Xylem cell size regulation is a key adaptive response to water deficit in Eucalyptus grandis.

Future climatic scenarios forecast increasingly frequent droughts that will pose substantial consequences on tree mortality. In light of this, drought-tolerant eucalypts have been propagated; however, the severity of these conditions will invoke adaptive responses, impacting the commercially valuable wood properties. To determine what mechanisms govern the wood anatomical adaptive response, highly controlled drought experiments were conducted in Eucalyptus grandis W. Hill ex Maiden, with the tree physiology and transcriptome closely monitored. In response to water deficit, E. grandis displays an isohydric stomatal response to conserve water and enable stem growth to continue, albeit at a reduced rate. Maintaining gaseous exchange is likely a critical short-term response that drives the formation of hydraulically safer xylem. For instance, the development of significantly smaller fibers and vessels was found to increase cellular density, thereby promoting drought tolerance through improved functional redundancy, as well as implosion and cavitation resistance. The transcriptome was explored to identify the molecular mechanisms responsible for controlling xylem cell size during prolonged water deficit. Downregulation of genes associated with cell wall remodeling and the biosynthesis of cellulose, hemicellulose and pectin appeared to coincide with a reduction in cellular enlargement during drought. Furthermore, transcript levels of NAC and MYB transcription factors, vital for cell wall component biosynthesis, were reduced, while those linked to lignification increased. The upregulation of EgCAD and various peroxidases under water deficit did not correlate with an increased lignin composition. However, with the elevated cellular density, a higher lignin content per xylem cross-sectional area was observed, potentially enhancing hydraulic safety. These results support the requirement for higher density, drought-adapted wood as a long-term adaptive response in E. grandis, which is largely influenced by the isohydric stomatal response coupled with cellular expansion-related molecular processes.

An open-source machine-learning approach for obtaining high-quality quantitative wood anatomy data from E. grandis and P. radiata xylem.

Quantitative wood anatomy is a subfield in dendrochronology that requires effective open-source image analysis tools. In this research, the bioimage analysis software QuPath (v0.4.4) is introduced as a candidate for accurately quantifying the cellular properties of the xylem in an automated manner. Additionally, the potential of QuPath to detect the transition of early- to latewood tracheids over the growing season was evaluated to assess a potential application in dendroecological studies. Various algorithms in QuPath were optimized to quantify different xylem cell types in Eucalyptus grandis and the transition of early- to latewood tracheids in Pinus radiata. These algorithms were coded into cell detection scripts for automatic quantification of stem microsections and compared to a manually curated method to assess the accuracy of the cell detections. The automatic cell detection approach, using QuPath, has been validated to be reproducible with an acceptable error when assessing fibers, vessels, early- and latewood tracheids. However, further optimization for parenchyma is still required. This proposed method developed in QuPath provides a scalable and accurate approach for quantifying anatomical features in stem microsections. With minor amendments to the detection and classification algorithms, this strategy is likely to be viable in other plant species.

Wood Formation Modeling - A Research Review and Future Perspectives.

Wood formation has received considerable attention across various research fields as a key process to model. Historical and contemporary models of wood formation from various disciplines have encapsulated hypotheses such as the influence of external (e.g., climatic) or internal (e.g., hormonal) factors on the successive stages of wood cell differentiation. This review covers 17 wood formation models from three different disciplines, the earliest from 1968 and the latest from 2020. The described processes, as well as their external and internal drivers and their level of complexity, are discussed. This work is the first systematic cataloging, characterization, and process-focused review of wood formation models. Remaining open questions concerning wood formation processes are identified, and relate to: (1) the extent of hormonal influence on the final tree ring structure; (2) the mechanism underlying the transition from earlywood to latewood in extratropical regions; and (3) the extent to which carbon plays a role as "active" driver or "passive" substrate for growth. We conclude by arguing that wood formation models remain to be fully exploited, with the potential to contribute to studies concerning individual tree carbon sequestration-storage dynamics and regional to global carbon sequestration dynamics in terrestrial vegetation models.

Stomatal responses of Eucalyptus spp. under drought can be predicted with a gain-risk optimization model.

The frequency and severity of drought events are expected to increase due to climate change, with optimal environmental conditions for forestry likely to shift. Modeling plant responses to a changing climate is therefore vital. We tested the process-based gain-risk model to predict stomatal responses to drought of two Eucalyptus hybrids. The process-based gain-risk model has the advantage that all the parameters used within the model are based on measurable plant traits. The gain-risk model proposes that plants optimize photosynthetic gain while minimizing a hydraulic cost. Previous versions of the model used hydraulic risk as a cost function; however, they did not account for delayed or reduced hydraulic recovery rates from embolism post-drought. Hydraulic recovery has been seen in many species, however it is still unclear how this inclusion of a partial or delayed hydraulic recovery would affect the predictive power of the gain-risk model. Many hydraulic parameters required by the model are also difficult to measure and are not freely available. We therefore tested a simplified gain-risk model that includes a delayed or reduced hydraulic recovery component post-drought. The simplified gain-risk model performed well at predicting stomatal responses in both Eucalyptus grandis × camaldulensis (GC) and Eucalyptus urophylla × grandis (UG). In this study two distinct strategies were seen between GC and UG, with GC being more resistant to embolism formation, however it could not recover hydraulic conductance compared with UG. The inclusion of a delayed or reduced hydraulic recovery component slightly improved model predictions for GC, however not for UG, which can be related to UG being able to recover lost hydraulic conductance and therefore can maintain stomatal conductance regardless of hydraulic risk. Even though the gain-risk model shows promise in predicting plant responses, more information is needed regarding hydraulic recovery after drought.

