Diffusion is the primary mechanism for the movement of lipids and proteins in a biological membrane. It is important in the formation of various macromolecular structures, such as lipid rafts. The commonly accepted theory for diffusion in membranes suggests that the molecules undergo continuous Brownian diffusion at long length scales, with a "rattling-in-the-cage" motion at short length scales, as shown in figure 1. However, this model has recently been challenged by experimental and simulation results. It has been observed that lipids move in loosely bound clusters rather than as individual molecules [1,2], and that there is a flow-like component to long range lipid diffusion [3]. Ballistic and sub-diffusive regimes have been observed in molecular dynamics simulations [4,5]. Diffusion is mainly studied by two experimental methods: fluorescence techniques and incoherent quasi-elastic neutron scattering. The two techniques access distinctly different length scales, resulting in a "blind spot" at mesoscopic distances. We note that the diffusion coefficients measured by these two techniques often differ by as much as orders of magnitude. The mechanism for diffusion, therefore, seems to depend on the length scale at which it is observed. The blind spot in the mesoscopic range will hopefully be closed in the future using high energy resolution lamor precession techniques performed with spin-echo spectrometers. To extend the window of length scales and investigate the motion of lipid molecules at very short distances, we used the unique capabilities of the IN13 thermal backscattering spectrometer. IN13 provides access to an exceptionally large Q range, covering length scales from 1.3 to 31 Å (0.2 Å -1 < Q < 5 Å -1 ). We used IN13 to study lipid diffusion at length scales smaller than a typical lipid-lipid distance in fluid bilayers. The aim of the experiment was to prove the validity of the Brownian diffusion model down to very small length scales. We chose a stacked model membrane system (DMPC) for this study and analysed the quasi-elastic neutron scattering response of the lipid molecules. Membranes were prepared as solid-supported, multi-lamellar membrane stacks on silicon wafers. Protonated lipids were hydrated by heavy water, so that the experiments were sensitive to the incoherent signal of the lipids. To increase the scattering signal, several wafers with thousands of highly oriented membranes were stacked. The membranes were studied in their physiologically relevant fluid state, at high temperature (T=30 °C) and full hydration. The width of the quasi-elastic energy response (full width at half maximum, FWHM) is shown in figure 2. If a particle diffuses via random Brownian motion, the time evolution of its displacement can be written as = 2Dt, and the quasi-elastic energy broadening has a Lorentzian shape, which demonstrate
