Kathy Steppe

Estimating evapotranspiration and drought stress with ground-based thermal remote sensing in agriculture: a review

As evaporation of water is an energy-demanding process, increasing evapotranspiration rates decrease the surface temperature (Ts) of leaves and plants. Based on this principle, ground-based thermal remote sensing has become one of the most important methods for estimating evapotranspiration and drought stress and for irrigation. This paper reviews its application in agriculture. The review consists of four parts. First, the basics of thermal remote sensing are briefly reviewed. Second, the theoretical relation between Ts and the sensible and latent heat flux is elaborated. A modelling approach was used to evaluate the effect of weather conditions and leaf or vegetation properties on leaf and canopy temperature. Ts increases with increasing air temperature and incoming radiation and with decreasing wind speed and relative humidity. At the leaf level, the leaf angle and leaf dimension have a large influence on Ts; at the vegetation level, Ts is strongly impacted by the roughness length; hence, by canopy height and structure. In the third part, an overview of the different ground-based thermal remote sensing techniques and approaches used to estimate drought stress or evapotranspiration in agriculture is provided. Among other methods, stress time, stress degree day, crop water stress index (CWSI), and stomatal conductance index are discussed. The theoretical models are used to evaluate the performance and sensitivity of the most important methods, corroborating the literature data. In the fourth and final part, a critical view on the future and remaining challenges of ground-based thermal remote sensing is presented.

Development and verification of a water and sugar transport model using measured stem diameter variations.

In trees, water and sugars are transported by xylem and phloem conduits which are hydraulically linked. A simultaneous study of both flows is interesting, since they concurrently influence important processes such as stomatal regulation and growth. A few mathematical models have already been developed to investigate the influence of both hydraulically coupled flows. However, none of these models has so far been tested using real measured field data. In the present study, a comprehensive whole-tree model is developed that enables simulation of the stem diameter variations driven by both the water and sugar transport. Stem diameter variations are calculated as volume changes of both the xylem and the phloem tissue. These volume changes are dependent on: (i) water transport according to the cohesion-tension theory; (ii) sugar transport according to the Münch hypothesis; (iii) loading and unloading of sugars; and (iv) irreversible turgor-driven growth. The model considers three main compartments (crown, stem, and roots) and is verified by comparison with actual measured stem diameter variations and xylem sap flow rates. These measurements were performed on a young oak (Quercus robur L.) tree in controlled conditions and on an adult beech (Fagus sylvatica L.) tree in a natural forest. A good agreement was found between simulated and measured data. Hence, the model seemed to be a realistic representation of the processes observed in reality. Furthermore, the model is able to simulate several physiological variables which are relatively difficult to measure: phloem turgor, phloem osmotic pressure, and Münchs counterflow. Simulation of these variables revealed daily dynamics in their behaviour which were mainly induced by transpiration. Some of these dynamics are experimentally confirmed in the literature, while others are not.

Diel growth dynamics in tree stems: linking anatomy and ecophysiology

Impacts of climate on stem growth in trees are studied in anatomical, ecophysiological, and ecological disciplines, but an integrative framework to assess those impacts remains lacking. In this opinion article, we argue that three research efforts are required to provide that integration. First, we need to identify the missing links in diel patterns in stem diameter and stem growth and relate those patterns to the underlying mechanisms that control water and carbon balance. Second, we should focus on the understudied mechanisms responsible for seasonal impacts on such diel patterns. Third, information on stem anatomy and ecophysiology should be integrated in the same experiments and mechanistic plant growth models to capture both diel and seasonal scales.

