ALBERT

Description: While natural spatial temperature gradients between measurement needles have been thoroughly investigated for continuous heat-based sap flow methods, little attention has been given to how natural changes in stem temperature impact heat pulse-based methods through temporal rather than spatial effects. By modelling the theoretical equation for both an ideal instantaneous pulse and a step pulse and applying a finite element model which included actual needle dimensions and wound effects, the influence of a varying stem temperature on heat pulse-based methods was investigated. It was shown that the heat ratio (HR) method was influenced, while for the compensation heat pulse and T max methods changes in stem temperatures of up to 0.002 °C s –1 did not lead to significantly different results. For the HR method, rising stem temperatures during measurements led to lower heat pulse velocity values, while decreasing stem temperatures led to both higher and lower heat pulse velocities, and to imaginary results for high flows. These errors of up to 40% can easily be prevented by including a temperature correction in the data analysis procedure, calculating the slope of the natural temperature change based on the measured temperatures before application of the heat pulse. Results of a greenhouse and outdoor experiment on Pinus pinea L. show the influence of this correction on low and average sap flux densities.

Description: 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.

Description: The formation of air emboli in the xylem during drought is one of the key processes leading to plant mortality due to loss in hydraulic conductivity, and strongly fuels the interest in quantifying vulnerability to cavitation. The acoustic emission (AE) technique can be used to measure hydraulic conductivity losses and construct vulnerability curves. For years, it has been believed that all the AE signals are produced by the formation of gas emboli in the xylem sap under tension. More recent experiments, however, demonstrate that gas emboli formation cannot explain all the signals detected during drought, suggesting that different sources of AE exist. This complicates the use of the AE technique to measure emboli formation in plants. We therefore analysed AE waveforms measured on branches of grapevine ( Vitis vinifera L. ‘Chardonnay’) during bench dehydration with broadband sensors, and applied an automated clustering algorithm in order to find natural clusters of AE signals. We used AE features and AE activity patterns during consecutive dehydration phases to identify the different AE sources. Based on the frequency spectrum of the signals, we distinguished three different types of AE signals, of which the frequency cluster with high 100–200 kHz frequency content was strongly correlated with cavitation. Our results indicate that cavitation-related AE signals can be filtered from other AE sources, which presents a promising avenue into quantifying xylem embolism in plants in laboratory and field conditions.

Description: The hydraulic performance of woody species during drought is currently of high interest in the context of climate change. It is known that woody species have the capacity to mitigate water shortage by using internally stored water. Elastic shrinkage of living cells and also water release during cavitation contribute to the so-called ‘hydraulic capacitance’ ( C ) of the plant, which adds water to the transpiration stream and buffers fluctuations in water potential. Although sap-conducting conduits may ultimately serve as a water pool, cavitation will hamper the conduction of sap. Both hydraulic conductivity and C are thus inextricably linked and the interaction between both should be studied to better understand hydraulic functioning of woody species during drought. However, measurements of C are scarce and no distinction is usually made between C from elastic storage and C supplied by cavitation. In this paper, we propose a new method to assess both the decrease in hydraulic conductivity and the change in C during bench dehydration of a whole-branch segment using continuous measurements of acoustic emissions, radial diameter shrinkage and gravimetrical water loss. With this method we could establish proper vulnerability curves for grapevine ( Vitis vinifera L. ‘Johanniter’) and quantify C during dehydration. Our results showed that loss in hydraulic conductivity during the cavitation phase was accompanied by 22–92% gain in hydraulic capacitance; therefore, a certain degree of cavitation may be tolerated in grapevine during periods of drought stress.

Description: Plant drought responses are still not fully understood. Improved knowledge on drought responses is, however, crucial to better predict their impact on individual plant and ecosystem functioning. Mechanistic models in combination with plant measurements are promising for obtaining information on plant water status and can assist us in understanding the effect of limiting soil water availability and drought stress. While existing models are reliable under sufficient soil water availability, they generally fail under dry conditions as not all appropriate mechanisms seem yet to have been implemented. We therefore aimed at identifying mechanisms underlying plant drought responses, and in particular investigated the behaviour of hydraulic resistances encountered in the soil and xylem for grapevine ( Vitis vinifera L.) and oak ( Quercus robur L.). A variable hydraulic soil-to-stem resistance was necessary to describe plant drought responses. In addition, implementation of a variable soil-to-stem hydraulic resistance enabled us to generate an in situ soil-to-stem vulnerability curve, which might be an alternative to the conventionally used vulnerability curves. Furthermore, a daily recalibration of the model revealed a drought-induced increase in radial hydraulic resistance between xylem and elastic living tissues. Accurate information on plant hydraulic resistances and simulation of plant drought responses can foster important discussions regarding the functioning of plants and ecosystems during droughts.

