Abstract
Eukaryotic cells organize their intracellular components into organelles that can be membrane-bound or membraneless. A large number of membraneless organelles, including nucleoli, Cajal bodies, P-bodies, and stress granules, exist as liquid droplets within the cell and arise from the condensation of cellular material in a process termed liquid-liquid phase separation (LLPS). Beyond a mere organizational tool, concentrating cellular components into membraneless organelles tunes biochemical reactions and improves cellular fitness during stress. In this review, we provide an overview of the molecular underpinnings of the formation and regulation of these membraneless organelles. This molecular understanding explains emergent properties of these membraneless organelles and shines new light on neurodegenerative diseases, which may originate from disturbances in LLPS and membraneless organelles.
Highlights
Eukaryotic cells organize their intracellular components into organelles that can be membrane-bound or membraneless
P granules are collections of RNA and RNA-binding proteins (RBPs) that accumulate on the posterior side of the C. elegans zygote before the cell divides into a posterior and anterior cell [9, 10]
By fluorescently labeling a constitutive P-granule protein, Hyman and co-workers [9] discovered that P granules display liquid-like properties: the granules are spherical, fuse with one another, deform under shear stress, have fast internal rearrangement as assessed by recovery after photobleaching, and drip off the surface of the nucleus like a liquid. These observations led to the conclusion that P granules are liquid droplets inside the cell that form via a process called liquid–liquid phase separation (LLPS) (Fig. 1A)
Summary
Edward Gomes and X James Shorter From the Department of Biochemistry and Biophysics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104 Eukaryotic cells organize their intracellular components into organelles that can be membrane-bound or membraneless. A large number of membraneless organelles, including nucleoli, Cajal bodies, P-bodies, and stress granules, exist as liquid droplets within the cell and arise from the condensation of cellular material in a process termed liquid–liquid phase separation (LLPS). We provide an overview of the molecular underpinnings of the formation and regulation of these membraneless organelles. We highlight advances in our understanding of the molecular language of these membraneless organelles with respect to how they form, what functions they serve, what rules regulate them, and how their dysregulation may contribute to human disease
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