Hydraulic constraints and stomatal optimization

<p>The classical Cowan-Farquhar approach to identifying optimal stomatal conductance treats total water loss as an imposed constraint. That approach can conflict, both physically and economically, with biophysical constraints on water transport. In this talk, I will illustrate these conflicts and discuss alternative approaches -- recently pioneered by Sperry, Wolf, Eller, and their colleagues -- that aim to penalize excessive transpiration by explicitly incorporating hydraulic risk, using hydraulic vulnerability curves (VCs). In this context, I will present preliminary efforts to determine whether VCs accurately reflect the actual probabilistic risk posed by low water potentials (that is, the expected reduction in total carbon gain), as well as an extension to the recent analytical solution by Eller et al.</p>

Quantifying Key Points of Hydraulic Vulnerability Curves From Drought-Rewatering Experiment Using Differential Method

Precise and accurate estimation of key hydraulic points of plants is conducive to mastering the hydraulic status of plants under drought stress. This is crucial to grasping the hydraulic status before the dieback period to predict and prevent forest mortality. We tested three key points and compared the experimental results to the calculated results by applying two methods. Saplings (n = 180) of Robinia pseudoacacia L. were separated into nine treatments according to the duration of the drought and rewatering. We established the hydraulic vulnerability curve and measured the stem water potential and loss of conductivity to determine the key points. We then compared the differences between the calculated [differential method (DM) and traditional method (TM)] and experimental results to identify the validity of the calculation method. From the drought-rewatering experiment, the calculated results from the DM can be an accurate estimation of the experimental results, whereas the TM overestimated them. Our results defined the hydraulic status of each period of plants. By combining the experimental and calculated results, we divided the hydraulic vulnerability curve into four parts. This will generate more comprehensive and accurate methods for future research.

Incorporating hydraulic traits within the functional strategy spectrum of woody plants globally

<p>Global-change-type droughts, resulting from climate warming and changes in precipitation patterns are believed to be accelerating the rates of tree mortality globally. Simulation of these changes, for instance in global vegetation models, requires parameterisation of plant strategies with respect to drought. However, our understanding of realised combinations of drought-relevant physiological and morphological traits across the range of global forest types is still limited.</p><p>We constructed a dataset consisting of 12 functional traits related to resource acquisition, growth rate, plant defence, water conductivity and hydraulic vulnerability for 11,000 woody species globally. We used a novel envelope-based analysis to assess the functional space occupied by these species whilst circumventing the problem of sparse sampling for many traits. We then subdivided the space into a continuum of strategic “clusters”. These clusters only map partially onto groupings of traditional plant functional types based on leaf type and phenology.</p><p>We found that the functional spaces for the plant strategies are highly interrelated globally, showing that the traits related to resource acquisition are positively associated with growth rates and leaf water conductivity, which together are negatively associated with conservative traits of plant defence, although these associations differ greatly between needleleaf and broadleaf species. However, the trait related to hydraulic failure demonstrated positive associations with resource acquisition, but no relationship with woody defence. Furthermore, there are clear linkages between water flow and hydraulic vulnerability traits, and climatic drivers relating to aridity and plant distribution.</p><p>The clusters identified in this systemic work can form a basis for new plant functional type definitions, facilitating including plant hydraulics in global vegetation models and, taking a step towards making reliable large-scale simulations of drought-driven tree mortality.</p>

A novel approach for fast prediction of hydraulic vulnerability in conifers

<p>Global warming calls for fast and easily applicable methods to measure hydraulic vulnerability in conifers since they are one of the most sensitive plant groups regarding drought stress. Classical methods to determine P<sub>50</sub>, i.e. the water potential resulting in 50% conductivity loss, are labor intensive and prone to errors. In this study, the empirical relationship between percent loss of hydraulic conductivity and relative water loss in sapwood of six conifer species was used to establish a novel proxy for P<sub>50</sub>. Our new proxy P<sub>25W</sub>, defined as 25% of relative water loss induced by air injection, is easy and fast to measure and correlates strongly with P<sub>50</sub> (r = 0.95) as well as with functional wood traits such as the tracheid wall/lumen ratio (r = -0.87). The method is regarded as a strong new phenotyping tool for screening trees for drought sensitivity.</p>

Phenotypic plasticity and genetic adaptation of functional traits influences intra-specific variation in hydraulic efficiency and safety

