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Plant phenological development is orchestrated through subtle changes in photoperiod, temperature, soil moisture and nutrient availability. Presently, the exact timing of plant development stages and their response to climate and management practices are crudely represented in land surface models. As visual observations of phenology are laborious, there is a need to supplement long-term observations with automated techniques such as those provided by digital repeat photography at high temporal and spatial resolution. We present the first synthesis from a growing observational network of digital cameras installed on towers across Europe above deciduous and evergreen forests, grasslands and croplands, where vegetation and atmosphere CO2 fluxes are measured continuously. Using colour indices from digital images and using piecewise regression analysis of time series, we explored whether key changes in canopy phenology could be detected automatically across different land use types in the network. The piecewise regression approach could capture the start and end of the growing season, in addition to identifying striking changes in colour signals caused by flowering and management practices such as mowing. Exploring the dates of green-up and senescence of deciduous forests extracted by the piecewise regression approach against dates estimated from visual observations, we found that these phenological events could be detected adequately (RMSE < 8 and 11 days for leaf out and leaf fall, respectively). We also investigated whether the seasonal patterns of red, green and blue colour fractions derived from digital images could be modelled mechanistically using the PROSAIL model parameterised with information of seasonal changes in canopy leaf area and leaf chlorophyll and carotenoid concentrations. From a model sensitivity analysis we found that variations in colour fractions, and in particular the late spring `green hump' observed repeatedly in deciduous broadleaf canopies across the network, are essentially dominated by changes in the respective pigment concentrations. Using the model we were able to explain why this spring maximum in green signal is often observed out of phase with the maximum period of canopy photosynthesis in ecosystems across Europe. Coupling such quasi-continuous digital records of canopy colours with co-located CO2 flux measurements will improve our understanding of how changes in growing season length are likely to shape the capacity of European ecosystems to sequester CO2 in the future.

Tree canopies are known to emit large amounts of VOCs (volatile organic compounds) such as monoterpenes into the surrounding air. High VOC emission rates from boreal forests have been observed during the transition from winter to summer activity. The most important sources of these are considered to be the green foliage, understory vegetation and soil organisms, but emissions from the living stand woody compartments have so far not been quantified. We analyzed whether the non-foliar components could partially explain the springtime high emission rates. We measured the monoterpene emissions from Scots pine (Pinus sylvestris L.) stem and shoots during the dehardening phase of trees in field conditions in two consecutive springs. We observed a large, transient monoterpene burst from the stem, while the shoot monoterpene emissions remained low. The burst lasted about 12 h. Simultaneously, an unusual nighttime sap flow and a non-systematic diurnal pattern of tree diameter were detected. Hence, we suggest that the monoterpene burst was a consequence of the recovery of the stem from wintertime, and likely related to the refilling of embolized tracheids and/or phenological changes in the living cells of the stem. This indicates that the dominant processes and environmental drivers triggering the monoterpene emissions are different between the stem and the foliage.

In spite of advances in greenhouse gas research, the spatiotemporal CH4 and N2O dynamics of boreal landscapes remain challenging, e.g., we need clarification of whether forest–mire transitions are occasional hotspots of landscape CH4 and N2O emissions during exceptionally high and low ground water level events. In our study, we tested the differences and drivers of CH4 and N2O dynamics of forest/mire types in field conditions along the soil moisture gradient of the forest–mire ecotone. Soils changed from Podzols to Histosols and ground water rose downslope from a depth of 10 m in upland sites to 0.1 m in mires. Yearly meteorological conditions changed from being exceptionally wet to typical and exceptionally dry for the local climate. The median fluxes measured with a static chamber technique varied from −51 to 586 μg m−2 h−1 for CH4 and from 0 to 6 μg m−2 h−1 for N2O between forest and mire types throughout the entire wet–dry period. In spite of the highly dynamic soil water fluctuations in carbon rich soils in forest–mire transitions, there were no large peak emissions in CH4 and N2O fluxes and the flux rates changed minimally between years. Methane uptake was significantly lower in poorly drained transitions than in the well-drained uplands. Water-saturated mires showed large CH4 emissions, which were reduced entirely during the exceptional summer drought period. Near-zero N2O fluxes did not differ significantly between the forest and mire types probably due to their low nitrification potential. When upscaling boreal landscapes, pristine forest–mire transitions should be regarded as CH4 sinks and minor N2O sources instead of CH4 and N2O emission hotspots.

