Neurosci. 2003 Jul 9;23(14):5984-97. doi: 10.1523/JNEUROSCI.23-14-05984.2003.
The blood vessels that nourish the inner retina cast shadows on photoreceptors, creating "angioscotomas" in the visual field. We have found the representations of angioscotomas in striate cortex of the squirrel monkey. They were detected in 9 of 12 normal adult animals by staining flatmounts for cytochrome oxidase activity after enucleation of one eye. They appeared as thin profiles in layer 4C radiating from the blind spot representation. Angioscotomas can be regarded as a local form of amblyopia. After birth, when light strikes the retina, photoreceptors beneath blood vessels are denied normal visual stimulation. This deprivation induces remodeling of geniculocortical afferents in a distribution that corresponds to the retinal vascular tree. Angioscotoma representations were most obvious in monkeys with fine ocular dominance columns and were invisible in monkeys with large, well segregated columns. In monkeys without columns, their width corresponded faithfully to the inducing retinal shadow, making it possible to calculate the minimum shadow required to produce a cortical representation. The "amblyogenic threshold" was calculated as the fraction of the pupil area eclipsed to trigger remodeling of geniculocortical afferents. It was found to be constant over retinal eccentricity, vessel size, and shadow size. Ambliogenic shadows only three to four cones wide were sufficient to generate a cortical representation, testifying to the remarkable precision of the cortical map. The representations of retinal blood vessels separated by only 0.65 degrees were resolvable in the cortex, yielding an upper limit on cortical resolution of 340 microm in layer 4C.
滋养视网膜内层的血管在光感受器上投射阴影,在视野中形成“血管性暗点”。我们在松鼠猴的纹状皮质中发现了血管性暗点的表征。在摘除一只眼睛后,通过对细胞色素氧化酶活性进行染色平片,在12只正常成年动物中的9只中检测到了它们。它们在4C层中表现为从盲点表征辐射出的细条纹。血管性暗点可被视为弱视的一种局部形式。出生后,当光线照射视网膜时,血管下方的光感受器无法获得正常的视觉刺激。这种剥夺会导致膝状体皮质传入纤维在与视网膜血管树相对应的分布中发生重塑。血管性暗点表征在具有精细眼优势柱的猴子中最为明显,而在具有大的、分隔良好的柱的猴子中则不可见。在没有柱的猴子中,它们的宽度与诱导视网膜阴影精确对应,从而有可能计算出产生皮质表征所需的最小阴影。“致弱视阈值”被计算为使膝状体皮质传入纤维发生重塑的瞳孔面积遮挡比例。发现它在视网膜离心率、血管大小和阴影大小方面是恒定的。仅三到四个视锥细胞宽的致弱视阴影就足以产生皮质表征,这证明了皮质图谱具有非凡的精确性。在皮质中可分辨出仅相隔0.65度的视网膜血管表征,在4C层中得出皮质分辨率的上限为340微米。