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1.  Development and persistence of sandsheaths of Lyginia barbata (Restionaceae): relation to root structural development and longevity 
Annals of Botany  2011;108(7):1307-1322.
Background and Aims
Strongly coherent sandsheaths that envelop perennial roots of many monocotyledonous species of arid environments have been described for over a century. This study, for the first time, details the roles played by the structural development of the subtending roots in the formation and persistence of the sheaths.
The structural development of root tissues associated with persistent sandsheaths was studied in Lyginia barbata, native to the Western Australian sand plains. Cryo-scanning electron microscopy CSEM, optical microscopy and specific staining methods were applied to fresh, field material. The role of root hairs was clarified by monitoring sheath development in roots separated from the sand profile by fine mesh.
Key Results and Conclusions
The formation of the sheaths depends entirely on the numerous living root hairs which extend into the sand and track closely around individual grains enmeshing, by approx. 12 cm from the root tip, a volume of sand more than 14 times that of the subtending root. The longevity of the perennial sheaths depends on the subsequent development of the root hairs and of the epidermis and cortex. Before dying, the root hairs develop cellulosic walls approx. 3 µm thick, incrusted with ferulic acid and lignin, which persist for the life of the sheath. The dead hairs remain in place fused to a persistent platform of sclerified epidermis and outer cortex. The mature cortex comprises this platform, a wide, sclerified inner rim and a lysigenous central region – all dead tissue. We propose that the sandsheath/root hair/epidermis/cortex complex is a structural unit facilitating water and nutrient uptake while the tissues are alive, recycling scarce phosphorus during senescence, and forming, when dead, a persistent essential structure for maintenance of a functional stele in the perennial Lyginia roots.
PMCID: PMC3197463  PMID: 21969258
CSEM; lignified and suberized root hairs; Lyginia barbata; perennial drought-tolerant roots; persistent root hairs; phosphorus recycling; Restionaceae; rhizosheaths; root hair histochemistry; sandbinding roots
2.  Neoformation of clay in lateral root catchments of mallee eucalypts: a chemical perspective 
Annals of Botany  2009;105(1):23-36.
Background and Aims
A previous paper (Annals of Botany 103: 673–685) described formation of clayey pavements in lateral root catchments of eucalypts colonizing a recently formed sand dune in south-west Western Australia. Here chemical and morphological aspects of their formation at the site are studied.
Chemical and physical examinations of soil cores through pavements and sand under adjacent heath assessed build-up of salts, clay and pH changes in or below pavements. Relationships of root morphology to clay deposition were examined and deposits subjected to scanning electron microscopy and energy-dispersive X-ray analysis. Xylem transport of mineral elements in eucalypt and non-eucalypt species was studied by analysis of xylem (tracheal) sap from lateral roots.
Key Results
The columns of which pavements are composed develop exclusively on lower-tier lateral roots. Such sites show intimate associations of fine roots, fungal filaments, microbiota and clay deposits rich in Si, Al and Fe. Time scales for construction of pavements by eucalypts were assessed. Cores through columns of pavemented profiles showed gross elevations of bulk density, Al, Fe and Si in columns and related increases in pH, Mg and Ca status in lower profiles. A cutting through the dune exhibited pronounced alkalinity (pH 7–10) under mallee woodland versus acidity (pH 5–6·5) under proteaceous heath. Xylem sap analyses showed unusually high concentrations of Al, Fe, Mg and Si in dry-season samples from column-bearing roots.
Deposition of Al–Fe–Si-rich clay is pivotal to pavement construction by eucalypts and leads to profound chemical and physical changes in relevant soil profiles. Microbial associates of roots are likely to be involved in clay genesis, with parent eucalypts supplying the required key mineral elements and carbon sources. Acquisition of the Al and Fe incorporated into clay derives principally from hydraulic uplift from ground water via deeply penetrating tap roots.
PMCID: PMC2794064  PMID: 19897459
Niche construction; eucalypts; root morphology; xylem transport; hydraulic lift; element mining; soil formation; biomineralization; soil pans; duplex soils
3.  Contemporary biogenic formation of clay pavements by eucalypts: further support for the phytotarium concept 
Annals of Botany  2009;103(5):673-685.
