Modeling the response of Norway spruce tree-ring carbon and oxygen isotopes to selection harvest on a drained peatland forest.

Continuous cover forestry (CCF) has gained interest as an alternative to even-aged management particularly on drained peatland forests. However, relatively little is known about the physiological response of suppressed trees when larger trees are removed as a part of CCF practices. Consequently, studies concentrating on process-level modeling of the response of trees to selection harvesting are also rare. Here, we compared, modeled and measured harvest response of previously suppressed Norway spruce (Picea abies) trees to a selection harvest. We quantified the harvest response by collecting Norway spruce tree-ring samples in a drained peatland forest site and measuring the change in stable carbon and oxygen isotopic ratios of wood formed during 2010-20, including five post-harvest years. The measured isotopic ratios were compared with ecosystem-level process model predictions for ${\kern0em }^{13}$C discrimination and ${\kern0em }^{18}$O leaf water enrichment. We found that the model predicted similar but lower harvest response than the measurements. Furthermore, accounting for mesophyll conductance was important for capturing the variation in ${\kern0em }^{13}$C discrimination. In addition, we performed sensitivity analysis on the model, which suggests that the modeled ${\kern0em }^{13}$C discrimination is sensitive to parameters related to CO2 transport through stomata to the mesophyll.

Estimating intraseasonal intrinsic water-use efficiency from high-resolution tree-ring δ13 C data in boreal Scots pine forests.

Intrinsic water-use efficiency (iWUE), a key index for carbon and water balance, has been widely estimated from tree-ring δ13 C at annual resolution, but rarely at high-resolution intraseasonal scale. We estimated high-resolution iWUE from laser-ablation δ13 C analysis of tree-rings (iWUEiso ) and compared it with iWUE derived from gas exchange (iWUEgas ) and eddy covariance (iWUEEC ) data for two Pinus sylvestris forests from 2002 to 2019. By carefully timing iWUEiso via modeled tree-ring growth, iWUEiso aligned well with iWUEgas and iWUEEC at intraseasonal scale. However, year-to-year patterns of iWUEgas , iWUEiso , and iWUEEC were different, possibly due to distinct environmental drivers on iWUE across leaf, tree, and ecosystem scales. We quantified the modification of iWUEiso by postphotosynthetic δ13 C enrichment from leaf sucrose to tree rings and by nonexplicit inclusion of mesophyll and photorespiration terms in photosynthetic discrimination model, which resulted in overestimation of iWUEiso by up to 11% and 14%, respectively. We thus extended the application of tree-ring δ13 C for iWUE estimates to high-resolution intraseasonal scale. The comparison of iWUEgas , iWUEiso , and iWUEEC provides important insights into physiological acclimation of trees across leaf, tree, and ecosystem scales under climate change and improves the upscaling of ecological models.

Tree organ growth and carbon allocation dynamics impact the magnitude and δ13C signal of stem and soil CO2 fluxes.

Incomplete knowledge of carbon (C) allocation dynamics in trees hinders accurate modeling and future predictions of tree growth. We studied C allocation dynamics in a mature Pinus sylvestris L. dominated forest with a novel analytical approach, allowing the first comparison of: (i) magnitude and δ13C of shoot, stem and soil CO2 fluxes (Ashoot, Rstem and Rsoil), (ii) concentration and δ13C of compound-specific and/or bulk non-structural carbohydrates (NSCs) in phloem and roots and (iii) growth of stem and fine roots. Results showed a significant effect of phloem NSC concentrations on tracheid growth, and both variables significantly impacted Rstem. Also, concentrations of root NSCs, especially starch, had a significant effect on fine root growth, although no effect of root NSC concentrations or root growth was detected on Rsoil. Time series analysis between δ13C of Ashoot and δ13C of Rstem or δ13C of Rsoil revealed strengthened C allocation to stem or roots under high C demands. Furthermore, we detected a significant correlation between δ13C of Rstem and δ13C of phloem sucrose and glucose, but not for starch or water-soluble carbohydrates. Our results indicate the need to include C allocation dynamics into tree growth models. We recommend using compound-specific concentration and δ13C analysis to reveal C allocation processes that may not be detected by the conventional approach that utilizes bulk organic matter.

Explicitly accounting for needle sugar pool size crucial for predicting intra-seasonal dynamics of needle carbohydrates δ18 O and δ13 C.

We explore needle sugar isotopic compositions (δ18 O and δ13 C) in boreal Scots pine (Pinus sylvestris) over two growing seasons. A leaf-level dynamic model driven by environmental conditions and based on current understanding of isotope fractionation processes was built to predict δ18 O and δ13 C of two hierarchical needle carbohydrate pools, accounting for the needle sugar pool size and the presence of an invariant pinitol pool. Model results agreed well with observed needle water δ18 O, δ18 O and δ13 C of needle water-soluble carbohydrates (sugars + pinitol), and needle sugar δ13 C (R2 = 0.95, 0.84, 0.60, 0.73, respectively). Relative humidity (RH) and intercellular to ambient CO2 concentration ratio (Ci /Ca ) were the dominant drivers of δ18 O and δ13 C variability, respectively. However, the variability of needle sugar δ18 O and δ13 C was reduced on diel and intra-seasonal timescales, compared to predictions based on instantaneous RH and Ci /Ca , due to the large needle sugar pool, which caused the signal formation period to vary seasonally from 2 d to more than 5 d. Furthermore, accounting for a temperature-sensitive biochemical 18 O-fractionation factor and mesophyll resistance in 13 C-discrimination were critical. Interpreting leaf-level isotopic signals requires understanding on time integration caused by mixing in the needle sugar pool.