Xylem as the main origin of stem radius changes in Eucalyptus.

The state-of-the-art interpretation of stem radius changes (DRTotal) for tree water relations is based on knowledge from mostly slow growing tree species. The ratio between diurnal size fluctuations of the rigid xylem (DRXylem) and the respective fluctuations of the elastic bark (DRBark) is known to be small (<0.4) and is of importance for the localisation of water storage dynamics in stems. In this study, fast growing Eucalyptus globulus Labill. in Tasmania were investigated by point dendrometers in order to investigate tree water relations. Unexpectedly, DRXylem was found to be the main driver of DRTotal with the bark acting as a passive layer on top of the fluctuating xylem under most conditions. Accordingly, the ratio between the diurnal fluctuations of the two tissues was found to be much higher (0.6-1.6) than everything reported before. Based on simulations using a hydraulic plant model, the high tissue-specific elasticity of the Eucalyptus xylem was found to explain this atypical response and not osmotically-driven processes or species-specific flow resistances. The wide zone of secondary thickening xylem in various stages of lignification is proposed to be an important component of the high wood elasticity. The tissue acts as additional water storage like the bark and may positively affect the water transport efficiency.

Wood properties in a long-lived conifer reveal strong climate signals where ring-width series do not.

Although tree-ring-width chronologies have been widely used for temperature reconstructions, there are many sites around the world at which there is little evidence of a clear climate signal in the ring-width chronologies. This is the case with the long-lived conifer Huon pine (Lagarostrobos franklinii (Hook. F.) Quinn), endemic to Tasmania, Australia, when the species grows at low elevation. In this study, we developed chronologies of several wood properties (e.g., tracheid radial diameter, microfibril angle) from Huon pine growing at a low-elevation site. We found that despite the absence of a climate signal in the ring-width chronologies, there were significant correlations between wood density, tracheid radial diameter and microfibril angle and temperature, stream flow and a drought index, enabling the development of robust chronologies. This novel finding suggests that chronologies based on these wood properties may have important potential for climate reconstructions from sites and species that have not yet been realized. In particular, a relatively extensive resource of ancient, low-elevation Huon pine in western Tasmania, in which climate signals have not been found using ring widths, may now be useful as part of the broader effort to reconstruct Southern Hemisphere climate.

The development of seasonal tree water deficit in Callitris intratropica.

The tropical conifer Callitris intratropica (Cupressaceae) produces clear annual growth rings, and has been shown to be potentially useful for understanding past climate variability in northern Australia. As climate patterns in this region become less predictable, an understanding of plant responses to different weather patterns is of importance. In this paper, we examine tree water relations using a parameter here called tree water deficit (ΔD), determined from de-trended stem size variability in densely grown ('grove') and isolated trees. This parameter provides an integrated measure of the trees' response to water supply and demand under constantly changing environmental conditions. The work, conducted over 12 months, found that daily variation in tree water deficit was determined mainly by soil water availability, but temperature and relative humidity contributed more to the variability over some periods. Isolated and grove trees exhibited quite distinct patterns of ΔD development during the year, but particularly during the transition between the dry and wet seasons. The results of this work suggest that the dendrochronological interpretation of tree rings in the context of strongly seasonal water availability should incorporate an understanding of the development of seasonal drought in isolated trees compared with trees experiencing strong intra-specific competition. Different responses based on the ecological situations of the trees will affect their patterns of stem growth, and ultimately the climatic information that is incorporated in ring width variability.

Use of a liter-scale microbial desalination cell as a platform to study bioelectrochemical desalination with salt solution or artificial seawater.

Bioelectrochemical desalination is potentially advantageous because of bioenergy production and integrated wastewater treatment and desalination. In this work, the performance and energy benefits of a liter-scale upflow microbial desalination cell (UMDC) were evaluated. The UMDC desalinated both salt solution (NaCl) and artificial seawater, and the removal rate of total dissolved solid (TDS) increased with an increased hydraulic retention time, although TDS reduction in artificial seawater was lower than that in salt solution. Our analysis suggested that electricity generation was a predominant factor in removing TDS (more than 70%), and that other factors, like water osmosis and unknown processes, also contributed to TDS reduction. It was more favorable given the high energy efficiency, when treating salt solution, to operate the UMDC under the condition of high power output compared with that of high current generation because of the amount of energy production; while high current generation was more desired with seawater desalination because of lower salinity in the effluent. Under the condition of the high power output and the assumption of the UMDC as a predesalination in connection with a reversal osmosis (RO) system, the UMDC could produce electrical energy that might potentially account for 58.1% (salt solution) and 16.5% (artificial seawater) of the energy required by the downstream RO system. Our results demonstrated the great potential of bioelectrochemical desalination.