In Finland, ditch network maintenance (DNM) is carried out annually on 60–70,000 ha of drained peatland to promote tree growth for forestry purposes. However, it is important to avoid ditching that contributes little to the stand growth and productivity, both to improve the economical profitability of forestry and to mitigate DNM-induced nutrient release to watercourses. We hypothesised that mature forest stands with substantial evapotranspiration potential do not necessarily need DNM, even if the ditch networks are in poor condition.We estimated evapotranspiration (EVT) of forest vegetation during the growing seasons (June–September) of 2007–2011 in four forested artificial peatland catchments dominated by Scots pine stands (Pinus sylvestris L.) (stand volume 93–151 m3 ha-1) located in southern, western, central and northern Finland. Precipitation (P), runoff (R) and water table level (WTL) were monitored continuously in the field. The water storage change (ΔS) was estimated on the basis of WTL measurements and peat pF curves determined from in situ peat samples. In addition, tree stand transpiration (T) was estimated in two of the catchments using the sap flow method. EVT was estimated as the residual term of the water balance equation.During the growing season, EVT (153–295 mm) was 49–161 % of the total accumulated P (155–368 mm), and decreased from south to north. Within each growing season, EVT was always largest in July or August. Tree transpiration was about 50 % of the total forest EVT in the two sites where it was measured directly. The mean WTL was at depth 36–63 cm during the growing seasons. Clear-cutting of a 100m3 ha-1 stand on one site resulted in an average rise of WTL by 18 cm.The results suggested that, in the southernmost site in particular, no drainage network management would be necessary to sustain satisfactory drainage conditions for tree growth because growing-season precipitation is transferred back to the atmosphere by forest EVT. In the northernmost site, ditch networks were considered important in controlling drainage conditions because of the low EVT potential of

The productivity of boreal forests is considered to be limited by low nitrogen (N) availability. Increased atmospheric N deposition has altered the functioning and N cycling of these N-sensitive ecosystems by increasing the availability of reactive nitrogen. The most important components of N pools and fluxes were measured in a boreal Scots pine stand in Hyytiälä, Southern Finland. The measurements at the site allowed direct estimations of nutrient pools in the soil and biomass, inputs from the atmosphere and outputs as drainage flow and gaseous losses from two micro-catchments. N was accumulating in the system, mainly in woody biomass, at a rate of 7 kg N ha−1 yr−1. Nitrogen input as atmospheric deposition was 7.4 kg N ha−1 yr−1. Dry deposition and organic N in wet deposition contributed over half of the inputs in deposition. Total outputs were 0.4 kg N ha−1 yr−1, the most important outputs being N2O emission to the atmosphere and organic N flux in drainage flow. Nitrogen uptake and retranslocation were equally important sources of N for plant growth. Most of the assimilated N originated from decomposition of organic matter, and the fraction of N that could originate directly from deposition was about 30%. In conclusion, atmospheric N deposition fertilizes the site considerably, but there are no signs of N saturation. Further research is needed to estimate soil N2 fluxes (emission and fixation), which may amount up to several kg N ha−1 yr−1.

Five years of carbon dioxide exchange measured with the eddy covariance technique at the world's northernmost urban flux station SMEAR III located in Helsinki, Finland, were analyzed. The long-term measurements and high-latitude location enabled us to examine the seasonal and annual variations of CO2 exchange, and to identify different factors controlling the measured exchange. Online traffic counts and soil respiration measurements were utilized in the study. Furthermore, the advantage of the station is that the complex surrounding area enables us to distinguish three different surface cover areas that can be evaluated separately. We also tested different methods (artificial neural networks and median diurnal cycles) to fill gaps in CO2 flux time series and examined their effect on annual emission estimates. The measured fluxes were highly dependent on the prevailing wind direction with the highest fluxes downwind from a large road and lowest downwind from the area of high fraction of vegetation cover. On an annual level, the area of the road emitted 3500 g C m−2 whereas the area of high fraction of vegetation cover emitted only 870 g C m−2 showing the effect of surface cover to be large in urban areas. Seasonal differences in the CO2 exchange downwind from the road were mainly caused by reduced traffic rates in summer, whereas in other directions seasonality was more determined by vegetation activity. Differences between the gap filling methods were small, but slightly better (0.6 μmol m−2 s−1 smaller RMSE) results were obtained when the artificial neural network with traffic counts was used instead of the one without traffic network and method based on median diurnal cycles. The measurement site was a net carbon source with average annual emissions of 1760 g C m−2, with a biased error of 6.1 g C m−2 caused by the gap filling. The annual value varied 16% between the different years.

