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'Summary' 14 I. 'Introduction' 14 II. 'The transpiration supply function' 15 III. 'The transpiration loss function' 17 IV. 'General properties of the supply–loss theory' 19 V. 'Variations on the theme' 19 VI. 'Predicting responses to climate' 21 VII. 'Implications for productivity and mortality: the chronic stress hypothesis' 22 VIII. 'Conclusion' 24 'Acknowledgements' 24 References 24 Summary Climate change exposes vegetation to unusual drought, causing declines in productivity and increased mortality. Drought responses are hard to anticipate because canopy transpiration and diffusive conductance (G) respond to drying soil and vapor pressure deficit (D) in complex ways. A growing database of hydraulic traits, combined with a parsimonious theory of tree water transport and its regulation, may improve predictions of at-risk vegetation. The theory uses the physics of flow through soil and xylem to quantify how canopy water supply declines with drought and ceases by hydraulic failure. This transpiration ‘supply function’ is used to predict a water ‘loss function’ by assuming that stomatal regulation exploits transport capacity while avoiding failure. Supply–loss theory incorporates root distribution, hydraulic redistribution, cavitation vulnerability, and cavitation reversal. The theory efficiently defines stomatal responses to D, drying soil, and hydraulic vulnerability. Driving the theory with climate predicts drought-induced loss of plant hydraulic conductance (k), canopy G, carbon assimilation, and productivity. Data lead to the ‘chronic stress hypothesis’ wherein > 60% loss of k increases mortality by multiple mechanisms. Supply–loss theory predicts the climatic conditions that push vegetation over this risk threshold. The theory's simplicity and predictive power encourage testing and application in large-scale modeling.

There is wide breadth of root function within ecosystems that should be considered when modeling the terrestrial biosphere. Root structure and function are closely associated with control of plant water and nutrient uptake from the soil, plant carbon (C) assimilation, partitioning and release to the soils, and control of biogeochemical cycles through interactions within the rhizosphere. Root function is extremely dynamic and dependent on internal plant signals, root traits and morphology, and the physical, chemical and biotic soil environment. While plant roots have significant structural and functional plasticity to changing environmental conditions, their dynamics are noticeably absent from the land component of process-based Earth system models used to simulate global biogeochemical cycling. Their dynamic representation in large-scale models should improve model veracity. Here, we describe current root inclusion in models across scales, ranging from mechanistic processes of single roots to parameterized root processes operating at the landscape scale. With this foundation we discuss how existing and future root functional knowledge, new data compilation efforts, and novel modeling platforms can be leveraged to enhance root functionality in large-scale terrestrial biosphere models by improving parameterization within models, and introducing new components such as dynamic root distribution and root functional traits linked to resource extraction.

Increasing evidence about hydraulic redistribution and its ecological consequences is emerging. Hydraulic redistribution results from an interplay between competing plant and soil water potential gradients. In this work, stem-mediated hydraulic redistribution was studied in a 53-year-old Douglas-fir tree during a period of drought. Sap flux density measurements using the heat field deformation method were performed at four locations: in two large opposing roots and on two sides of the tree stem. Hydraulic redistribution was induced by localized irrigation on one of the measured roots, creating heterogeneous soil water conditions. Stem-mediated hydraulic redistribution was detected during night-time conditions when water was redistributed from the wet side of the tree to the nonirrigated dry side. In addition to stem-mediated hydraulic redistribution, bidirectional flow in the dry root was observed, indicating radial sectoring in the xylem. It was observed that, through stem-mediated hydraulic redistribution, Douglas-fir was unable to increase its transpiration despite the fact that sufficient water was available to one part of the root system. This resulted from the strong water potential gradient created by the dry soil in contact with the nonirrigated part of the root system. A mechanism of stem-mediated hydraulic redistribution is proposed and its possible implications are discussed.

Summary • Here, we tested the alternation of root summer dormancy and winter growth as a critical survival strategy for a long-lived monocotyledon (Restionaceae) adapted to harsh seasonal extremes of Mediterranean southwest Western Australia. • Measurements of growth and the results of comparative studies of the physiology, water content, metabolites, osmotic adjustments, and proteomics of the dormant and growing perennial roots of Lyginia barbata (Restionaceae) were assessed in field-grown plants. • Formation of dormant roots occurred before the onset of summer extremes. They resumed growth (average 2.3 mm d−1) the following winter to eventually reach depths of 2–4 m. Compared with winter-growing roots, summer dormant roots had decreased respiration and protein concentration and c. 70% water content, sustained by sand-sheaths, osmotic adjustment and presumably hydraulic redistribution. Concentrations of compatible solutes (e.g. sucrose and proline) were significantly greater during dormancy, presumably mitigating the effects of heat and drought. Fifteen root proteins showed differential abundance and were correlated with either winter growth or summer dormancy. None matched currently available libraries. • The specific features of the root dormancy strategy of L. barbata revealed in this study are likely to be important to understanding similar behaviour in roots of many long-lived monocotyledons, including overwintering and oversummering crop species.

Summary • While there is strong evidence for hydraulic redistribution (HR) of soil water by trees, it is not known if common mycorrhizal networks (CMN) can facilitate HR from mature trees to seedlings under field conditions. • Ponderosa pine (Pinus ponderosa) seedlings were planted into root-excluding 61-µm mesh barrier chambers buried in an old-growth pine forest. After 2 yr, several mature trees were cut and water enriched in D2O and acid fuchsin dye was applied to the stumps. • Fine roots and mycorrhizal root tips of source trees became heavily dyed, indicating reverse sap flow in root xylem transported water from stems throughout root systems to the root hyphal mantle that interfaces with CMN. Within 3 d, D2O was found in mesh-chamber seedling foliage > 1 m from source trees; after 3 wk, eight of 10 mesh-chamber seedling stem samples were significantly enriched above background levels. Average mesh-chamber enrichment was 1.8× greater than that for two seedlings for which the connections to CMN were broken by trenching before D2O application. • Even small amounts of water provided to mycorrhizas by HR may maintain hyphal viability and facilitate nutrient uptake under drying conditions, which may provide an advantage to seedlings hydraulically linked by CMN to large trees.

Summary • Although hydraulic redistribution of soil water (HR) by roots is a widespread phenomenon, the processes governing spatial and temporal patterns of HR are not well understood. We incorporated soil/plant biophysical properties into a simple model based on Darcy's law to predict seasonal trajectories of HR. • We investigated the spatial and temporal variability of HR across multiple years in two old-growth coniferous forest ecosystems with contrasting species and moisture regimes by measurement of soil water content (θ) and water potential (Ψ) throughout the upper soil profile, root distribution and conductivity, and relevant climate variables. • Large HR variability within sites (0–0.5 mm d−1) was attributed to spatial patterns of roots, soil moisture and depletion. HR accounted for 3–9% of estimated total site water depletion seasonally, peaking at 0.16 mm d−1 (ponderosa pine; Pinus ponderosa) or 0.30 mm d−1 (Douglas-fir; Pseudotsuga menziesii), then declining as modeled pathway conductance dropped with increasing root cavitation. • While HR can vary tremendously within a site, among years and among ecosystems, this variability can be explained by natural variability in Ψ gradients and seasonal courses of root conductivity.