Linking canopy-scale mesophyll conductance and phloem sugar δ13 C using empirical and modelling approaches.

Interpreting phloem carbohydrate or xylem tissue carbon isotopic composition as measures of water-use efficiency or past tree productivity requires in-depth knowledge of the factors altering the isotopic composition within the pathway from ambient air to phloem contents and tree ring. One of least understood of these factors is mesophyll conductance (gm ). We formulated a dynamic model describing the leaf photosynthetic pathway including seven alternative gm descriptions and a simple transport of sugars from foliage down the trunk. We parameterised the model for a boreal Scots pine stand and compared simulated gm responses with weather variations. We further compared the simulated δ13 C of new photosynthates among the different gm descriptions and against measured phloem sugar δ13 C. Simulated gm estimates of the seven descriptions varied according to weather conditions, resulting in varying estimates of phloem δ13 C during cold/moist and warm/dry periods. The model succeeded in predicting a drought response and a postdrought release in phloem sugar δ13 C indicating suitability of the model for inverse prediction of leaf processes from phloem isotopic composition. We suggest short-interval phloem sampling during and after extreme weather conditions to distinguish between mesophyll conductance drivers for future model development.

Temperature and moisture dependence of daily growth of Scots pine (Pinus sylvestris L.) roots in Southern Finland.

Scots pine (Pinus sylvestris L.) is one of the most important conifers in Northern Europe. In boreal forests, over one-third of net primary production is allocated to roots. Pioneer roots expand the horizontal and vertical root systems and transport nutrients and water from belowground to aboveground. Fibrous roots, often colonized by mycorrhiza, emerge from the pioneer roots and absorb water and nutrients from the soil. In this study, we installed three flatbed scanners to detect the daily growth of both pioneer and fibrous roots of Scots pine during the growing season of 2018, a year with an unexpected summer drought in Southern Finland. The growth rate of both types of roots had a positive relationship with temperature. However, the relations between root elongation rate and soil moisture differed significantly between scanners and between root types indicating spatial heterogeneity in soil moisture. The pioneer roots were more tolerant to severe environmental conditions than the fibrous roots. The pioneer roots initiated elongation earlier and ceased it later than the fibrous roots. Elongation ended when the temperature dropped below the threshold temperature of 4 °C for pioneer roots and 6 °C for fibrous roots. During the summer drought, the fibrous roots halted root surface area growth at the beginning of the drought, but there was no drought effect on the pioneer roots over the same period. To compare the timing of root production and the aboveground organs' production, we used the CASSIA model, which estimates the aboveground tree carbon dynamics. In this study, root growth started and ceased later than growth of aboveground organs. Pioneer roots accounted for 87% of total root productivity. We suggest that future carbon allocation models should separate the roots by root types (pioneer and fibrous), as their growth patterns are different and they have different reactions to changes in the soil environment.

Current-year shoot hydraulic structure in two boreal conifers-implications of growth habit on water potential.

Metabolic scaling theory allows us to link plant hydraulic structure with metabolic rates in a quantitative framework. In this theoretical framework, we considered the hydraulic structure of current-year shoots in Pinus sylvestris and Picea abies, focusing on two properties unaccounted for by metabolic scaling theories: conifer needles are attached to the entire length of shoots, and the shoot as a terminal element does not display invariant properties. We measured shoot length and diameter as well as conduit diameter and density in two locations of 14 current-year non-leader shoots of pine and spruce saplings, and calculated conductivities of shoots from measured conduit properties. We evaluated scaling exponents for the hydraulic structure of shoots at the end of the water transport pathway from the data and applied the results to simulate water potential of shoots in the crown. Shoot shape was intermediate between cylindrical and paraboloid. Contrary to previous findings, we found that conduit diameter scaled with relative, not absolute, distance from the apex and absolute under-bark shoot diameter independently of species within the first-year shoots. Shoot hydraulic conductivity scaled with shoot diameter and hydraulic diameter. Larger shoots had higher hydraulic conductance. We further demonstrate by novel model calculations that ignoring foliage distribution along the hydraulic pathway overestimates water potential loss in shoots and branches and therefore overestimates related water stress effects. Scaling of hydraulic properties with shoot size enhances apical dominance and may contribute to the decline of whole-tree conductance in large trees.

Newtonian boreal forest ecology: The Scots pine ecosystem as an example.

