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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.
Methods
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.
doi:10.1093/aob/mcr244
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.  Multiple adaptive responses of Australian native perennial legumes with pasture potential to grow in phosphorus- and moisture-limited environments 
Annals of Botany  2010;105(5):755-767.
Background and Aims
Many Australian legumes have evolved in low-phosphorus (P) soils and low-rainfall areas. Therefore a study was made of the interaction of soil [P] and water availability on growth, photosynthesis, water-use efficiency (WUE) and P nutrition of two Australian native legumes with pasture potential, Cullen australasicum and C. pallidum, and the widely grown exotic pasture legume, lucerne (Medicago sativa).
Methods
Plants were grown in a glasshouse at 3, 10 and 30 mg P kg−1 dry soil for 5 months. At week 10, two drought treatments were imposed, total pot dried (all-dry) and only top soil dried (top-dry), while control pots were maintained at field capacity.
Key Results
Shoot dry weight produced by lucerne was never higher than that of C. australasicum. For C. pallidum only, shoot dry weight was reduced at 30 mg P kg−1 dry soil. The small root system of the Cullen species was quite plastic, allowing plants to access P and moisture efficiently. Lucerne always had a higher proportion of its large root system in the top soil layer compared with Cullen species. All species showed decreased photosynthesis, leaf water potential and stomatal conductance when exposed to drought, but the reductions were less for Cullen species, due to tighter stomatal control, and consequently they achieved a higher WUE. All species showed highest rhizosphere carboxylate concentrations in the all-dry treatment. For lucerne only, carboxylates decreased as P supply increased. Citrate was the main carboxylate in the control and top-dry treatments, and malate in the all-dry treatment.
Conclusions
Multiple adaptive responses of Cullen species and lucerne favoured exploitation of low-P soils under drought. The performance of undomesticated Cullen species, relative to that of lucerne, shows their promise as pasture species for environments such as in south-western Australia where water and P are limiting, especially in view of a predicted drying and warming climate.
doi:10.1093/aob/mcq040
PMCID: PMC2859915  PMID: 20421234
Australian native legumes; carboxylates; climate change; Cullen spp.; drought; Medicago sativa; novel crops; perennial pastures; phosphorus; photosynthesis; root distribution; water-use efficiency
3.  Localized application of soil organic matter shifts distribution of cluster roots of white lupin in the soil profile due to localized release of phosphorus 
Annals of Botany  2010;105(4):585-593.
Background and Aims
Phosphorus (P) is a major factor controlling cluster-root formation. Cluster-root proliferation tends to concentrate in organic matter (OM)-rich surface-soil layers, but the nature of this response of cluster-root formation to OM is not clear. Cluster-root proliferation in response to localized application of OM was characterized in Lupinus albus (white lupin) grown in stratified soil columns to test if the stimulating effect of OM on cluster-root formation was due to (a) P release from breakdown of OM; (b) a decrease in soil density; or (c) effects of micro-organisms other than releasing P from OM.
Methods
Lupin plants were grown in three-layer stratified soil columns where P was applied at 0 or 330 mg P kg−1 to create a P-deficient or P-sufficient background, and OM, phytate mixed with OM, or perlite was applied to the top or middle layers with or without sterilization.
Key Results
Non-sterile OM stimulated cluster-root proliferation and root length, and this effect became greater when phytate was supplied in the presence of OM. Both sterile OM and perlite significantly decreased cluster-root formation in the localized layers. The OM position did not change the proportion of total cluster roots to total roots in dry biomass among no-P treatments, but more cluster roots were concentrated in the OM layers with a decreased proportion in other places.
Conclusions
Localized application of non-sterile OM or phytate plus OM stimulated cluster-root proliferation of L. albus in the localized layers. This effect is predominantly accounted for by P release from breakdown of OM or phytate, but not due to a change in soil density associated with OM. No evidence was found for effects of micro-organisms in OM other than those responsible for P release.
doi:10.1093/aob/mcq012
PMCID: PMC2850796  PMID: 20150198
Citrate exudation; cluster-root formation; root proliferation; Lupinus albus; phosphorus; soil patch; soil micro-organisms; soil organic matter
4.  Effects of phosphorus supply on growth, phosphate concentration and cluster-root formation in three Lupinus species 
Annals of Botany  2009;105(3):365-374.
Background and Aims
In some lupin species, phosphate deficiency induces cluster-root formation, which enhances P uptake by increasing root surface area and, more importantly, the release of root exudates which enhances P availability.
Methods
Three species of Lupinus, L. albus, L. atlanticus and L. micranthus, with inherently different relative growth rates were cultivated under hydroponics in a greenhouse at four phosphate concentrations (1, 10, 50 and 150 µm) to compare the role of internal P in regulating cluster-root formation.