Background and Aims
Clayey (sodosolic) pavements were studied in lateral root catchments of eucalypts where mixed myrtaceous:proteaceous vegetation was colonizing a dune of quartzitic sand blown out from a playa lake during late Pleistocene times. The site at Chillinup in south-west Western Australia provided an opportunity to examine these signature pavements in an unequivocally recent setting, and to assess their effects on competing non-eucalypt vegetation.
Pavements were located, and their extents and depths assessed by probing with steel rods, followed by corings and pit excavations using an air spade. Listings of plant species, growth forms and root morphologies were assembled for different vegetation zones in a representative transect across the dune. A deep cutting through the dune provided details on pavement morphology and modifications to the sand deposit bioengineered by eucalypt and heath vegetation.
Key Results
Clay pavements comprised closely spaced, round-topped columns whose mean diameters and depths varied between eucalypt species. Incipient pavement formation was characterized by clumps of clay deposited around fine root material. Pavements appeared to have been synthesized in situ from locally accessed and imported constituents. Understoreys on superficial pavements of a tree eucalypt were considerably less dense and biodiverse than on the deeper pavements of two mallee species, whilst most profuse vegetation cover was encountered in heath on unmodified (non-pavemented) sand. Certain species were restricted to superficially located pavements, whilst other ‘generalist’ species occurred widely across the dune. Relict pavements formed by earlier generations of eucalypts were present in certain areas of the transect and in soil profiles of the cutting. Some relict pavements colonized by proteaceous shrubs were overprinted with ferricrete.
Clay pavements formed by eucalypts have pronounced effects on understorey vegetation and may have been instrumental in establishment of the complex mosaics of mallee-woodland and proteaceous heathland observed across semi-arid landscapes of south-west Western Australia. Findings are related to earlier observations on the range of plant-mediated changes in soil profiles discussed in the recently advanced ‘Phytotarium’ concept.
PMCID: PMC2707863  PMID: 19141601
Biomineralization; eucalypt roots; niche construction; soil profiles; woody plants; semi-arid ecosystems; vegetation mosaics; competing phytotaria
4.  Quantifying Above‐ and Below‐ground Growth Responses of the Western Australian Oil Mallee, Eucalyptus kochii subsp. plenissima, to Contrasting Decapitation Regimes 
Annals of Botany  2002;90(2):185-197.
Resprouting in the oil mallee, Eucalyptus kochii Maiden & Blakely subsp. plenissima Gardner (Brooker), involves generation of new shoots from preformed meristematic foci on the lignotuber. Numbers of such foci escalated from 200 per lignotuber in trees aged 1 year to 3000 on 4‐ to 5‐year‐old trees. Removal of shoot biomass by decapitation 5 cm above ground in summer (February) or spring (October) resulted in initiation of 140–170 new shoots, but approx. 400 shoots were induced to form if crops of new shoots were successively removed until sprouting ceased and rootstocks senesced. Initially, the new shoot biomass of regenerating coppices increased slowly and the root biomass failed to increase appreciably until 1·7–2·5 years after cutting. Newly cut trees showed loss of fine root biomass, and structural roots failed to secondarily thicken to the extent shown by uncut trees. After 2 years, the biomass of shoots of coppiced plants was only one‐third that of uncut control trees and shoot : root dry mass ratios of coppiced plants were still low (1·5–2·0) compared with those of the controls (average ratio of 3·1). Spring cutting promoted quicker and greater biomass recovery than summer cutting. Starch in below‐ground biomass fell quickly following decapitation and remained low for a 12–18 month period. Utilization of starch reserves in naturally regenerating coppices was estimated to provide only a small proportion of the dry matter accumulated in new shoots. Results are discussed in relation to their impact on coppicing ability of the species under natural conditions or when successively coppiced for shoot biomass production.
PMCID: PMC4240411  PMID: 12197516
Coppice; short‐rotation forestry; shoot growth; root growth; starch utilization; shading; rootstock decline; oil mallee; Eucalyptus kochii Maiden & Blakely subsp. plenissima Gardner (Brooker)

Results 1-4 (4)