High-resolution stem diameter variations (SDV) are widely recognized as a useful drought stress indicator and have therefore been used in many irrigation scheduling studies. More recently, SDV have been used in combination with other plant measurements and biophysical modelling to study fundamental mechanisms underlying whole-plant functioning and growth. The present review aims to scrutinize the important insights emerging from these more recent SDV applications to identify trends in ongoing fundamental research. The main mechanism underlying SDV is variation in water content in stem tissues, originating from reversible shrinkage and swelling of dead and living tissues, and irreversible growth. The contribution of different stem tissues to the overall SDV signal is currently under debate and shows variation with species and plant age, but can be investigated by combining SDV with state-of-the-art technology like magnetic resonance imaging. Various physiological mechanisms, such as water and carbon transport, and mechanical properties influence the SDV pattern, making it an extensive source of information on dynamic plant behaviour. To unravel these dynamics and to extract information on plant physiology or plant biophysics from SDV, mechanistic modelling has proved to be valuable. Biophysical models integrate different mechanisms underlying SDV, and help us to explain the resulting SDV signal. Using an elementary modelling approach, we demonstrate the application of SDV as a tool to examine plant water relations, plant hydraulics, plant carbon relations, plant nutrition, freezing effects, plant phenology and dendroclimatology. In the ever-expanding SDV knowledge base we identified two principal research tracks. First, in detailed short-term experiments, SDV measurements are combined with other plant measurements and modelling to discover patterns in phloem turgor, phloem osmotic concentrations, root pressure and plant endogenous control. Second, long-term SDV time series covering many different species, regions and climates provide an expanding amount of phenotypic data of growth, phenology and survival in relation to microclimate, soil water availability, species or genotype, which can be coupled with genetic information to support ecological and breeding research under on-going global change. This under-exploited source of information has now encouraged research groups to set up coordinated initiatives to explore this data pool via global analysis techniques and data-mining.
Mangrove forests depend on a dense structure of sufficiently large trees to fulfil their essential functions as providers of food and wood for animals and people, CO2 sinks and protection from storms. Growth of these forests is known to be dependent on the salinity of soil water, but the influence of foliar uptake of rainwater as a freshwater source, additional to soil water, has hardly been investigated. Under field conditions in Australia, stem diameter variation, sap flow and stem water potential of the grey mangrove (Avicennia marina (Forssk.) Vierh.) were simultaneously measured during alternating dry and rainy periods. We found that sap flow in A. marina was reversed, from canopy to roots, during and shortly after rainfall events. Simultaneously, stem diameters rapidly increased with growth rates up to 70 μm h-1, which is about 25-75 times the normal growth rate reported in temperate trees. A mechanistic tree model was applied to provide evidence that A. marina trees take up water through their leaves, and that this water contributes to turgor-driven stem growth. Our results indicate that direct uptake of freshwater by the canopy during rainfall supports mangrove tree growth and serve as a call to consider this water uptake pathway if we aspire to correctly assess influences of changing rainfall patterns on mangrove tree growth.
See also the Commentary by Cernusak and Cheesman
Sap-flow measurements have become increasingly important in plant science. Since the early experiments with dyes, many methods have been developed. Most of these are based on the application of heat in the sapwood which is transported by the moving sap. By measuring changes in the temperature field around the heater, sap flow can be derived. Although these methods all have the same basis, their working principles vary widely. A first distinction can be made between those measuring the sap-flow rate (g h–1) such as the stem heat balance and trunk sector heat balance method and those measuring sap-flux density (cm3 cm–2 h–1). Within the latter, the thermal dissipation and heat field deformation methods are based on continuous heating, whereas the compensation heat pulse velocity, Tmax,heat ratio, calibrated average gradient and Sapflow+ methods are based on the application of heat pulses. Each of these methods has its advantages and limitations. Although the sap-flow rate methods have been adequately described in previous reviews, recent developments in sap-flux density methods prompted a synthesis of the existing but scattered literature. This paper reviews sap-flux density methods to enable users to make a well founded choice, whether for practical applications or fundamental research questions, and to encourage further improvement in sap-flux density measurement techniques
Summary• To our knowledge, to date, no nonempirical method exists to measure reverse, low or high sap flux density. Moreover, existing sap flow methods require destructive wood core measurements to determine sapwood water content, necessary to convert heat velocity to sap flux density, not only damaging the tree, but also neglecting seasonal variability in sapwood water content.• Here, we present a nonempirical heat-pulse-based method and coupled sensor which measure temperature changes around a linear heater in both axial and tangential directions after application of a heat pulse. By fitting the correct heat conduction-convection equation to the measured temperature profiles, the heat velocity and water content of the sapwood can be determined.• An identifiability analysis and validation tests on artificial and real stem segments of European beech (Fagus sylvatica L.) confirm the applicability of the method, leading to accurate determinations of heat velocity, water content and hence sap flux density.• The proposed method enables sap flux density measurements to be made across the entire natural occurring sap flux density range of woody plants. Moreover, the water content during low flows can be determined accurately, enabling a correct conversion from heat velocity to sap flux density without destructive core measurements.
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