在2009年9月21日的Journal of Cell Biology的一篇報道中,，Nowak等人介紹了上皮細胞脂膜中維持細胞形狀的蛋白質網絡,，以及在老鼠晶狀體中該網絡維持細胞形狀和有序排列的機制,。
蛋白網絡中的血影蛋白（Spectrin），F(xiàn)肌動蛋白（F-actin）以及其他的相關蛋白能夠支撐和定形紅細胞的脂膜,。在其他類型的細胞中同樣存在著相似的網絡,，包括晶體上皮細胞。脂膜骨架中的肌絲（Actin filament）,，是通過網絡中的相關蛋白固定的,，如肌動蛋白結合蛋白（actin-binding protein）的原肌球蛋白家族和原肌球調節(jié)蛋白家族的成員。
在缺少肌球調節(jié)蛋白1的老鼠中,，其晶體上皮細胞質膜骨架中的gamma原肌球蛋白同樣也會缺失,。F肌動蛋白和血影蛋白雖然仍在細胞的脂膜中，以往連續(xù)的網絡中卻出現(xiàn)了裂口,，表明這兩個肌動蛋白結合蛋白穩(wěn)固了肌絲,，而肌絲對網絡來數是必需的。電子顯微掃描顯示,，缺少肌球調節(jié)蛋白1的細胞出現(xiàn)形狀扭曲以及不規(guī)則的排列,。
Velia Fowler解釋說，血影肌動蛋白網絡的破壞改變了相鄰細胞間的相互粘合,，導致細胞形狀改變和晶狀體相關的機械應力混亂,。（生物谷Bioon.com）
生物谷推薦原始出處：
The Journal of Cell Biology, Vol. 186, No. 6, 915-928 doi:10.1083/jcb.200905065
Tropomodulin1 is required for membrane skeleton organization and hexagonal geometry of fiber cells in the mouse lens
Roberta B. Nowak1, Robert S. Fischer1, Rebecca K. Zoltoski2, Jerome R. Kuszak3, and Velia M. Fowler1
1 Department of Cell Biology, The Scripps Research Institute, La Jolla, CA 92037
2 Department of Biological Health Sciences, Illinois College of Optometry, Chicago, IL 60616
3 Departments of Pathology and Ophthalmology, Rush University Medical Center, Chicago, IL 60612
Hexagonal packing geometry is a hallmark of close-packed epithelial cells in metazoans. Here, we used fiber cells of the vertebrate eye lens as a model system to determine how the membrane skeleton controls hexagonal packing of post-mitotic cells. The membrane skeleton consists of spectrin tetramers linked to actin filaments (F-actin), which are capped by tropomodulin1 (Tmod1) and stabilized by tropomyosin (TM). In mouse lenses lacking Tmod1, initial fiber cell morphogenesis is normal, but fiber cell hexagonal shapes and packing geometry are not maintained as fiber cells mature. Absence of Tmod1 leads to decreased TM levels, loss of F-actin from membranes, and disrupted distribution of β2-spectrin along fiber cell membranes. Regular interlocking membrane protrusions on fiber cells are replaced by irregularly spaced and misshapen protrusions. We conclude that Tmod1 and TM regulation of F-actin stability on fiber cell membranes is critical for the long-range connectivity of the spectrin–actin network, which functions to maintain regular fiber cell hexagonal morphology and packing geometry.