Description: Several heat-based sap flow methods, such as the heat field deformation method and the heat ratio method, include the thermal diffusivity D of the sapwood as a crucial parameter. Despite its importance, little attention has been paid to determine D in a plant physiological context. Therefore, D is mostly set as a constant, calculated during zero flow conditions or from a method of mixtures, taking into account wood density and moisture content. In this latter method, however, the meaning of the moisture content is misinterpreted, making it theoretically incorrect for D calculations in sapwood. A correction to this method, which includes the correct application of the moisture content, is proposed. This correction was tested for European and American beech and Eucalyptus caliginosa Blakely & McKie. Depending on the dry wood density and moisture content, the original approach over- or underestimates D and, hence, sap flux density by 10% and more.

Description: Sap flow measurements conducted with thermal dissipation probes (TDPs) are vulnerable to natural temperature gradient (NTG) bias. Few studies, however, attempted to explain the dynamics underlying the NTG formation and its influence on the sensors' signal. This study focused on understanding how the TDP signals are affected by negative and positive temperature influences from NTG and tested the novel cyclic heat dissipation (CHD) method to filter out the NTG bias. A series of three experiments were performed in which gravity-driven water flow was enforced on freshly cut stem segments of Fagus sylvatica L., while an artificial temperature gradient (ATG) was induced. The first experiment sought to confirm the incidence of the ATG on sensors. The second experiment established the mis-estimations caused by the biasing effect of the ATG on standard TDP measurements. The third experiment tested the accuracy of the CHD method to account for the ATG biasing effect, as compared with other cyclic correction methods. During experiments, sap flow measured by TDP was assessed against gravimetric measurements. The results show that negative and positive ATGs were comparable in pattern but substantially larger than field NTGs. Second, the ATG bias caused an overestimation of the standard TDP sap flux density of ~17 cm 3 cm –2 h –1 by 76%, and the sap flux density of ~2 cm 3 cm –2 h –1 by over 800%. Finally, the proposed CHD method successfully reduced the max. ATG bias to 25% at ~11 cm 3 cm –2 h –1 and to 40% at ~1 cm 3 cm –2 h –1 . We concluded that: (i) the TDP method is susceptible to NTG especially at low flows; (ii) the CHD method successfully corrected the TDP signal and resulted in generally more accurate sap flux density estimates (mean absolute percentage error ranging between 11 and 21%) than standard constant power TDP method and other cyclic power methods; and (iii) the ATG enforcing system is a suitable way of re-creating NTG for future tests.

Description: Transport phloem is no longer associated with impermeable pipes, but is instead considered as a leaky system in which loss and retrieval mechanisms occur. Local stem chilling is often used to study these phenomena. In this study, 5-cm- lengths of stems of 3-year-old oak trees ( Quercus robur L.) were locally chilled for 1 week to investigate whether observations at stem and leaf level can be explained by the leakage-retrieval mechanism. The chilling experiment was repeated three times across the growing season. Measurements were made of leaf photosynthesis, carbohydrate concentrations in leaves and bark, stem growth and maximum daily stem shrinkage. Across the growing season, a feedback inhibition in leaf photosynthesis was observed, causing increased dark respiration and starch concentration. This inhibition was attributed to the total phloem resistance which locally increased due to the cold temperatures. It is hypothesized that this higher phloem resistance increased the phloem pressure above the cold block up to the source leaves, inducing feedback inhibition. In addition, an increase in radial stem growth and carbohydrate concentration was observed above the cold block, while the opposite occurred below the block. These observations indicate that net lateral leakage of carbohydrates from the phloem was enhanced above the cold block and that translocation towards regions below the block decreased. This behaviour is probably also attributable to the higher phloem resistance. The chilling effects on radial stem growth and carbohydrate concentration were significant in the middle of the growing season, while they were not at the beginning and near the end of the growing season. Furthermore, maximum daily shrinkages were larger above the cold block during all chilling experiments, indicating an increased resistance in the xylem vessels, also generated by low temperatures. In conclusion, localized stem chilling altered multiple carbon processes in the source leaves and the main stem by changing hydraulic resistances.