Phloem transport: a review of mechanisms and controls

It is generally believed that an osmotically generated pressure gradient drives the phloem mass flow. So far, this widely accepted Münch theory has required remarkably few adaptations, but the debate on alternative and additional hypotheses is still ongoing. Recently, a possible shortcoming of the Münch theory has been pointed out, suggesting that the Münch pressure flow is more suitable for herbs than for trees. Estimation of the phloem resistance indicates that a point might be reached in long sieve tubes where the pressure required to drive the Münch flow cannot be generated. Therefore, the relay hypothesis regained belief as it implies that the sieve tubes are shorter then the plants axial axis. In the source phloem, three different loading strategies exist which probably result from evolutionary advantages. Passive diffusion seems to be the most primitive one, whereas active loading strategies substantially increase the growth potential. Along the transport phloem, a leakage-retrieval mechanism is observed. Appreciable amounts of carbohydrates are lost from the sieve tubes to feed the lateral sinks, while a part of these lost carbohydrates is subsequently reloaded into the sieve tubes. This mechanism is probably involved to buffer short-term irregularities in phloem turgor and gradient. In the long term, the mechanism controls the replenishment and remobilization of lateral stem storage tissues. As phloem of higher plants has multiple functions in plant development, reproduction, signalling, and growth, the fundamental understanding of the mechanisms behind phloem transport should be elucidated to increase our ability to influence plant growth and development.

Responses of tree species to heat waves and extreme heat events

The number and intensity of heat waves has increased, and this trend is likely to continue throughout the 21st century. Often, heat waves are accompanied by drought conditions. It is projected that the global land area experiencing heat waves will double by 2020, and quadruple by 2040. Extreme heat events can impact a wide variety of tree functions. At the leaf level, photosynthesis is reduced, photooxidative stress increases, leaves abscise and the growth rate of remaining leaves decreases. In some species, stomatal conductance increases at high temperatures, which may be a mechanism for leaf cooling. At the whole plant level, heat stress can decrease growth and shift biomass allocation. When drought stress accompanies heat waves, the negative effects of heat stress are exacerbated and can lead to tree mortality. However, some species exhibit remarkable tolerance to thermal stress. Responses include changes that minimize stress on photosynthesis and reductions in dark respiration. Although there have been few studies to date, there is evidence of within-species genetic variation in thermal tolerance, which could be important to exploit in production forestry systems. Understanding the mechanisms of differing tree responses to extreme temperature events may be critically important for understanding how tree species will be affected by climate change.

Transport of root‐respired CO2 via the transpiration stream affects aboveground carbon assimilation and CO2 efflux in trees

Upward transport of CO₂ via the transpiration stream from belowground to aboveground tissues occurs in tree stems. Despite potentially important implications for our understanding of plant physiology, the fate of internally transported CO₂ derived from autotrophic respiratory processes remains unclear. We infused a ¹³CO₂-labeled aqueous solution into the base of 7-yr-old field-grown eastern cottonwood (Populus deltoides) trees to investigate the effect of xylem-transported CO₂ derived from the root system on aboveground carbon assimilation and CO₂ efflux. The ¹³C label was transported internally and detected throughout the tree. Up to 17% of the infused label was assimilated, while the remainder diffused to the atmosphere via stem and branch efflux. The largest amount of assimilated ¹³C was found in branch woody tissues, while only a small quantity was assimilated in the foliage. Petioles were more highly enriched in ¹³C than other leaf tissues. Our results confirm a recycling pathway for respired CO₂ and indicate that internal transport of CO₂ from the root system may confound the interpretation of efflux-based estimates of woody tissue respiration and patterns of carbohydrate allocation.

An experimental system for analysis of the dynamic sap-flow characteristics in young trees: results of a beech tree