Understanding which hydraulic traits are under genetic control and/or are phenotypically plastic is essential in understanding how tree species will respond to rapid shifts in climate. We quantified hydraulic traits in Eucalyptus obliqua L'Her. across a precipitation gradient in the field to describe (i) trait variation in relation to long-term climate and (ii) the short-term (seasonal) ability of traits to adjust (i.e., phenotypic plasticity). Seedlings from each field population were raised under controlled conditions to assess (iii) which traits are under strong genetic control. In the field, drier populations had smaller leaves with anatomically thicker xylem vessel walls, a lower leaf hydraulic vulnerability and a lower water potential at turgor loss point, which likely confers higher hydraulic safety. Traits such as the water potential at turgor loss point and ratio of sapwood to leaf area (Huber value) showed significant adjustment from wet to dry conditions in the field, indicating phenotypic plasticity and importantly, the ability to increase hydraulic safety in the short term. In the nursery, seedlings from drier populations had smaller leaves and a lower leaf hydraulic vulnerability, suggesting that key traits associated with hydraulic safety are under strong genetic control. Overall, our study suggests a strong genetic control over traits associated with hydraulic safety, which may compromise the survival of wet-origin populations in drier future climates. However, phenotypic plasticity in physiological and morphological traits may confer sufficient hydraulic safety to facilitate genetic adaptation.

The stability enigma of hydraulic vulnerability curves: addressing the link between hydraulic conductivity and drought-induced embolism

Maintaining xylem water transport under drought is vital for plants, but xylem failure does occur when drought-induced embolisms form and progressively spread through the xylem. The hydraulic method is widely considered the gold standard to quantify drought-induced xylem embolism. The method determines hydraulic conductivity (Kh) in cut branch samples, dehydrated to specific drought levels, by pushing water through them. The technique is widely considered for its reliable Kh measurements, but there is some uncertainty in the literature over how to define stable Kh and how that relates to the degree of xylem embolism formation. Therefore, the most common setup for this method was extended to measure four parameters: (i) inlet Kh, (ii) outlet Kh, (iii) radial flow from xylem to surrounding living tissue and (iv) the pressure difference across the sample. From a strictly theoretical viewpoint, hydraulic steady state, where inflow equals outflow and radial flow is zero, will result in stable Kh. Application of the setup to Malus domestica Borkh. branches showed that achieving hydraulic steady state takes considerable time (up to 300 min) and that time to reach steady state increased with declining xylem water potentials. During each experimental run, Kh and xylem water potentials dynamically increased, which was supported by X-ray computed microtomography visualizations of embolism refilling under both high- (8 kPa) and low-pressure (2 kPa) heads. Supplying pressurized water can hence cause artificial refilling of vessels, which makes it difficult to achieve a truly stable Kh in partially embolized xylem.

Drought-Induced Mortality Is Related to Hydraulic Vulnerability Segmentation of Tree Species in a Savanna Ecosystem

Vulnerability segmentation (VS) has been widely suggested to protect stems and trunks from hydraulic failure during drought events. In many ecosystems, some species have been shown to be non-segmented (NS species). However, it is unclear whether drought-induced mortality is related to VS. To understand this, we surveyed the mortality and recruitment rate and measured the hydraulic traits of leaves and stems as well as the photosynthesis of six tree species over five years (2012–2017) in a savanna ecosystem in Southwest China. Our results showed that the NS species exhibited a higher mortality rate than the co-occurring VS species. Across species, the mortality rate was not correlated with xylem tension at 50% loss of stem hydraulic conductivity (P50stem), but was rather significantly correlated with leaf water potential at 50% loss of leaf hydraulic conductance (P50leaf) and the difference in water potential at 50% loss of hydraulic conductance between the leaves and terminal stems (P50leaf-stem). The NS species had higher Huber values and maximum net photosynthetic rates based on leaf area, which compensated for a higher mortality rate and promoted rapid regeneration under the conditions of dry–wet cycles. To our knowledge, this study is the first to identify the difference in drought-induced mortality between NS species and VS species. Our results emphasize the importance of VS in maintaining hydraulic safety in VS species. Furthermore, the high mortality rate and fast regeneration in NS species may be another hydraulic strategy in regions where severe seasonal droughts are frequent.