We analyzed the dynamics of carbon balance components: gross primary production (GPP) and total ecosystem respiration (TER), of a boreal Scots pine forest in Southern Finland. The main focus is on investigations of environmental drivers of GPP and TER and how they affect the inter-annual variation in the carbon balance in autumn (September–December). We used standard climate data and CO2 exchange measurements collected by the eddy covariance (EC) technique over 11 years. EC data revealed that increasing autumn temperature significantly enhances TER: the temperature sensitivity was 9.5 gC m−2 °C−1 for the period September–October (early autumn when high radiation levels still occur) and 3.8 gC m−2 °C−1 for November–December (late autumn with suppressed radiation level). The cumulative GPP was practically independent of the temperature in early autumn. In late autumn, air temperature could explain part of the variation in GPP but the temperature sensitivity was very weak, less than 1 gC m−2 °C−1. Two models, a stand photosynthesis model (COCA) and a global vegetation model (ORCHIDEE), were used for estimating stand GPP and its sensitivity to the temperature. The ORCHIDEE model was tested against the observations of GPP derived from EC data. The stand photosynthesis model COCA predicted that under a predescribed 3–6 °C temperature increase, the temperature sensitivity of 4–5 gC m−2 °C−1 in GPP may appear in early autumn. The analysis by the ORCHIDEE model revealed the model sensitivity to the temporal treatment of meteorological forcing. The model predictions were similar to observed ones when the site level 1/2-hourly time step was applied, but the results calculated by using daily meteorological forcing, interpolated to 1/2-hourly time step, were biased. This is due to the nonlinear relationship between the processes and the environmental factors.

No abstract available.

Gross primary production (GPP) is the primary source of all carbon fluxes in the ecosystem. Understanding variation in this flux is vital to understanding variation in the carbon sink of forest ecosystems, and this would serve as input to forest production models. Using GPP derived from eddy-covariance (EC) measurements, it is now possible to determine the most important factor to scale GPP across sites. We use long-term EC measurements for six coniferous forest stands in Europe, for a total of 25 site-years, located on a gradient between southern France and northern Finland. Eddy-derived GPP varied threefold across the six sites, peak ecosystem leaf area index (LAI) (all-sided) varied from 4 to 22 m2 m–2 and mean annual temperature varied from –1 to 13 °C. A process-based model operating at a half-hourly time-step was parameterized with available information for each site, and explained 71–96% in variation between daily totals of GPP within site-years and 62% of annual total GPP across site-years. Using the parameterized model, we performed two simulation experiments: weather datasets were interchanged between sites, so that the model was used to predict GPP at some site using data from either a different year or a different site. The resulting bias in GPP prediction was related to several aggregated weather variables and was found to be closely related to the change in the effective temperature sum or mean annual temperature. High R2s resulted even when using weather datasets from unrelated sites, providing a cautionary note on the interpretation of R2 in model comparisons. A second experiment interchanged stand-structure information between sites, and the resulting bias was strongly related to the difference in LAI, or the difference in integrated absorbed light. Across the six sites, variation in mean annual temperature had more effect on simulated GPP than the variation in LAI, but both were important determinants of GPP. A sensitivity analysis of leaf physiology parameters showed that the quantum yield was the most influential parameter on annual GPP, followed by a parameter controlling the seasonality of photosynthesis and photosynthetic capacity. Overall, the results are promising for the development of a parsimonious model of GPP. [ABSTRACT FROM AUTHOR]

Current methods to study relations between stem respiration and stem growth have been hampered by problems in quantifying stem growth from dendrometer measurements, particularly on a daily time scale. This is mainly due to the water-related influences within these measurements that mask growth. A previously published model was used to remove water-related influences from measured radial stem variations to reveal a daily radial growth signal (ΔˆGm). We analysed the intra- and inter-annual relations between ΔˆGm and estimated growth respiration rates (Rg) on a daily scale for 5 years. Results showed that Rg was weakly correlated to stem growth prior to tracheid formation, but was significant during the early summer. In the late summer, the correlation decreased slightly relative to the early summer. A 1-day time lag was found of ΔˆGm preceding Rg. Using wavelet analysis and measurements from eddy covariance, it was found that Rg followed gross primary production and temperature with a 2 and 3 h time lag, respectively.This study shows that further in-depth analysis of in-situ growth and growth respiration dynamics is greatly needed, with a focus on cellular respiration at specific developmental stages, its woody tissue costs and linkages to source-sink processes and environmental drivers.

