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Traditional photosynthesis-driven growth models have considerable uncertainties in predicting tree growth under changing climates, partially because sink activities are directly affected by the environment but not adequately addressed in growth modelling. Therefore, we developed a semi-mechanistic model coupling stomatal optimality, temperature control of enzymatic activities and phenology of cambial growth. Parameterized using Bayesian inference and measured data on Picea abies and Pinus sylvestris in peatland and mineral soils in Finland, the coupled model simulates transpiration and assimilation rates and stem radial dimension (SRD) simultaneously at 30 min resolution. The results suggest that both the sink and phenological formulations with environmental effects are indispensable for capturing SRD dynamics across hourly to seasonal scales. Simulated using the model, growth was more sensitive than assimilation to temperature and soil water, suggesting carbon gain is not driving growth at the current temporal scale. Also, leaf-specific production was occasionally positively correlated with growth duration but not with growth onset timing or annual cambial area increment. Thus, as it is hardly explained by carbon gain, phenology itself should be included in sink-driven growth models of the trees in the boreal zone and possibly other environments where sink activities and photosynthesis are both restrained by harsh conditions., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

The transport processes of methane (CH 4 ) in tree stems remain largely unknown, although they are critical in assessing the whole-forest CH4 dynamics. We used a physically based dynamic model to study the spatial and diurnal dynamics of stem CH 4 transport and fluxes. We parameterised the model using data from laboratory experiments with Pinus sylvestris and Betula pendula and compared the model to experimental data from a field study. Stem CH 4 flux in laboratory and field conditions were explained by the axial advective CH 4 transport from soil with xylem sap flow and the radial CH 4 diffusion through the stem conditions. Diffusion resistance caused by the bark permeability did not significantly affect gas transport or stem CH 4 flux in the laboratory experiments. The role of axial diffusion of CH 4 in trees was unresolved and requires further studies. Due to the transit time of CH 4 in the stem, the diurnal dynamics of stem CH 4 fluxes can deviate markedly from the diurnal dynamics of sap flow., (© 2023 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Carbon isotope composition of tree-ring (δ 13 C Ring ) is a commonly used proxy for environmental change and ecophysiology. δ 13 C Ring reconstructions are based on a solid knowledge of isotope fractionations during formation of primary photosynthates (δ 13 C P ), such as sucrose. However, δ 13 C Ring is not merely a record of δ 13 C P . Isotope fractionation processes, which are not yet fully understood, modify δ 13 C P during sucrose transport. We traced, how the environmental intra-seasonal δ 13 C P signal changes from leaves to phloem, tree-ring and roots, for 7 year old Pinus sylvestris, using δ 13 C analysis of individual carbohydrates, δ 13 C Ring laser ablation, leaf gas exchange and enzyme activity measurements. The intra-seasonal δ 13 C P dynamics was clearly reflected by δ 13 C Ring , suggesting negligible impact of reserve use on δ 13 C Ring . However, δ 13 C P became increasingly 13 C-enriched during down-stem transport, probably due to post-photosynthetic fractionations such as sink organ catabolism. In contrast, δ 13 C of water-soluble carbohydrates, analysed for the same extracts, did not reflect the same isotope dynamics and fractionations as δ 13 C P , but recorded intra-seasonal δ 13 C P variability. The impact of environmental signals on δ 13 C Ring , and the 0.5 and 1.7‰ depletion in photosynthates compared ring organic matter and tree-ring cellulose, respectively, are useful pieces of information for studies exploiting δ 13 C Ring ., (© 2023 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Tree stems have been identified as sources of volatile organic compounds (VOCs) that play important roles in tree defence and atmospheric chemistry. Yet, we lack understanding on the magnitude and environmental drivers of stem VOC emissions in various forest ecosystems. Due to the increasing importance of extreme drought, we studied drought effects on the VOC emissions from mature Scots pine (Pinus sylvestris L.) stems. We measured monoterpenes, acetone, acetaldehyde and methanol emissions with custom-made stem chambers, online PTR-MS and adsorbent sampling in a drought-prone forest over the hot-dry summer of 2018 and compared the emission rates and dynamics between trees in naturally dry conditions and under long-term irrigation (drought release). The pine stems were significant monoterpene sources. The stem monoterpene emissions potentially originated from resin, based on their similar monoterpene spectra. The emission dynamics of all VOCs followed temperature at a daily scale, but monoterpene and acetaldehyde emission rates decreased nonlinearly with drought over the summer. Despite the dry conditions, large peaks of monoterpene, acetaldehyde and acetone emissions occurred in late summer potentially due to abiotic or biotic stressors. Our results highlight the potential importance of stem emissions in the ecosystem VOC budget, encouraging further studies in diverse environments., (© 2021 John Wiley & Sons Ltd.)

