SummaryThe 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' (CAS-SIA) 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.

Abstract. Estimates of volatile organic compound (VOC)emissions from forests are based on the assumption that foliage has a steady emission potential over its lifetime, and that emissions are mainly modified by short-term variations in light and temperature. However, in many field studies this has been challenged, and high emissions and atmospheric concentrations have been measured during periods of low biological activity, such as in springtime. We conducted measurements during three years, using an online gas-exchange monitoring system to observe volatile organic emissions from a mature (1 year-old) and a growing Scots pine shoot. The emission rates of organic vapors from vegetative buds of Scots pine during the dehardening and rapid shoot growth stages were one to two orders of magnitude higher than those from mature foliage; this difference decreased and finally disappeared when the new shoot was maturing in late summer. On average, the springtime monoterpene emission rate of the bud was about 500 times higher than that of the mature needles; during the most intensive needle elongation period, the monoterpene emission rate of the growing needles was 3.5 higher than that of the mature needles, and in September the monoterpene emission rate of the same years' needles was even lower (50 %) than that of the previous years' needles. For other measured compounds (methanol, acetone and methylbutenol) the values were of the same order of magnitude, except before bud break in spring, when the emission rates of buds for those compounds were on average about 20-30 times higher than that of mature needles. During spring and early summer the buds and growing shoots are a strong source of several VOCs, and if they are not accounted for in emission modeling a significant proportion of the emissions -from a few percent to even half of the annual cumulative emissions -will remain concealed. The diurnal emission pattern of growing shoots differed from the diurnal cycle in temperature as well as from the diurnal emission pattern of mature shoots, which may be related to processes involved in shoot or needle elongation. Our findings imply that global estimations of monoterpene emission rates from forests are in need of revision, and that the physiological state of the plants should be taken into account when emissions of the reactive gases such as monoterpenes are estimated.

Photosynthesis is not entirely synchronized with carbon sinks, implying that trees are capable of storing non-structural carbohydrates (NSC), such as soluble sugars and starch. These storages provide a buffer between carbohydrate supply and demand and also allow trees to resist drought through osmoregulation. However, estimates of the total pool size and seasonal dynamics of the NSC storage of mature trees are still rare. Part of NSC is allocated outside roots, mainly to symbiotic, root-associated mycorrhizal fungi. The quantity and dynamics of this allocation are difficult to estimate in field conditions due to the close interaction between the symbionts. The aims of this study were to (1) determine the temporal development of NSC concentrations in tree organs, (2) upscale the storage compounds to whole-tree level and (3) analyse the significance of NSC allocation to belowground symbionts as part of the carbon balance in mature pines in a boreal Scots pine stand in southern Finland. We took samples every 2-4 weeks of needles, fine roots, stem wood, shoot wood and phloem from 1 to 3 trees in 2015. Concentrations of soluble sugars and starch were analyzed from the samples and upscaled to tree level. For quantifying the third aim, we used a whole-tree carbon balance model CASSIA that incorporates daily photosynthesis, respiration and organ-specific growth as functions of environmental factors. In this study, we included the allocation to belowground symbionts as an additional carbon sink and scaled the flux using the NSC pool over the whole tree. We observed that organ-specific NSC concentrations were highest in phloem, needles and fine roots. Total NSC increased in spring, peaked during midsummer and decreased again in autumn without any notable decrease during the most intensive growth period at midsummer. In the model analysis, 6% of yearly photosynthesis was allocated to the root-associated symbionts. The study highlights the applicability of the carbon balance approach in evaluating the importance of processes that cannot yet be directly measured.

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.

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.

