Kuszak Jer R, Mazurkiewicz Mike, Zoltoski Rebecca
Department of Ophthalmology, Rush University Medical Center, Chicago, IL 60612, USA.
Mol Vis. 2006 Apr 3;12:251-70.
The purpose of this study was to use qualitative and quantitative structural data from nonprimate lenses with branched (Y and line) sutures to generate computer models (animations) of secondary fiber development and suture formation.
A minimum of 12-18 adult lenses/species (mice, cows, frogs, and rabbits) were used in this study. Lenses were analyzed by light (LM), transmission (TEM), and scanning electron microscopy (SEM). Fiber width, thickness, and length were ascertained from micrographs and by using formulations to calculate distances between degrees of latitude and longitude on asymmetrical oblate spheroids. This information was then used to create scale computer assisted drawings (CADs) of fibers at different stages of their development. The CADs were then placed on a timeline and animated to produce dynamic representations of secondary fiber development and growth.
Animating secondary fiber development and suture formation with the inclusion of quantifiable differences in fiber dimensions at progressive stages of their differentiation revealed the following: first, there is the presumption that fibers migrate, rotate, and elongate until they reach their sutural destinations is not likely to be correct. When developing fibers reach approximately 60-65% of their eventual total length, their migration and rotation is complete. The remaining fiber elongation (the production of end segments) occurs without either concomitant cellular migration or rotation. Second, it is presumed that suture branches originate peripherally and are then constructed sequentially until all of the branches come to confluence at the poles is also not likely to be correct. While suture branches do originate peripherally, if the rate of elongation is constant in the anterior and posterior directions (intrafiber elongation speed) and between developing fibers within a forming growth shell (interfiber elongation speed), then only a part of their construction proceeds sequentially toward the poles. A second suture branch origin will be established at the poles resulting in a short distal portion of suture branches being formed sequentially in the reverse direction. Suture formation will conclude when a long proximal and a short distal portion of branches come to confluence within unequal anterior and posterior polar cap regions. This segmented suture formation scheme will be more pronounced in line suture lenses than in Y suture lenses. Third, because lenses with branched sutures have growth shells consisting of fibers of unequal length, fiber maturation is likely to be initiated in these lenses before a growth shell as well as suture formation is completed and would proceed in distinct patterns over a period of time. This is in marked contrast to avian lens fiber maturation which does not begin until growth shell and suture (branchless umbilical) formation is completed and then occurs rapidly and essentially simultaneously across the entire growth shell.
Animations of secondary fiber development and suture formation based on quantitative analysis of electron micrographs reveals important novel aspects of these processes that have not been apparent from the results of previous mechanistic studies. The more complex schemes of fiber differentiation and suture formation presented herein are consistent with the notion that lens function (dynamic focusing) is interdependent on lens structure and physiology. The animations confirm that while all vertebrate lenses have a similar structure, differences in the level of their structural complexity established early in development and maintained throughout life can account for the varying amount of optical quality known to exist between species.
本研究的目的是利用来自具有分支(Y形和线状)缝线的非灵长类动物晶状体的定性和定量结构数据,生成次级纤维发育和缝线形成的计算机模型(动画)。
本研究使用了至少12 - 18个成年晶状体/物种(小鼠、牛、青蛙和兔子)。通过光学显微镜(LM)、透射电子显微镜(TEM)和扫描电子显微镜(SEM)对晶状体进行分析。从显微照片中确定纤维的宽度、厚度和长度,并使用公式计算不对称扁球体上不同纬度和经度之间的距离。然后利用这些信息创建不同发育阶段纤维的比例计算机辅助绘图(CAD)。接着将CAD放置在时间轴上并制作动画,以生成次级纤维发育和生长的动态表示。
通过在次级纤维发育和缝线形成的动画中纳入其分化各阶段纤维尺寸的可量化差异,揭示了以下内容:第一,认为纤维迁移、旋转并伸长直至到达其缝线目的地的推测不太可能正确。当发育中的纤维达到其最终总长度的约60 - 65%时,它们的迁移和旋转就完成了。剩余的纤维伸长(末端段的产生)在没有伴随细胞迁移或旋转的情况下发生。第二,认为缝线分支起源于周边,然后依次构建直至所有分支在极点汇合的推测也不太可能正确。虽然缝线分支确实起源于周边,但如果在前后方向(纤维内伸长速度)以及形成的生长壳内发育中的纤维之间(纤维间伸长速度)伸长速率恒定,那么它们的构建只有一部分会依次向极点进行。在极点会形成第二个缝线分支起源,导致缝线分支的短远端部分以相反方向依次形成。当分支的长近端和短远端部分在不等的前后极帽区域内汇合时,缝线形成结束。这种分段缝线形成方案在线状缝线晶状体中比在Y形缝线晶状体中更明显。第三,由于具有分支缝线的晶状体的生长壳由长度不等的纤维组成,纤维成熟可能在这些晶状体中在生长壳以及缝线形成完成之前就开始,并会在一段时间内以不同模式进行。这与鸟类晶状体纤维成熟形成显著对比,鸟类晶状体纤维成熟直到生长壳和缝线(无分支脐部)形成完成才开始,然后在整个生长壳内迅速且基本同时发生。
基于电子显微镜定量分析的次级纤维发育和缝线形成动画揭示了这些过程中重要的新方面,这些方面在以前的机制研究结果中并不明显。本文提出的更复杂的纤维分化和缝线形成方案与晶状体功能(动态聚焦)依赖于晶状体结构和生理学的观点一致。这些动画证实,虽然所有脊椎动物晶状体都具有相似的结构,但在发育早期建立并在整个生命过程中维持的结构复杂程度差异可以解释已知存在于不同物种之间的不同光学质量。