“…Understanding the dynamics of this expansion, and the processes responsible for them, is fundamental to understanding biofilm development (Lambert et al, 2014;Nadell et al, 2016;Wille and Coenye, 2020). Many studies have focused on understanding the horizontal "range expansion," detailing the impact of physical interactions, the environment, inter-strain competitive dynamics, and more (D'Acunto et al, 2015;Ghosh and Levine, 2017;Wang et al, 2017;Echten et al, 2020;Fei et al, 2020;Xiong et al, 2020;Fortune et al, 2022;Tam et al, 2022;Moore-Ott et al, 2022). Conversely, less is known about the vertical growth of biofilms, despite its importance for determining access to nutrients and oxygen (De Beer et al, 1994;Stewart et al, 2016).…”