This paper describes an experimental system designed for analysis of the dynamic water flow through young trees, complemented with test results of a 2-year-old beech tree (Fagus sylvatica L.). The system allows automatic and simultaneous measurements of a complex set of plant physiological processes at leaf, branch, stem and root levels [transpiration (E), sap flow (F) and diameter fluctuations (Δd)], in combination with the micrometeorological variables that control these processes [soil and air temperature (Ts and Ta), vapour pressure deficit of the air (D) and photosynthetically active radiation (PAR)]. A 2-d experimental period was used to study the whole-tree water transport dynamics of the young beech tree. Good correspondence between E of the leaves and F in the supporting branch was found. An increased time lag between the F measurements along the hydraulic pathway down towards the root system was observed, indicating the non-steady-state nature of the water flow. The daytime Δd of stem and branch revealed the depletion and the replenishment of internal water reserves. The daily amount of water withdrawn from internal storage was 5% of the total daily transpiration. A good linear relationship was found between the rate of change in internal water storage and the rate of change in stem diameter, having no time lag. We conclude that the data obtained with this system will allow experimental assessment of hydraulic properties in young trees and facilitate calibration of models for non-steady-state conditions of water flow in young trees.

Stem respiration and carbon dioxide efflux of young Populus deltoides trees in relation to temperature and xylem carbon dioxide concentration

Oxidative respiration is strongly temperature driven. However, in woody stems, efflux of CO2 to the atmosphere (EA), commonly used to estimate the rate of respiration (RS), and stem temperature (Tst) have often been poorly correlated, which we hypothesized was due to transport of respired CO2 in xylem sap, especially under high rates of sap flow (fs). To test this, we measured EA, Tst, fs and xylem sap CO2 concentrations ([CO2*]) in 3-year-old Populus deltoides trees under different weather conditions (sunny and rainy days) in autumn. We also calculated RS by mass balance as the sum of both outward and internal CO2 fluxes and hypothesized that RS would correlate better with Tst than EA. We found that EA sometimes correlated well with Tst, but not on sunny mornings and afternoons or on rainy days. When the temperature effect on EA was accounted for, a clear positive relationship between EA and xylem [CO2*] was found. [CO2*] varied diurnally and increased substantially at night and during periods of rain. Changes in [CO2*] were related to changes in fs but not Tst. We conclude that changes in both respiration and internal CO2 transport altered EA. The dominant component flux of RS was EA. However, on a 24-h basis, the internal transport flux represented 9–18% and 3–7% of RS on sunny and rainy days, respectively, indicating that the contribution of stem respiration to forest C balance may be larger than previously estimated based on EA measurements. Unexpectedly, the relationship between RS and Tst was sometimes weak in two of the three trees. We conclude that in addition to temperature, other factors such as water deficits or substrate availability exert control on the rate of stem respiration so that simple temperature functions are not sufficient to predict stem respiration.

Sap-flux density measurement methods : working principles and applicability

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 (gh-1) such as the stem heat balance and trunk sector heat balance method and those measuring sap-flux density (cm3cm-2h-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.

Resistance to radial CO2 diffusion contributes to between-tree variation in CO2 efflux of Populus deltoides stems

Rates of CO2 efflux of stems and branches are highly variable among and within trees and across stands. Scaling factors have only partially succeeded in accounting for the observed variations. In this study, the resistance to radial CO2 diffusion was quantified for tree stems of an eastern cottonwood (Populus deltoides Bartr. ex Marsh.) clone by direct manipulation of the CO2 concentration ([CO2]) of xylem sap under controlled conditions. Tree-specific linear relationships between rates of stem CO2 efflux (JO) and xylem [CO2] were found. The resistance to radial CO2 diffusion differed 6-fold among the trees and influenced the balance between the amount of CO2 retained in the xylem v. that which diffused to the atmosphere. Therefore, we hypothesised that variability in the resistance to radial CO2 diffusion might be an overlooked cause for the inconsistencies and large variations in woody tissue CO2 efflux. It was found that transition from light to dark conditions caused a rapid increase in JO and xylem [CO2], both in manipulated trees and in an intact tree with no sap manipulation. This resulted in an increased resistance to radial CO2 diffusion during the dark, at least for trees with smaller daytime resistances. Stem diameter changes measured in the intact tree supported the idea that higher actual respiration rates occurred at night owing to higher metabolism in relation to an improved water status and higher turgor pressure.