Isaac Newton's approach to developing theories in his book Principia Mathematica proceeds in four steps. First, he defines various concepts, second, he formulates axioms utilising the concepts, third, he mathematically analyses the behaviour of the system defined by the concepts and axioms obtaining predictions and fourth, he tests the predictions with measurements. In this study, we formulated our theory of boreal forest ecosystems, called NewtonForest, following the four steps introduced by Newton. The forest ecosystem is a complicated entity and hence we needed altogether 27 concepts to describe the material and energy flows in the metabolism of trees, ground vegetation and microbes in the soil, and to describe the regularities in tree structure. Thirtyfour axioms described the most important features in the behaviour of the forest ecosystem. We utilised numerical simulations in the analysis of the behaviour of the system resulting in clear predictions that could be tested with field data. We collected retrospective time series of diameters and heights for test material from 6 stands in southern Finland and five stands in Estonia. The numerical simulations succeeded to predict the measured diameters and heights, providing clear corroboration with our theory.

Temperature dependence of needle and shoot elongation before bud break in Scots pine.

Knowledge about the early part of needle growth is deficient compared with what is known about shoot growth. It is however important to understand growth of different organs to be able to estimate the changes in whole tree growth in a changing environment. The onset of growth in spring has been observed to occur over some certain threshold value of momentary temperature or temperature accumulation. We measured the length growth of Scots pine (Pinus sylvestris L.) needles and shoots from March until bud break over 3 years. We first compared needle growth with concurrent shoot growth. Then, we quantified threshold temperature of growth (i) with a logistic regression based on momentary temperatures and (ii) with the temperature sum accumulation method. Temperature sum was calculated with combinations of various time steps, starting dates and threshold temperature values. Needle elongation began almost concurrently with shoot elongation and proceeded linearly in relation to shoot growth until bud break. When studying the threshold temperature for growth, the method with momentary temperature effect on growth onset yielded ambiguous results in our conditions. The best fit of an exponential regression between needle growth or length and temperature sum was obtained with threshold temperatures -1 to +2 °C, with several combinations of starting date and time step. We conclude that although growth onset is a momentary event the process leading to it is a long-term continuum where past time temperatures have to be accounted for, rather than a sudden switch from quiescence to active growth. Further, our results indicate that lower temperatures than the commonly used +5 °C are sufficient for actuating the growth process.

CASSIA--a dynamic model for predicting intra-annual sink demand and interannual growth variation in Scots pine.

The control of tree growth vs environment by carbon sources or sinks remains unresolved although it is widely studied. This study investigates growth of tree components and carbon sink-source dynamics at different temporal scales. We constructed a dynamic growth model 'carbon allocation sink source interaction' (CASSIA) that calculates tree-level carbon balance from photosynthesis, respiration, phenology and temperature-driven potential structural growth of tree organs and dynamics of stored nonstructural carbon (NSC) and their modifying influence on growth. With the model, we tested hypotheses that sink demand explains the intra-annual growth dynamics of the meristems, and that the source supply is further needed to explain year-to-year growth variation. The predicted intra-annual dimensional growth of shoots and needles and the number of cells in xylogenesis phases corresponded with measurements, whereas NSC hardly limited the growth, supporting the first hypothesis. Delayed GPP influence on potential growth was necessary for simulating the yearly growth variation, indicating also at least an indirect source limitation. CASSIA combines seasonal growth and carbon balance dynamics with long-term source dynamics affecting growth and thus provides a first step to understanding the complex processes regulating intra- and interannual growth and sink-source dynamics.

Duration of shoot elongation in Scots pine varies within the crown and between years.

BACKGROUND AND AIMS: Shoot elongation in boreal and temperate trees typically follows a sigmoid pattern where the onset and cessation of growth are related to accumulated effective temperature (thermal time). Previous studies on leader shoots suggest that while the maximum daily growth rate depends on the availability of resources to the shoot, the duration of the growth period may be an adaptation to long-term temperature conditions. However, other results indicate that the growth period may be longer in faster growing lateral shoots with higher availability of resources. This study investigates the interactions between the rate of elongation and the duration of the growth period in units of thermal time in lateral shoots of Scots pine (Pinus sylvestris). METHODS: Length development of 202 lateral shoots were measured approximately three times per week during seven growing seasons in 2-5 trees per year in a mature stand and in three trees during one growing season in a sapling stand. A dynamic shoot growth model was adapted for the analysis to determine (1) the maximum growth rate and (2) the thermal time reached at growth completion. The relationship between those two parameters and its variation between trees and years was analysed using linear mixed models. KEY RESULTS: The shoots with higher maximum growth rate within a crown continued to grow for a longer period in any one year. Higher July-August temperature of the previous summer implied a higher requirement of thermal time for growth completion. CONCLUSIONS: The results provide evidence that the requirement of thermal time for completion of lateral shoot extension in Scots pine may interact with resource availability to the shoot both from year to year and among shoots in a crown each year. If growing season temperatures rise in the future, this will affect not only the rate of shoot growth but its duration also.