Tree stems are an overlooked source of volatile organic compounds (VOCs). Their contribution to ecosystem processes and total VOC fluxes is not well studied, and assessing it requires better understanding of stem emission dynamics and their driving processes. To gain more mechanistic insight into stem emission patterns, we measured monoterpene, methanol and acetaldehyde emissions from the stems of mature Scots pines (Pinus sylvestris L.) in a boreal forest over three summers. We analysed the effects of temperature, soil water content, tree water status, transpiration and growth on the VOC emissions and used generalized linear models to test their relative importance in explaining the emissions. We show that Scots pine stems are considerable sources of monoterpenes, methanol and acetaldehyde, and their emissions are strongly regulated by temperature. However, even small changes in water availability affected the emission potentials: increased soil water content increased the monoterpene emissions within a day, whereas acetaldehyde and methanol emissions responded within 2-4 days. This lag corresponded to their transport time in the xylem sap from the roots to the stem. Moreover, the emissions of monoterpenes, methanol and acetaldehyde were influenced by the cambial growth rate of the stem with 6-10-day lags., (© 2020 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Functional relationships between wood density and measures of xylem hydraulic safety and efficiency are ambiguous, especially in wet tropical forests. In this meta-analysis, we move beyond wood density per se and identify relationships between xylem allocated to fibers, parenchyma, and vessels and measures of hydraulic safety and efficiency. We analyzed published data of xylem traits, hydraulic properties and measures of drought resistance from neotropical tree species retrieved from 346 sources. We found that xylem volume allocation to fiber walls increases embolism resistance, but at the expense of specific conductivity and sapwood capacitance. Xylem volume investment in fiber lumen increases capacitance, while investment in axial parenchyma is associated with higher specific conductivity. Dominant tree taxa from wet forests prioritize xylem allocation to axial parenchyma at the expense of fiber walls, resulting in a low embolism resistance for a given wood density and a high vulnerability to drought-induced mortality. We conclude that strong trade-offs between xylem allocation to fiber walls, fiber lumen, and axial parenchyma drive drought resistance in neotropical trees. Moreover, the benefits of xylem allocation to axial parenchyma in wet tropical trees might not outweigh the consequential low embolism resistance under more frequent and severe droughts in a changing climate., (© 2019 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Understanding stomatal regulation is fundamental to predicting the impact of changing environmental conditions on vegetation. However, the influence of soil temperature (ST) and soil water content (SWC) on canopy conductance (g s ) through changes in belowground hydraulic conductance (k bg ) remains poorly understood, because k bg has seldom been measured in field conditions. Our aim was to (a) examine the dependence of k bg on ST and SWC, (b) examine the dependence of g s on k bg and (c) test a recent stomatal optimization model according to which g s and soil-to-leaf hydraulic conductance are strongly coupled. We estimated k bg from continuous sap flow and xylem diameter measurements in three boreal species. k bg increased strongly with increasing ST when ST was below +8°C, and typically increased with increasing SWC when ST was not limiting. g s was correlated with k bg in all three species, and modelled and measured g s were well correlated in Pinus sylvestris (a model comparison was only possible for this species). These results imply an important role for k bg in mediating linkages between the soil environment and leaf gas exchange. In particular, our finding that ST strongly influences k bg in mature trees may help us to better understand tree behaviour in cold environments., (© 2019 John Wiley & Sons Ltd.)

