, Volume 222, Issue 3, pp 472–483 | Cite as

Cuticular wax deposition in growing barley (Hordeum vulgare) leaves commences in relation to the point of emergence of epidermal cells from the sheaths of older leaves

  • Andrew Richardson
  • Rochus Franke
  • Gerhard Kerstiens
  • Mike Jarvis
  • Lukas Schreiber
  • Wieland Fricke
Original Article


In grasses, leaf cells divide and expand within the sheaths of older leaves, where the micro-environment differs from the open atmosphere. By the time epidermal cells are displaced into the atmosphere, they must have a functional cuticle to minimize uncontrolled water loss. In the present study, gas chromatography and scanning electron microscopy were used to follow cuticular wax deposition along the growing leaf three of barley (Hordeum vulgare L.). 1-Hexacosanol (C26 alcohol) comprised more than 75% of extractable cuticular wax and was used as a marker for wax deposition. There was no detectable wax along the first 20 mm from the point of leaf insertion. Deposition started within the distal portion of the elongation zone (23–45 mm) and continued beyond the point of leaf emergence from the sheath of leaf two. The region where wax deposition commenced shifted towards more proximal (basal) positions when the point of leaf emergence was lowered by stripping back part of the sheath. When relative humidity in the shoot environment was elevated from 70% (standard growth conditions) to 92–96% for up to 4  days prior to analysis, wax deposition did not change significantly. The results show that cuticular waxes are deposited along the growing grass leaf independent of cell age or developmental stage. Instead, the reference point for wax deposition appears to be the point of emergence of cells into the atmosphere. The possibility of changes in relative humidity between enclosed and emerged leaf regions triggering wax deposition is discussed.


Cell development Cuticle Hordeum Leaf growth Relative humidity Wax 



Flame ionization detector


Gas chromatography


Leaf elongation velocity


Mass spectrometry


Point of emergence


Relative elemental growth rate


Relative humidity


Scanning electron microscopy



This research was supported by the Biotechnology and Biological Sciences Research Council (BBSRC), UK, Grant 61/P18283. We would like to thank Eva Simanova and Tobias Wojciechowski from the University of Bonn for their help and advice concerning the GC analysis, Margaret Corrigan and Frank Placido from the University of Paisley for their help and tuition with the SEM and the referees for their helpful comments.


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Copyright information

© Springer-Verlag 2005

Authors and Affiliations

  • Andrew Richardson
    • 1
  • Rochus Franke
    • 2
  • Gerhard Kerstiens
    • 3
  • Mike Jarvis
    • 4
  • Lukas Schreiber
    • 2
  • Wieland Fricke
    • 1
  1. 1.Division of Biological SciencesUniversity of PaisleyPaisleyUK
  2. 2.Department of Ecophysiology, IZMBUniversity of BonnBonnGermany
  3. 3.Lancaster Environment Centre, Department of Biological SciencesLancaster UniversityLancasterUK
  4. 4.Department of ChemistryUniversity of GlasgowGlasgowUK

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