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. 2019 Jan 22;9:292. doi: 10.1038/s41598-018-36646-z

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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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Collagen gel invasion of A549 cells (green) from spheroids with different types of fibroblasts. (a) A549/WI-38 spheroid incubated for 22 h (day 1) (left panel) and 44 h (day 2) (right panel). Inset, the original spheroid at day 0. (b) A549/OUS-11 spheroid incubated for 22 h (day 1). Inset, day 0. (c) Time-lapse images of the box “c” in (b) Arrows trace one cancer cell migrating on four fibroblasts in sequence. See Video 4. (d) Time-lapse images of the box “d” in (b) after 25-h incubation. Arrows trace two cancer cells showing reciprocating movement on a string of fibroblast. (e) A549/LuCAF (left panel) and A549/Ctr-NLF (right panel) spheroids after 22-h incubation (day 1). Note that both LuCAF and Ctr-NLF migrate very actively in the collagen matrix compared with WI-38 (a) and OUS-11 (b) at Day 1. Scale bars, 50 μm, except for 20 μm in (d) In all images, small arrows point cancer cells binding to fibroblasts.