Most plant-based emissions of volatile organic compounds are considered mainly temperature dependent. However, certain oxygenated volatile organic compounds (OVOCs) have high water solubility; thus, also stomatal conductance could regulate their emissions from shoots. Due to their water solubility and sources in stem and roots, it has also been suggested that their emissions could be affected by transport in the xylem sap. Yet further understanding on the role of transport has been lacking until present. We used shoot-scale long-term dynamic flux data from Scots pines (Pinus sylvestris) to analyse the effects of transpiration and transport in xylem sap flow on emissions of 3 water-soluble OVOCs: methanol, acetone, and acetaldehyde. We found a direct effect of transpiration on the shoot emissions of the 3 OVOCs. The emissions were best explained by a regression model that combined linear transpiration and exponential temperature effects. In addition, a structural equation model indicated that stomatal conductance affects emissions mainly indirectly, by regulating transpiration. A part of the temperature's effect is also indirect. The tight coupling of shoot emissions to transpiration clearly evidences that these OVOCs are transported in the xylem sap from their sources in roots and stem to leaves and to ambient air., (© 2018 John Wiley & Sons Ltd.)

Preconditions of phloem transport in conifers are relatively unknown. We studied the variation of needle and inner bark axial osmotic gradients and xylem water potential in Scots pine and Norway spruce by measuring needle and inner bark osmolality in saplings and mature trees over several periods within a growing season. The needle and inner bark osmolality was strongly related to xylem water potential in all studied trees. Sugar concentrations were measured in Scots pine, and they had similar dynamics to inner bark osmolality. The sucrose quantity remained fairly constant over time and position, whereas the other sugars exhibited a larger change with time and position. A small osmotic gradient existed from branch to stem base under pre-dawn conditions, and the osmotic gradient between upper stem and stem base was close to zero. The turgor in branches was significantly driven by xylem water potential, and the turgor loss point in branches was relatively close to daily minimum needle water potentials typically reported for Scots pine. Our results imply that xylem water potential considerably impacts the turgor pressure gradient driving phloem transport and that gravitation has a relatively large role in phloem transport in the stems of mature Scots pine trees., (© 2017 John Wiley & Sons Ltd.)

Substantial uncertainty surrounds our knowledge of tree stem growth, with some of the most basic questions, such as when stem radial growth occurs through the daily cycle, still unanswered. We employed high-resolution point dendrometers, sap flow sensors, and developed theory and statistical approaches, to devise a novel method separating irreversible radial growth from elastic tension-driven and elastic osmotically driven changes in bark water content. We tested this method using data from five case study species. Experimental manipulations, namely a field irrigation experiment on Scots pine and a stem girdling experiment on red forest gum trees, were used to validate the theory. Time courses of stem radial growth following irrigation and stem girdling were consistent with a-priori predictions. Patterns of stem radial growth varied across case studies, with growth occurring during the day and/or night, consistent with the available literature. Importantly, our approach provides a valuable alternative to existing methods, as it can be approximated by a simple empirical interpolation routine that derives irreversible radial growth using standard regression techniques. Our novel method provides an improved understanding of the relative source-sink carbon dynamics of tree stems at a sub-daily time scale., (© 2016 The Authors Plant, Cell & Environment Published by John Wiley & Sons Ltd.)

Coniferous tree stems contain large amounts of oleoresin under positive pressure in the resin ducts. Studies in North-American pines indicated that the stem oleoresin exudation pressure (OEP) correlates negatively with transpiration rate and soil water content. However, it is not known how the OEP changes affect the emissions of volatile vapours from the trees. We measured the OEP, xylem diameter changes indicating changes in xylem water potential and monoterpene emissions under field conditions in mature Scots pine (Pinus sylvestris L.) trees in southern Finland. Contrary to earlier reports, the diurnal OEP changes were positively correlated with temperature and transpiration rate. OEP was lowest at the top part of the stem, where water potentials were also more negative, and often closely linked to ambient temperature and stem monoterpene emissions. However, occasionally OEP was affected by sudden changes in vapour pressure deficit (VPD), indicating the importance of xylem water potential on OEP as well. We conclude that the oleoresin storage pools in tree stems are in a dynamic relationship with ambient temperature and xylem water potential, and that the canopy monoterpene emission rates may therefore be also regulated by whole tree processes and not only by the conditions prevailing in the upper canopy., (© 2015 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