Background and Aims: Functional-structural plant models (FSPMs) allow simulation of tree crown development as the sum of modular (e.g. shoot-level) responses triggered by the local environmental conditions. The actual process of space filling by the crowns can be studied. Although the FSPM simulations are at organ scale, the data for their validation have usually been at more aggregated levels (whole-crown or whole-tree). Measurements made by terrestrial laser scanning (TLS) that have been segmented into elementary units (internodes) offer a phenotyping tool to validate the FSPM predictions at levels comparable with their detail. We demonstrate the testing of different formulations of crown development of Scots pine trees in the LIGNUM model using segmented TLS data., Methods: We made TLS measurements from four sample trees growing in a forest on a relatively poor soil from sapling size to mature stage. The TLS data were segmented into internodes. The segmentation also produced information on whether needles were present in the internode. We applied different formulations of crown development (flushing of buds and length of growth of new internodes) in LIGNUM. We optimized the parameter values of each formulation using genetic algorithms to observe the best fit of LIGNUM simulations to the measured trees. The fitness function in the estimation combined both tree-level characteristics (e.g. tree height and crown length) and measures of crown shape (e.g. spatial distribution of needle area)., Key Results: Comparison of different formulations against the data indicates that the Extended Borchert-Honda model for shoot elongation works best within LIGNUM. Control of growth by local density in the crown was important for all shoot elongation formulations. Modifying the number of lateral buds as a function of local density in the crown was the best way to accomplish density control., Conclusions: It was demonstrated how segmented TLS data can be used in the context of a shoot-based model to select model components.

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.)

Trees must simultaneously balance their CO2 uptake rate via stomata, photosynthesis, the transport rate of sugars and rate of sugar utilization in sinks while maintaining a favourable water and carbon balance. We demonstrate using a numerical model that it is possible to understand stomatal functioning from the viewpoint of maximizing the simultaneous photosynthetic production, phloem transport and sink sugar utilization rate under the limitation that the transpiration-driven hydrostatic pressure gradient sets for those processes. A key feature in our model is that non-stomatal limitations to photosynthesis increase with decreasing leaf water potential and/or increasing leaf sugar concentration and are thus coupled to stomatal conductance. Maximizing the photosynthetic production rate using a numerical steady-state model leads to stomatal behaviour that is able to reproduce the well-known trends of stomatal behaviour in response to, e.g., light, vapour concentration difference, ambient CO2 concentration, soil water status, sink strength and xylem and phloem hydraulic conductance. We show that our results for stomatal behaviour are very similar to the solutions given by the earlier models of stomatal conductance derived solely from gas exchange considerations. Our modelling results also demonstrate how the 'marginal cost of water' in the unified stomatal conductance model and the optimal stomatal model could be related to plant structural and physiological traits, most importantly, the soil-to-leaf hydraulic conductance and soil moisture., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We collected relevant observational and measured annual-resolution time series dealing with climate in northern Europe, focusing in Finland. We analysed these series for the reliability of their temperature signal at annual and seasonal resolutions. Importantly, we analysed all of the indicators within the same statistical framework, which allows for their meaningful comparison. In this framework, we employed a cross-validation procedure designed to reduce the adverse effects of estimation bias that may inflate the reliability of various temperature indicators, especially when several indicators are used in a multiple regression model. In our data sets, timing of phenological observations and ice break-up were connected with spring, tree ring characteristics (width, density, carbon isotopic composition) with summer and ice formation with autumn temperatures. Baltic Sea ice extent and the duration of ice cover in different watercourses were good indicators of winter temperatures. Using combinations of various temperature indicator series resulted in reliable temperature signals for each of the four seasons, as well as a reliable annual temperature signal. The results hence demonstrated that we can obtain reliable temperature information over different seasons, using a careful selection of indicators, combining the results with regression analysis, and by determining the reliability of the obtained indicator.

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.

