title: Fibre cables in the lacunae of Typha leaves contribute to a tensegrity structure.
abstract: OBJECTIVE
  Cables composed of long, non-lignified fibre cells enclosed in a cover of much shorter thin-walled, crystal-containing cells traverse the air chambers (lacunae) in leaves of the taller species of Typha. The non-lignified fibre cables are anchored in diaphragms composed of stellate cells of aerenchyma tissue that segment the long air chambers into smaller compartments. Although the fibre cables are easily observed and can be pulled free from the porous-to-air diaphragms, their structure and function have been ignored or misinterpreted.
  
  
  METHODS
  Leaves of various species of Typha were dissected and fibre cables were pulled free and observed with a microscope using bright-field and polarizing optics. Maximal tensile strength of freshly removed cables was measured by hanging weights from fibre cables, and Instron analysis was used to produce curves of load versus extension until cables broke.
  
  
  CONCLUSIONS
  Polarized light microscopy revealed that the cellulose microfibrils that make up the walls of the cable fibres are oriented parallel to the long axis of the fibres. This orientation ensures that the fibre cables are mechanically stiff and strong under tension. Accordingly, the measured stiffness and tensile strength of the fibre cables were in the gigapascal range. In combination with the dorsal and ventral leaf surfaces and partitions that contain lignified fibre bundles and vascular strands that are strong in compression, the very fine fibre cables that are strong under tension form a tensegrity structure. The tensegrity structure creates multiple load paths through which stresses are redistributed throughout the 1-3 m tall upright leaves of Typha angustifolia, T. latifolia, T. × glauca, T. domingensis and T. shuttleworthii. The length of the fibre cables relative to the length of the leaf blades is reduced in the last-formed leaves of flowering individuals. Fibre cables are absent in the shorter leaves of Typha minima and, if present, only extend for a few centimetres from the sheath into the leaf blade of Typha laxmannii. The advantage of the structure of the Typha leaf blade, which enables stiffness to give way to flexibility under windy conditions, is discussed for both vegetative and flowering plants.
authors.affiliation: Department of Life Sciences, Ben-Gurion University of the Negev, Beer Sheva, Israel.
authors.forename: Allan
authors.initials: A
authors.lastname: Witztum
authors.forename: Randy
authors.initials: R
authors.lastname: Wayne
mesh_terms: D002473:Cell Wall / Q000648:ultrastructure*; D008855:Microscopy, Electron, Scanning; D018515:Plant Leaves / Q000166:cytology* / Q000648:ultrastructure; D013718:Tensile Strength; D032401:Typhaceae / Q000166:cytology* / Q000648:ultrastructure
keywords: Biomechanics; Typha; cattail; cellulose microfibrils; fibre cables; fibre cells; hierarchical structure; pleochroism; polarized light microscopy; tensegrity