Biological membranes are composed of lipid bilayers that are often asymmetric with regards to the lipid composition and/or aqueous solvent they separate. Studying lipid asymmetry both experimentally and computationally is challenging. Molecular dynamics simulations of lipid bilayers with asymmetry are difficult due to finite system sizes and time scales accessible to simulations. Due to the very slow flip-flop rate for phospholipids, one must first choose how many lipids are on each side of the bilayer, but the resulting bilayer may be unstable (or metastable) due to differing tensile and compressive forces between leaflets. Here we use molecular dynamics simulations to investigate a number of different asymmetric membrane systems, both with atomistic and coarse-grained models. Asymmetries studied include differences in number of lipids, lipid composition (unsaturated and saturated tails and different headgroups), and chemical gradients between the aqueous phases. Extensive analysis of the bilayers' properties such as area per lipid, density, and lateral pressure profiles are used to characterize bilayer asymmetry. We also address how cholesterol (which flip-flops relatively quickly) influences membrane asymmetries. Our results show how each leaflet is influenced by the other and can mitigate the structural changes to the bilayer overall structure. Cholesterol can respond to changes in bilayer asymmetry to alleviate some of the effect on the bilayer structure, but that will alter its leaflet distribution, which in turn affects its chemical potential. Ionic imbalances are shown to have a modest change in bilayer structure, despite large changes in the electrostatic potential. Bilayer asymmetry can also induce a modest electrostatic potential across the membrane. Our results highlight the importance of membrane asymmetry on bilayer properties, the influence of lipid headgroups, tails and cholesterol on asymmetry, and the ability of lipids to adapt to different environments.

The Weak Equivalence Principle states that the geodesics of a test particle in a gravitational field are independent of the particle's constitution. To constrain violations of the Weak Equivalence Principle, we use the one-meter telescope at Table Mountain Observatory near Los Angeles to monitor the relative arrival times of pulses from the Crab Pulsar in the optical (λ ≈ 585 nm) and near-infrared (λ ≈ 814 nm) using an instrument which detects single photons with nanosecond-timing resolution in those two bands. The infrared pulse arrives slightly before the visible pulse. Our three analysis methods give delays with statistical errors of ∆t obs = 7.41±0.58, 0.4±3.6, and 7.35±4.48 microseconds (at most 1/4000 of the pulsar period). We attribute this discrepancy to systematic error from the fact that the visible and infrared pulses have slightly different shapes. Whether this delay emerges from the pulsar, is caused by passing through wavelength-dependent media, or is caused by a violation of the equivalence principle, unless there is a fine-tuned cancellation among these, we set the first upper limit on the differential post-Newtonian parameter at these wavelengths of ∆γ < 1.07 × 10 −10 (3σ). This result falls in an unexplored region of parameter space and complements existing limits on equivalence-principle violation from fast radio bursts, gamma ray bursts, as well as previous limits from the Crab.