of them maturate into solids (gel-like or even crystalline, depending on the extent of spatial ordering within the condensate). Many proteins participating in the formation of MLO contain so-called intrinsically disordered regions (IDRs), parts of the polypeptide chain that can adopt multiple secondary structures and form random coils in solution. IDRs can provide interactions responsible for LLPS, but over time conformational changes (sometimes promoted by mutations or post-translational modifications) may lead to the formation of higher ordered structures and solidification of MLOs. Some MLOs serve their biological role as solid particles (e.g. Balbiani body stabilised by β-sheet structure), but in many cases transformation from liquid to solid results in the formation of pathological aggregates. Examples of both liquid-liquid phase separating and aggregation-prone proteins include FUS, TDP-43 and hnRNPA1. Aggregates of these proteins are associated with neurodegenerative diseases (e.g. amyotrophic lateral sclerosis, or frontotemporal dementia). History At the start of the 20th century, scientists had become interested in the stability of colloids, both the dispersions of solid particles and the solutions of polymeric molecules. It was known that salts and temperature could often be used to cause flocculation of a colloid. The German chemist F.W. Tiebackx reported in 1911 that flocculation could also be induced in certain polymer solutions by mixing them together. In particular, he reported the observation of opalescence (a turbid mixture) when equal volumes of acidified 0.5% “washed” gelatine solution, and 2% gum arabic solution were mixed. Tiebackx did not further analyse the nature of the flocs, but it is likely that this was an example of complex coacervation. Dutch chemist H. G. Bungenberg-de Jong reported in his PhD thesis (Utrecht, 1921) two types of flocculation in agar solutions: one that leads to a suspensoid state, and one that leads to an emulsoid state. He observed the emulsoid state under the microscope and described small particles that merged into larger particles (Thesis, p. 82), most likely a description of coalescing coacervate droplets. Several years later, in 1929, Bungenberg-de Jong published a seminal paper with his PhD advisor, H. R. Kruyt, entitled “Coacervation. Partial miscibility in colloid systems”. In their paper, they give many more examples of colloid systems that flocculate into an emulsoid state, either by varying the temperature, by adding salts, co-solvents or by mixing together two oppositely charged polymer colloids, and illustrate their observations with the first microscope pictures of coacervate droplets. They term this phenomenon