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Analysis of the simulations was carried out in the GROMACS suite of programs. VMD was used for molecular graphics [36] . The first 2 µs of the CG simulations were discarded as equilibration periods for calculation of ensemble-average properties.

The structural properties of the bilayer, the density distribution of TO and POPC, and the flip flop rates of TO from the two copies each of the UNI5 and RAND5 were qualitatively and quantitatively similar, showing that the results discussed in the following sections are independent of the initial conformations of TO in the POPC bilayer. Furthermore, results from the larger 4X5 and 4XMID5 simulations were identical, indicating that the final conformations were independent of the initial state of the system. For clarity, results from only one out of the four CG simulations with 5.2% TO will be presented, unless otherwise indicated. Similarly, the results from only one of the two copies of UNI2 are presented.

The density distribution of the glycerol backbone of TO from the UNI2 (2.3% TO), UNI5 (5.2% TO) and 4X5 (5.2% TO, larger simulation size) simulations is shown in Fig. 1 . Simulations snapshots from the three simulations are shown in Fig. 2 . The density of TO components has been multiplied by a factor of 10 for clarity. Counter-intuitively, there is a significant density of the polar TO glycerol backbone residing in the hydrophobic center of the membrane in all three cases. There is a higher density of TO in the UNI5 simulation, compared to the UNI2 simulation. In the UNI5 simulation, some TO molecules partition into the bilayer center in the form of a TO aggregate, which is devoid of any specific arrangement of TO molecules. The number of TO molecules which remained at the interface were similar for the 2.3% and 5.2% simulations, indicating the limited interfacial solubility of TO in lipid bilayers. The excess lipids that could not be accommodated at the interface in the 5.2% simulations formed an aggregate in the middle of the membrane. However, as the system size increased, very little TO was left at the interface ( Fig. 1a ) in the 4XMID5 simulation. Thus, TO preferred to partition to the aggregate, rather than stay at the membrane interface in a sufficiently large system with a larger aggregate. There was very little TO at the interface in the 4X5, 4XMID5, 9X5 and 16X5 simulations, and the aggregate occupied an area almost half of that of the membrane patch ( Fig. 2a ). No aggregate was formed in the UNI2 simulations. We note that the non-zero density of TO at the bilayer center in the UNI2 system demonstrates that TO can partition into the bilayer center without an aggregate being present. The density profile of UNI2 is similar to that of the larger 4X2 system of the same concentration. The aggregate at the center of the membrane is biologically relevant. It is similar to triglyceride-rich mobile lipid domains previously detected using 1 H-NMR spectra of cell suspensions that detect mobile lipid domains that carry a high amount of triglycerides [11] , [26] , [28] . The aggregate is also similar to a nascent lipid droplet observed in the ER membrane. We will return to this in the Discussion .

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