Contemporary phospholipid based cell membranes are formidable barriers to the uptake of polar and charged molecules ranging from metal ions to complex nutrients. Modern cells therefore require sophisticated protein channels and pumps to mediate the exchange of molecules with their environment. The strong barrier function of membranes has made it difficult to understand the origin of cellular life and has been thought to preclude a heterotrophic lifestyle for primitive cells. Although nucleotides can cross DMPC membranes through defects formed at the gel to liquid transition temperature 1, 2 , phospholipid membranes lack the dynamic properties required for membrane growth. Fatty acids and their corresponding alcohols and glycerol monoesters are attractive candidates for the components of protocell membranes because they are simple amphiphiles that form bilayer membrane vesicles 3-5 that retain encapsulated oligonucleotides 3,6 and are capable of growth and division 7-9 . Here we show that such membranes allow the passage of charged molecules such as nucleotides, so that activated nucleotides added to the outside of a model protocell (Fig. 1) spontaneously cross the membrane and take part in efficient template copying in the protocell interior. The permeability properties of prebiotically plausible membranes suggest that primitive protocells could have acquired complex nutrients from their environment in the absence of any macromolecular transport machinery, i.e. could have been obligate heterotrophs.Previous observations of slow permeation of UMP across fatty acid based membranes 6 stimulated us to explore the structural factors that control the permeability of these membranes. We examined membrane compositions with varied surface charge density, fluidity, and stability of regions of high local curvature. We began by studying the permeability of ribose, because this sugar is a key building block of the nucleic acid RNA, and because sugar permeability is conveniently measured with a real-time fluorescence readout of vesicle volume following solute addition 10, 11 . We used pure myristoleic acid (C14:1 fatty acid, myristoleate in its ionized form) as a reference composition, because this compound generates robust vesicles that are more permeable to solutes than the more common longer chain oleic acid. Both myristoleyl alcohol (MA-OH) and the glycerol monoester of myristoleic acid (monomyristolein, GMM) stabilize myristoleate vesicles to the disruptive effects of divalent cations 3,6 . Addition of these amphiphiles should decrease the surface charge density of myristoleate vesicles, while myristoleyl phosphate (MP) should increase the surface charge

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It has been proposed that the initiation of meiotic recombination involves either single-strand or double-strand breaks in DNA. It is difficult to distinguish between these on the basis of genetic evidence because they give rise to similar predictions. All models invoke initiation at specific sites to explain polarity, which is a gradient in gene conversion frequency from one end of a gene to the other. In the accompanying paper we describe the localization of an initiation site for gene conversion to the promoter region of the ARG4 gene of the yeast Saccharomyces cerevisiae. Here, we show that a double-strand break appears at the ARG4 recombination initiation site at the time of recombination, and that the broken DNA molecules end in long single-stranded tails.