Efficient salt removal in a continuously operated upflow microbial desalination cell with an air cathode.

Microbial desalination cells (MDCs) hold great promise for drinking water production because of potential energy savings during the desalination process. In this study, we developed a continuously operated MDC--upflow microbial desalination cell (UMDC) for the purpose of salt removal. During the 4-month operation, the UMDC constantly removed salts and generated bio-electricity. At a hydraulic retention time (HRT) of 4 days (salt solution) and current production of ∼62 mA, the UMDC was able to remove more than 99% of NaCl from the salt solution that had an initial salt concentration of 30 g total dissolved solids (TDS)/L. In addition, the TDS removal rate was 7.50 g TDSL(-1)d(-1) (salt solution volume) or 5.25 g TDSL(-1)d(-1) (wastewater volume), and the desalinated water met the drinking water standard, in terms of TDS concentration. A high charge transfer efficiency of 98.6% or 81% was achieved at HRT 1 or 4d. The UMDC produced a maximum power density of 30.8 W/m(3). The phenomena of bipolar electrodialysis and proton transport in the UMDC were discussed. These results demonstrated the potential of the UMDC as either a sole desalination process or a pre-desalination reactor for downstream desalination processes.

CAMBIUM, a process-based model of daily xylem development in Eucalyptus.

In hardwoods such as Eucalyptus spp., xylem (wood) is a heterogeneous tissue consisting of multiple cell types. As such, xylem development involves multiple complex interactions. To describe and understand xylem development, and ultimately predict the resultant wood properties, a process-based approach to modelling wood property variation is potentially very useful. In this paper, a new model (CAMBIUM), which incorporates concepts of these processes, is described. CAMBIUM predicts how wood density and fibre and vessel anatomical properties vary from pith-to-bark at a daily time step as a function of changing environmental conditions and a set of simulated physiological processes. Simulations from an existing process-based model of stand development (CABALA) are used as inputs. A key feature of CAMBIUM is a model of the interaction between different xylem cell types. Some weaknesses were identified in the ability of the model to simulate vessel spatial patterns and frequencies, emphasizing the complexities inherent in this aspect of angiosperm wood formation. The model was, however, able to provide realistic estimates of short-term variation and temporal ranges in eucalypt fibre diameter and secondary wall development and wood density.

Temporal variation in δ13C, wood density and microfibril angle in variously irrigated Eucalyptus nitens.

Wood can serve as a record of past climate, recording tree responses to changing conditions. It is also valuable in understanding tree responses to environment to optimise forest management. Stable carbon isotope ratios (δ13C), wood density and microfibril angle (MFA) are potentially useful wood property parameters for these purposes. The goal of this study was to understand how δ13C varied over time in response to cycles of soil drying and wetting and to variation in temperature in Eucalyptus nitens Deane & Maiden, in concert with wood density and MFA. δ13C increases did not necessarily occur when water stress was highest, but, rather, when it was relieved. Our hypothesis is that this was a result of the use of previously fixed carbohydrate reserves when growth and metabolic activity was resumed after a period of dormancy. MFA in particular showed concomitant temporal variation with δ13C. A peak in δ13C may not coincide temporally with an increase in water stress, but with a decrease, when higher growth rates enable the final incorporation of earlier stored photosynthate into mature wood. This has implications for using δ13C as a tool to understand past environmental conditions using radial measurements of wood properties. However, interpreting this data with other wood properties may be helpful for understanding past tree responses.

Daily patterns of stem size variation in irrigated and unirrigated Eucalyptus globulus.

High resolution measurements of stem diameter variation provide a means to study short-term dynamics of tree growth and water status. In this 14-month study, daily changes in stem radius of Eucalyptus globulus Labill. seedlings were measured with electronic point dendrometers in a plantation in southern Tasmania, Australia. The daily patterns of stem expansion and shrinkage were classified into three phases: shrinkage; recovery; and increase in diameter from one maximum to the next, or increment. This study showed that rapid onset of even mild drought in irrigated trees caused distinct changes in daily patterns of stem diameter variation, particularly the duration of daily stem increment. The duration of the daily increment phase was directly related to increment magnitude. The dynamics of daily increment were significantly affected by mean minimum temperature, indicating a temperature limitation on metabolic processes underlying diameter growth in these trees. Most likely due to differences in conductance, the duration but not rate of the incremental daily expansion was greater in fast- than in slow-growing trees.