The bacterial flagellum is constructed through a highly ordered process in which gene expression and physical assembly are coordinated.
How does a bacterial cell construct a flagellum in the correct sequence?
A flagellum does not appear in the cell as a completed structure. Its components are assembled in a controlled sequence, generally proceeding from internal structures toward the exterior.
Completion of particular stages can influence expression or export of components needed for later stages.
Flagellar assembly involves regulatory hierarchies, protein export, self-assembly, molecular recognition, checkpoints, and structural transitions. Components destined for external portions of the flagellum must be transported through the developing structure.
The assembly process adds another level to the investigation. Explaining the finished machine requires explaining not only its parts but also how biological systems manufacture and assemble those parts in the proper places and sequence.
Sequential flagellar assembly and its coupling to gene regulation have been demonstrated experimentally. Important checkpoints coordinate completion of one portion of the structure with production or export of later components.
Evolutionary models propose that simpler ancestral secretion or motility systems could have acquired additional components and regulatory relationships progressively.
Although much is known about assembly of modern flagella, the sequence in which the assembly pathway itself evolved is reconstructed rather than directly observed.
More than 60 genes participate in flagellar assembly and function in some extensively studied bacterial systems.
Ordered assembly is relevant to design because functional organization includes not only the final structure but also the information and processes required to build it. At the same time, self-assembly and regulatory evolution are genuine natural processes that must be included in any fair evaluation.
The flagellum illustrates two related problems: the origin of a molecular machine and the origin of the coordinated process that constructs that machine. Modern biology provides extensive knowledge of how existing flagella are assembled. The historical question of how the assembly system itself arose requires reconstruction from comparative, genetic, structural, and experimental evidence.