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. 2019 Jul 25;10:3321. doi: 10.1038/s41467-019-11316-4

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 3 — Protocell differentiation in opposing gradients. a Graphical representation showing counter-directional SDS (left, red) and POM (right, blue) chemical gradients along the x-axis. b Graphic showing possible sequence of morphogen-mediated transformations in the intersection zone shown in (a) for a single row of PDDA/ATP coacervate droplets (green filled circles, time t = 0). The various spatiotemporal responses produce a row of differentiated protocells with different morphologies (see (c) for details). Relative positions are along the x-axis; t_n = time intervals. c Panel of protocell morphological types produced in the SDS/POM intersection zone displaying representative graphics (row 1) and corresponding optical (row 2) and fluorescence microscopy images (row 3, TNP-ATP green fluorescence; row 4, RITC-PAH red fluorescence). From left to right; native coacervate droplet (C, filled green circle), multi-compartmentalized coacervate vesicle (M_CV, green circle with dots), balloon-shaped POM/coacervate vesicle (P_CB, (light blue graphic), spherical POM/coacervate vesicle (P_CV, dark blue circle), POM/SDS/coacervate vesicle (PS_CV, red circle) and POM/SDS wrinkled vesicle (PS_WV, black graphic). d, e Optical (d) and corresponding fluorescence microscopy (e) images showing a 2D array of differentiated protocells viewed in the observation window after intersection of an opposing reaction-diffusion gradient of SDS (from left to right) and POM (from right to left) at an initial SDS : POM molar ratio of 2.3. Four spatially distinct populations are observed (white dashed lines). Images were recorded after no further changes in morphology were observed (30 min); displayed grid size, 5 × 42. f, g As for d, e, respectively, but for an initial SDS : POM morphogen ratio of 9.0 showing a spatially interpenetrating community. Images were recorded after no further changes in morphology were observed (20 min); displayed grid size, 5 × 42; scale bar 500 µm. h, i Plots showing correlated changes in relative number densities of four different types of differentiated protocells (% population) with spatial position along the direction (x-axis) of opposing morphogen gradients produced at SDS : POM molar ratios of 2.3 (h) and 9.0 (i). j Plot showing a landscape of protocell morphological types produced at different SDS : POM molar ratios under non-diffusive equilibrium conditions. Graphics corresponding to the different forms shown in c except for the SDS/PDDA vesicle (S_V; red circle). All the forms develop irreversibly via a M_CV intermediate except for PS_WV which is derived from PS_CV. Dashed line indicates the presence or absence of a coacervate phase (with/without ATP, respectively). k–n 2D plots showing the spatial distributions for final populations of PS_WV (k), P_CV (l), P_CB (m) and PS_CV (n) morphological types produced in opposing morphogen gradients prepared by injection of different amounts of SDS and POM (% values). Colour scale represents percentage of a given population. Source data are provided as a Source Data file. Error bars represent the standard deviation of the statistics count of the different lines of different protocells (n = 3)