Abstract. Biogenic volatile organic compounds (BVOCs) emitted by the forests are known to have strong impacts in the atmosphere. However, lots of missing reactivity is found, especially in the forest air. Therefore better characterization of sources and identification/quantification of unknown reactive compounds is needed. While isoprene and monoterpene (MT) emissions of boreal needle trees have been studied quite intensively, there is much less knowledge on the emissions of boreal deciduous trees and emissions of larger terpenes and oxygenated volatile organic compounds (OVOCs). Here we quantified the downy birch (Betula pubescens) leaf emissions of terpenes, oxygenated terpenes and green leaf volatiles (GLVs) at the SMEAR II boreal forest site using in situ gas chromatographs with mass spectrometers. Sesquiterpenes (SQTs) and oxygenated sesquiterpenes (OSQTs) were the main emitted compounds. Mean emission rates of SQTs and OSQTs were significantly higher in the early growing season (510 and 650 ng gdw-1 h−1, respectively) compared to in the main (40 and 130 ng gdw-1 h−1, respectively) and late (14 and 46 ng gdw-1 h−1, respectively) periods, indicating that early leaf growth is a strong source of these compounds. The emissions had a very clear diurnal variation with afternoon maxima being on average 4 to 8 times higher than seasonal means for SQTs and OSQTs, respectively. β-Caryophyllene and β-farnesene were the main SQTs emitted. The main emitted OSQTs were tentatively identified as 14-hydroxy-β-caryophyllene acetate (M=262 g mol−1) and 6-hydroxy-β-caryophyllene (M=220 g mol−1). Over the whole growing season, the total MT emissions were only 24 % and 17 % of the total SQT and OSQT emissions, respectively. A stressed tree growing in a pot was also studied, and high emissions of α-farnesene and an unidentified SQT were detected together with high emissions of GLVs. Due to the relatively low volatility and the high reactivity of SQTs and OSQTs, downy birch emissions are expected to have strong impacts on atmospheric chemistry, especially on secondary organic aerosol (SOA) production.

Estimates of volatile organic compound (VOC) emissions from forests are based on the assumption that foliage has a steady emission potential over its lifetime, and that emissions are mainly modified by short-term variations in light and temperature. However, in many field studies this has been challenged, and high emissions and atmospheric concentrations have been measured during periods of low biological activity, such as in springtime. We conducted measurements during three years, using an online gas-exchange monitoring system to observe volatile organic emissions from a mature (1 year-old) and a growing Scots pine shoot. The emission rates of organic vapors from vegetative buds of Scots pine during the dehardening and rapid shoot growth stages were one to two orders of magnitude higher than those from mature foliage; this difference decreased and finally disappeared when the new shoot was maturing in late summer. On average, the springtime monoterpene emission rate of the bud was about 500 times higher than that of the mature needles; during the most intensive needle elongation period, the monoterpene emission rate of the growing needles was 3.5 higher than that of the mature needles, and in September the monoterpene emission rate of the same years' needles was even lower (50 %) than that of the previous years' needles. For other measured compounds (methanol, acetone and methylbutenol) the values were of the same order of magnitude, except before bud break in spring, when the emission rates of buds for those compounds were on average about 20-30 times higher than that of mature needles. During spring and early summer the buds and growing shoots are a strong source of several VOCs, and if they are not accounted for in emission modeling a significant proportion of the emissions -from a few percent to even half of the annual cumulative emissions -will remain concealed. The diurnal emission pattern of growing shoots differed from the diurnal cycle in temperature as well as from the diurnal emission pattern of mature shoots, which may be related to processes involved in shoot or needle elongation. Our findings imply that global estimations of monoterpene emission rates from forests are in need of revision, and that the physiological state of the plants should be taken into account when emissions of the reactive gases such as monoterpenes are estimated.Published by Copernicus Publications on behalf of the European Geosciences Union.

 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 (g m).  We formulated a dynamic model describing the leaf photosynthetic pathway including seven alternative g m descriptions and a simple transport of sugars from foliage down the trunk. We parameterized the model for a boreal Scots pine stand and compared simulated g m responses to weather variations. We further compared the simulated δ 13 C of new photosynthates among the different g m descriptions and against measured phloem sugar δ 13 C.  Simulated g m estimates of the seven descriptions varied according to weather conditions resulting in varying estimates of phloem δ 13 C during cold / moist and warm / dry periods. The model succeeded in predicting a drought response and a post-drought release in phloem sugar δ 13 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.