Key Results
The highest growth rate was observed in L. atlanticus, followed by L. albus and L. micranthus. At 1 µm P, cluster-root formation was markedly induced in all three species. The highest P uptake and accumulation was observed in L. micranthus, followed by L. atlanticus and then L. albus. Inhibition of cluster-root formation was severe at 10 µm P in L. atlanticus, but occurred stepwise with increasing P concentration in the root medium in L. albus.
Conclusions
In L. atlanticus and L. albus cluster-root formation was suppressed by P treatments above 10 µm, indicating a P-inducible regulating system for cluster-root formation, as expected. By contrast, production of cluster roots in L. micranthus, in spite of a high internal P concentration, indicated a lower sensitivity to P status, which allowed P-toxicity symptoms to develop.
doi:10.1093/aob/mcp297
PMCID: PMC2826247  PMID: 20037142
Cluster roots; lupin; Lupinus; phosphate nutrition; toxicity; uptake
5.  Change in Uptake, Transport and Accumulation of Ions in Nerium oleander (Rosebay) as Affected by Different Nitrogen Sources and Salinity 
Annals of Botany  2008;102(5):735-746.
Background and Aims
The source of nitrogen plays an important role in salt tolerance of plants. In this study, the effects of NaCl on net uptake, accumulation and transport of ions were investigated in Nerium oleander with ammonium or nitrate as the nitrogen source in order to analyse differences in uptake and cycling of ions within plants.
Methods
Plants were grown in a greenhouse in hydroponics under different salt treatments (control vs. 100 mm NaCl) with ammonium or nitrate as the nitrogen source, and changes in ion concentration in plants, xylem sap exuded from roots and stems, and phloem sap were determined.
Key Results
Plant weight, leaf area and photosynthetic rate showed a higher salt tolerance of nitrate-fed plants compared with that of ammonium-fed plants. The total amount of Na+ transported in the xylem in roots, accumulated in the shoot and retranslocated in the phloem of ammonium-fed plants under salt treatment was 1·8, 1·9 and 2·7 times more, respectively, than that of nitrate-treated plants. However, the amount of Na+ accumulated in roots in nitrate-fed plants was about 1·5 times higher than that in ammonium-fed plants. Similarly, Cl− transport via the xylem to the shoot and its retranslocation via the phloem (Cl− cycling) were far greater with ammonium treatment than with nitrate treatment under conditions of salinity. The uptake and accumulation of K+ in shoots decreased more due to salinity in ammonium-fed plants compared with nitrate-fed plants. In contrast, K+ cycling in shoots increased due to salinity, with higher rates in the ammonium-treated plants.
Conclusions
The faster growth of nitrate-fed plants under conditions of salinity was associated with a lower transport and accumulation of Na+ and Cl− in the shoot, whereas in ammonium-fed plants accumulation and cycling of Na+ and Cl− in shoots probably caused harmful effects and reduced growth of plants.
doi:10.1093/aob/mcn156
PMCID: PMC2712379  PMID: 18772147
Mineral cycling; Nerium oleander; nitrogen source; salinity; xylem and phloem transport
6.  Plant Growth Modelling and Applications: The Increasing Importance of Plant Architecture in Growth Models 
Annals of Botany  2008;101(8):1053-1063.
Background
Modelling plant growth allows us to test hypotheses and carry out virtual experiments concerning plant growth processes that could otherwise take years in field conditions. The visualization of growth simulations allows us to see directly and vividly the outcome of a given model and provides us with an instructive tool useful for agronomists and foresters, as well as for teaching. Functional–structural (FS) plant growth models are nowadays particularly important for integrating biological processes with environmental conditions in 3-D virtual plants, and provide the basis for more advanced research in plant sciences.
Scope
In this viewpoint paper, we ask the following questions. Are we modelling the correct processes that drive plant growth, and is growth driven mostly by sink or source activity? In current models, is the importance of soil resources (nutrients, water, temperature and their interaction with meristematic activity) considered adequately? Do classic models account for architectural adjustment as well as integrating the fundamental principles of development? Whilst answering these questions with the available data in the literature, we put forward the opinion that plant architecture and sink activity must be pushed to the centre of plant growth models. In natural conditions, sinks will more often drive growth than source activity, because sink activity is often controlled by finite soil resources or developmental constraints.
PMA06
This viewpoint paper also serves as an introduction to this Special Issue devoted to plant growth modelling, which includes new research covering areas stretching from cell growth to biomechanics. All papers were presented at the Second International Symposium on Plant Growth Modeling, Simulation, Visualization and Applications (PMA06), held in Beijing, China, from 13–17 November, 2006. Although a large number of papers are devoted to FS models of agricultural and forest crop species, physiological and genetic processes have recently been included and point the way to a new direction in plant modelling research.
doi:10.1093/aob/mcn050
PMCID: PMC2710283  PMID: 18387970
Biomechanics; carbon allocation; functional–structural plant models; meristem; nitrogen; phenotypic plasticity; root architecture; simulation; sink; source; PMA06

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