Origin and secretion of milk lipids.

The cream fraction of milk comprises droplets of triacylglycerol coated with cellular membranes. In this review, we discuss how these droplets are formed and secreted from mammary epithelial cells during lactation. This secretory system is especially interesting because the assembled lipid droplets are secreted from the cytoplasm enveloped by cellular membranes. In other cells, such as hepatocytes and enterocytes, lipid is secreted by exocytosis from membrane-bounded compartments of the secretory pathway. Milk lipids originate as small droplets of triacylglycerol, synthesized in or on the surfaces of rough endoplasmic reticulum (ER)4 membranes. These droplets are released into the cytoplasm as microlipid droplets (MLDs) with a surface coat of protein and polar lipid. MLDs may fuse with each other to form larger cytoplasmic lipid droplets (CLDs). Droplets of varying size, are transported to the apical cytoplasm by unknown mechanisms and are secreted from the cell coated with an outer bilayer membrane. CLDs may increase in size in all regions of the cell, especially at the plasma membrane during secretion. Two possible mechanisms for lipid secretion have been proposed: an apical mechanism, in which lipid droplets are enveloped with apical plasma membrane, and a secretory-vesicle mechanism, in which fat droplets are surrounded by secretory vesicles in the cytoplasm and are released from the surface by exocytosis from intracytoplasmic vacuoles. A combination of both mechanisms may be possible. Following secretion, a fraction of the membrane surrounding the globules may be shed from the droplets and give rise to membrane fragments in the skim milk phase.