Not Just Fat: The Structure and Function of the Lipid Droplet

Lipid droplets (LDs) are independent organelles that are composed of a lipid ester core and a surface phospholipid monolayer. Recent studies have revealed many new proteins, functions, and phenomena associated with LDs. In addition, a number of diseases related to LDs are beginning to be understood at the molecular level. It is now clear that LDs are not an inert store of excess lipids but are dynamically engaged in various cellular functions, some of which are not directly related to lipid metabolism. Compared to conventional membrane organelles, there are still many uncertainties concerning the molecular architecture of LDs and how each function is placed in a structural context. Recent findings and remaining questions are discussed.Lipid droplets (LDs) were recognized as a mere deposit of lipid esters for many years. Recently, LDs have been redefined as authentic organelles that are actively engaged in multiple functions, and these have been reviewed in many recent articles (Murphy 2001; Martin et al. 2006; Ducharme et al. 2008; Fujimoto et al. 2008; Goodman 2008; Olofsson et al. 2008; Thiele et al. 2008; Murphy et al. 2009; Walther et al. 2009; Beller et al. 2010). The name, lipid droplets, implies the opposite: a static and inert nature. Alternative names have been proposed, but lipid droplet is now a generally agreed nomenclature (Martin et al. 2006).LDs are intimately related to many aspects of lipid metabolism. The lipid storage function of LDs is most conspicuous in white adipocytes, which have a gigantic unilocular LD. The regulation of the process of lipid storage and utilization has been the focus of many studies because of the prevalence of obesity, type II diabetes, and metabolic syndrome in the modern world.Besides the canonical lipid-related function, various new functions have been ascribed to LDs more recently, including some that do not appear to be directly related to lipid metabolism. However, both the canonical and noncanonical functions of LDs, are rooted in the unique architecture of LDs. In contrast to other organelles that have aqueous content within a phospholipid bilayer membrane, the basic structure of LDs is thought to be a mass of lipid esters covered by a phospholipid monolayer (Murphy et al. 1999; Tauchi-Sato et al. 2002). In this article, we first review what is known about LD structure and then discuss various functions. We review this information by considering the extent to which the functions can be understood based on the current structural paradigm.