The ability of many living systems to actively self-propel underlies critical biomedical, environmental, and industrial processes. While such active transport is well-studied in uniform settings, environmental complexities such as geometric constraints, mechanical cues, and external stimuli such as chemical gradients and fluid flow can strongly influence transport. In this chapter, we describe recent progress in the study of active transport in such complex environments, focusing on two prominent biological systems-bacteria and eukaryotic cells-as archetypes of active matter. We review research findings highlighting how environmental factors can fundamentally alter cellular motility, hindering or promoting active transport in unexpected ways, and giving rise to fascinating behaviors such as directed migration and large-scale clustering. In parallel, we describe specific open questions and promising avenues for future research. Furthermore, given the diverse forms of active matterranging from enzymes and driven biopolymer assemblies, to microorganisms and synthetic microswimmers, to larger animals and even robots-we also describe connections to other active systems as well as more general theoretical/computational models of transport processes in complex environments. * Preprint of a chapter in the book Out-of-Equilibrium Soft Matter: Active Fluids, to be published by the Royal Society of Chemistry.

The planar cell polarity (PCP) complex orients cytoskeletal and multicellular organization throughout vertebrate development. PCP is speculated to function in formation of the murine lung, where branching morphogenesis generates a complex tree of tubular epithelia whose distal tips expand dramatically during sacculation in preparation for gas exchange after birth. Here, using tissue-specific knockouts, we show that the PCP complex is dispensable in the airway epithelium for sacculation. Rather, we find a novel, Celsr1-independent role for the PCP component Vangl in the pulmonary mesenchyme: loss of Vangl1/2 inhibits mesenchymal thinning and expansion of the saccular epithelium. Further, loss of mesenchymal Wnt5a mimics the sacculation defects observed in Vangl2-mutant lungs, implicating mesenchymal Wnt5a/Vangl signaling as a key regulator of late lung morphogenesis. By mathematically modeling sacculation, we predict that the process of sacculation requires a fluid mesenchymal compartment. Finally, lineage-tracing and cell-shape analyses are consistent with the pulmonary mesenchyme acting as a fluid tissue, and suggest that loss of Vangl1/2 likely impacts the ability of mesenchymal cells to exchange neighbors. Our data thus uncover an explicit function for Vangl and the pulmonary mesenchyme during late lung morphogenesis to actively shape the saccular epithelium.

The ability of many living systems to actively self-propel underlies critical biomedical, environmental, and industrial processes. While such active transport is well-studied in uniform settings, environmental complexities such as geometric constraints, mechanical cues, and external stimuli such as chemical gradients and fluid flow can strongly influence transport. In this chapter, we describe recent progress in the study of active transport in such complex environments, focusing on two prominent biological systems—bacteria and eukaryotic cells—as archetypes of active matter. We review research findings highlighting how environmental factors can fundamentally alter cellular motility, hindering or promoting active transport in unexpected ways, and giving rise to fascinating behaviors such as directed migration and large-scale clustering. In parallel, we describe specific open questions and promising avenues for future research. Furthermore, given the diverse forms of active matter—ranging from enzymes and driven biopolymer assemblies, to microorganisms and synthetic microswimmers, to larger animals and even robots—we also describe connections to other active systems as well as more general theoretical/computational models of transport processes in complex environments.