We present a generic theory for the dynamics of a stiff filament under tension, in an active medium with orientational correlations, such as a microtubule in contractile actin. In sharp contrast to the case of a passive medium, we find the filament can stiffen, and possibly oscillate or buckle, depending on both the contractile or tensile nature of the activity and the filament-medium anchoring interaction. We also demonstrate a strong violation of the fluctuation-dissipation (FD) relation in the effective dynamics of the filament, including a negative FD ratio. Our approach is also of relevance to the dynamics of axons, and our model equations bear a remarkable formal similarity to those in recent work [Martin P, Hudspeth AJ, Juelicher F (2001) Proc Natl Acad Sci USA 98:14380-14385] on auditory hair cells. Detailed tests of our predictions can be made by using a single filament in actomyosin extracts or bacterial suspensions.cytoskeleton | active hydrodynamics | microrheology | fluctuation-dissipation ratio | neuronal growth cone T he cytoskeleton (1) is a dense multicomponent meshwork of semiflexible polymers which interact sterically as well as through active (1-3) processes. Although the blending of polymers industrially requires special effort, the active environment of the living cell provides a setting in which polymers that differ substantially in their stiffness are naturally mixed and interact. Moreover, active processes, such as polymerization and the working of molecular motors, lead to the generation of stresses without the external imposition of flow fields. These two mechanisms combine to yield a rich range of novel physical phenomena. The role of activity in cytoskeletal mechanics is receiving increasing attention, as seen from many recent theoretical and experimental studies of the rheology of cells and cell extracts (4-6). It is clear in particular (7) that interactions between different species of filaments are crucial for cell motility, cell division, vesicular transport, and organelle positioning and integrity.In this paper, we make a study of the effect of these interactions by modeling the dynamics of a stiff filament, which we will call a "microtubule", immersed in an active medium ( Fig. 1) with orientational degrees of freedom, which we will call "F-actin". We emphasize here that the names microtubule and F-actin are introduced for convenience: We consider both contractile and tensile activity, although only the former applies to actomyosin. Our treatment applies more generally to semiflexible polymers under tension in a wide variety of active media. We describe the medium by the active generalization of liquid-crystal hydrodynamics (2,3,(8)(9)(10)(11)(12)(13)(14). For the purposes of this paper, an active medium is one whose constituent particles possess the ability to extract energy from an ambient nutrient bath and dissipate it, executing some kind of systematic motion in the process. This activity endows each such particle with a permanent uniaxial stress. The other central ingredie...