title: Growth dynamics of mechanically impeded lupin roots: does altered morphology induce hypoxia?
abstract: OBJECTIVE
  Root axes elongate slowly and swell radially under mechanical impedance. However, temporal and spatial changes to impeded root apices have only been described qualitatively. This paper aims (a) to quantify morphological changes to root apices and (b) assess whether these changes pre-dispose young root tissues to hypoxia.
  
  
  METHODS
  Lupin (Lupinus angustifolius) seedlings were grown into coarse sand that was pressurized through a diaphragm to generate mechanical impedance on growing root axes. In situ observations yielded growth rates and root response to hypoxia. Roots were then removed to assess morphology, cell lengths and local growth velocities. Oxygen uptake into excised segments was measured.
  
  
  RESULTS
  An applied pressure of 15 kPa slowed root extension by 75% after 10-20 h while the same axes thickened by about 50%. The most terminal 2-3 mm of axes did not respond morphologically to impedance, in spite of the slower flux of cells out of this region. The basal boundary of root extension encroached to within 4 mm of the apex (cf. 10 mm in unimpeded roots), while radial swelling extended 10 mm behind the apex in impeded roots. Oxygen demand by segments of these short, thick, impeded roots was significantly different from segments of unimpeded roots when the zones of elongation in each treatment were compared. Specifically, impeded roots consumed O2 faster and O2 consumption was more likely to be O2-limited over a substantial proportion of the elongation zone, making these roots more susceptible to O2 deficit. Impeded roots used more O2 per unit growth (measured as either unit of elongation or unit of volumetric expansion) than unimpeded roots. Extension of impeded roots in situ was O2-limited at sub-atmospheric O2 levels (21% O2), while unimpeded roots were only limited below 11% O2.
  
  
  CONCLUSIONS
  The shift in the zone of extension towards the apex in impeded roots coincided with greater vulnerability to hypoxia even after soil was removed. Roots still encased in impeded soil are likely to suffer from marked O2 deficits.
authors.affiliation: Department of Agriculture, 3 Baron-Hay Court, South Perth, WA 6151, and Department of Biological Sciences, Macquarie University, Sydney, NSW, Australia.
authors.forename: Colin D
authors.initials: CD
authors.lastname: Hanbury
authors.forename: Brian J
authors.initials: BJ
authors.lastname: Atwell
mesh_terms: D015687:Cell Hypoxia; D031297:Lupinus / Q000187:drug effects / Q000254:growth & development* / Q000378:metabolism*; D010100:Oxygen / Q000378:metabolism / Q000494:pharmacology; D010101:Oxygen Consumption / Q000187:drug effects; D018517:Plant Roots / Q000166:cytology / Q000187:drug effects / Q000254:growth & development* / Q000378:metabolism*; D011312:Pressure
chemical_list: D010100:Oxygen