Cilium

A cellular projection whose structure and functions differ from the bacterial flagellar apparatus. Motile cilia can move fluid; a transport estimate must compare advection with diffusion.

A eukaryotic cilium contains an organized microtubule-based core. Motile forms use dynein-dependent sliding and bending, while many nonmotile primary cilia participate in sensing. These functions distinguish cilia from the rotary filament of a bacterial flagellum. ^[ref]

An axoneme is the organized microtubule-based core of a eukaryotic cilium or flagellum. It supplies a structural framework and, in motile forms, carries motor and linking components that turn relative microtubule sliding into movement. The axoneme is an internal assembly, not the entire membrane-covered projection.^[ref]

The familiar 9+2 arrangement contains nine outer doublets surrounding two central singlets. It describes spatial organization, not eleven identical independent tubules: each outer doublet has two associated tubules. Other axonemal arrangements occur, so the term does not require every cilium to have a central pair or to be motile.

An important mechanical test used isolated sea-urchin sperm axonemes. After brief treatment with trypsin, the preparations initially retained the characteristic organization, but addition of ATP caused active separation into individual tubules and groups. K. E. Summers and I. R. Gibbons interpreted these observations as evidence for sliding between the structural elements.^[ref]

The experiment separates two roles that can otherwise be conflated. Motors such as dynein generate relative motion, while structural constraints normally keep sliding components in an organized assembly. Weakening the constraints reveals sliding as separation; preserving them permits bending. The disintegration experiment does not imply that a normally beating motile cilium repeatedly falls apart and rebuilds itself.

A motile cilium is a eukaryotic cilium that actively bends or rotates to move the cell or the surrounding fluid. Its internal axoneme contains organized microtubules and motor machinery. In familiar bending cilia, dynein uses energy from ATP to drive relative sliding of neighboring microtubule doublets; constraints on that sliding convert motor activity into a change of shape.^[ref]

Many motile cilia have a 9+2 arrangement: nine outer doublets around two central singlets. The nine counts doublets, rather than nine individual outer tubules. Motility should not be defined solely by this pattern, since some motile cilia have other arrangements. Conversely, a ciliary-looking projection is not necessarily motile.

A classic experiment separated the force-producing mechanism from the intact structure's constraints. K. E. Summers and I. R. Gibbons isolated sea-urchin sperm axonemes and briefly treated them with trypsin. Adding ATP then caused active separation into tubules and groups of tubules. The experiment supported sliding between structural elements as a source of motion, rather than requiring each tubule to shorten like a contracting thread.^[ref]

The long beating projection of a eukaryotic sperm cell and short motile cilia share this general mechanical framework. A bacterial flagellum has a different construction and motor. Comparing their functions alone would hide the difference between bending a microtubule assembly and rotating a bacterial filament.