The quantification of cambial growth over short time periods has been hampered by problems to discern between growth and the swelling and shrinking of a tree stem. This paper presents a model, which separates cambial growth and reversible water-potential induced diurnal changes from simultaneously measured whole stem and xylem radial variations, from field-measured Scots pine trees in Finland. The modelled growth, which includes osmotic concentration changes, was compared with (direct) dendrometer measurements and microcore samples. In addition, the relationship of modelled growth and dendrometer measurements to environmental factors was analysed. The results showed that the water-potential induced changes of tree radius were successfully separated from stem growth. Daily growth predicted by the model exhibited a high correlation with the modelled daily changes of osmotic concentration in phloem, and a temperature dependency in early summer. Late-summer growth saw higher dependency on water availability and temperature. Evaluation of the model against dendrometer measurements showed that the latter masked a true environmental signal in stem growth due to water-potential induced changes. The model provides better understanding of radial growth physiology and offers potential to examine growth dynamics and changes due to osmotic concentration, and how the environment affects growth., (© 2015 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Carbon uptake and transpiration in plant leaves occurs through stomata that open and close. Stomatal action is usually considered a response to environmental driving factors. Here we show that leaf gas exchange is more strongly related to whole tree level transport of assimilates than previously thought, and that transport of assimilates is a restriction of stomatal opening comparable with hydraulic limitation. Assimilate transport in the phloem requires that osmotic pressure at phloem loading sites in leaves exceeds the drop in hydrostatic pressure that is due to transpiration. Assimilate transport thus competes with transpiration for water. Excess sugar loading, however, may block the assimilate transport because of viscosity build-up in phloem sap. Therefore, for given conditions, there is a stomatal opening that maximizes phloem transport if we assume that sugar loading is proportional to photosynthetic rate. Here we show that such opening produces the observed behaviour of leaf gas exchange. Our approach connects stomatal regulation directly with sink activity, plant structure and soil water availability as they all influence assimilate transport. It produces similar behaviour as the optimal stomatal control approach, but does not require determination of marginal cost of water parameter., (© 2012 Blackwell Publishing Ltd.)

Efficient water transport from the soil to the leaves is essential for plant function, while building and maintaining the water transport structure in the xylem require a major proportion of the assimilated carbon of the tree. Xylem transport also faces additional challenges as water in the xylem is under tension and therefore cavitation cannot be completely avoided. We constructed a model that calculates the xylem structure that maximizes carbon-use efficiency while simultaneously taking into account pit structure in increasing the resistance to water transport and constricting the spreading of embolisms. The optimal xylem structure predicted by the model was found to correspond well to the generally observed trends: xylem conduits grew in size from the apex towards the base while simultaneously decreasing in number, and vulnerability to cavitation increased with conduit size. These trends were caused primarily by the axial water potential gradient in the xylem. The pits have to be less porous near the apex where water potential is lower to restrict the spreading of embolisms, while whole-plant carbon-use efficiency demands that conduit size decreases and conduit number increases simultaneously. The model predictions remained qualitatively the same regardless of the exact optimality criterion used for defining carbon-use efficiency., (© 2011 Blackwell Publishing Ltd.)

Measurements of diurnal diameter variations of the xylem and phloem are a promising tool for studying plant hydraulics and xylem-phloem interactions in field conditions. However, both the theoretical framework and the experimental verification needed to interpret phloem diameter data are incomplete. In this study, we analytically evaluate the effects of changing the radial conductance between the xylem and the phloem on phloem diameter variations and test the theory using simple manipulation experiments. Our results show that phloem diameter variations are mainly caused by changes in the radial flow rate of water between the xylem and the phloem. Reducing the hydraulic conductance between these tissues decreases the amplitude of phloem diameter variation and increases the time lag between xylem and phloem diameter variation in a predictable manner. Variation in the amplitude and timing of diameter variations that cannot be explained by changes in the hydraulic conductance, could be related to changes in the osmotic concentration in the phloem., (© 2011 Blackwell Publishing Ltd.)

Embolisms decrease plant hydraulic conductance and therefore reduce the ability of the xylem to transport water to leaves provided that embolized conduits are not refilled. However, as a xylem conduit is filled with gas during cavitation, water is freed to the transpiration stream and this transiently increases xylem water potential. This capacitive effect of embolism formation on plant function has not been explicitly quantified in the past. A dynamic model is presented that models xylem water potential, xylem sap flow and cavitation, taking into account both the decreasing hydraulic conductance and the water release effect of xylem embolism. The significance of the capacitive effect increases in relation to the decreasing hydraulic conductance effect when transpiration rate is low in relation to the total amount of water in xylem conduits. This ratio is typically large in large trees and during drought.