Trees experience low apoplastic water potential frequently in most environments. Low apoplastic water potential increases the risk of embolism formation in xylem conduits and creates dehydration stress for the living cells. We studied the magnitude and rate of xylem diameter change in response to decreasing apoplastic water potential and the role of living parenchyma cells in it to better understand xylem diameter changes in different environmental conditions. We compared responses of control and heat-injured xylem of Pinus sylvestris (L.) and Populus tremula (L.) branches to decreasing apoplastic water potential created by osmotic stress, desiccation and freezing. It was shown that xylem in control branches shrank more in response to decreasing apoplastic water potential in comparison with the samples that were preheated to damage living xylem parenchyma. By manipulating the osmotic pressure of the xylem sap, we observed xylem shrinkage due to decreasing apoplastic water potential even in the absence of water tension within the conduits. These results indicate that decreasing apoplastic water potential led to withdrawal of intracellular water from the xylem parenchyma, causing tissue shrinkage. The amount of xylem shrinkage per decrease in apoplastic water potential was higher during osmotic stress or desiccation compared with freezing. During desiccation, xylem diameter shrinkage involved both dehydration-related shrinkage of xylem parenchyma and water tension-induced shrinkage of conduits, whereas dehydration-related shrinkage of xylem parenchyma was accompanied by swelling of apoplastic ice during freezing. It was also shown that the exchange of water between symplast and apoplast within xylem is clearly faster than previously reported between the phloem and the xylem. Time constant of xylem shrinkage was 40 and 2 times higher during osmotic stress than during freezing stress in P. sylvestris and P. tremula, respectively. Finally, it was concluded that the amount of water stored in the xylem parenchyma is an important reservoir for trees to buffer daily fluctuations in water relations., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

The pull of water from the soil to the leaves causes water in the transpiration stream to be under negative pressure decreasing the water potential below zero. The osmotic concentration also contributes to the decrease in leaf water potential but with much lesser extent. Thus, the surface tension force is approximately balanced by a force induced by negative water potential resulting in concavely curved water-air interfaces in leaves. The lowered water potential causes a reduction in the equilibrium water vapor pressure in internal (sub-stomatal/intercellular) cavities in relation to that over water with the potential of zero, i.e., over the flat surface. The curved surface causes a reduction also in the equilibrium vapor pressure of dissolved CO 2 , thus enhancing its physical solubility to water. Although the water vapor reduction is acknowledged by plant physiologists its consequences for water vapor exchange at low water potential values have received very little attention. Consequences of the enhanced CO 2 solubility to a leaf water-carbon budget have not been considered at all before this study. We use theoretical calculations and modeling to show how the reduction in the vapor pressures affects transpiration and carbon assimilation rates. Our results indicate that the reduction in vapor pressures of water and CO 2 could enhance plant water use efficiency up to about 10% at a leaf water potential of -2 MPa, and much more when water potential decreases further. The low water potential allows for a direct stomatal water vapor uptake from the ambient air even at sub-100% relative humidity values. This alone could explain the observed rates of foliar water uptake by e.g., the coastal redwood in the fog belt region of coastal California provided the stomata are sufficiently open. The omission of the reduction in the water vapor pressure causes a bias in the estimates of the stomatal conductance and leaf internal CO 2 concentration based on leaf gas exchange measurements. Manufactures of leaf gas exchange measurement systems should incorporate leaf water potentials in measurement set-ups.

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.)

Boreal trees experience repeated freeze-thaw cycles annually. While freezing has been extensively studied in trees, the dynamic responses occurring during the freezing and thawing remain poorly understood. At freezing and thawing, rapid changes take place in the water relations of living cells in needles and in stem. While freezing is mostly limited to extracellular spaces, living cells dehydrate, shrink and their osmotic concentration increases. We studied how the freezing-thawing dynamics reflected on leaf gas exchange, chlorophyll fluorescence and xylem and living bark diameter changes of Scots pine (Pinus sylvestris L.) saplings in controlled experiments. Photosynthetic rate quickly declined following ice nucleation and extracellular freezing in xylem and needles, almost parallel to a rapid shrinking of xylem diameter, while that of living bark followed with a slightly longer delay. While xylem and living bark diameters responded well to decreasing temperature and water potential of ice, the relationship was less consistent in the case of increasing temperature. Xylem showed strong temporal swelling at thawing suggesting water movement from bark. After thawing xylem diameter recovered to a pre-freezing level but living bark remained shrunk. We found that freezing affected photosynthesis at multiple levels. The distinct dynamics of photosynthetic rate and stomatal conductance reveals that the decreased photosynthetic rate reflects impaired dark reactions rather than stomatal closure. Freezing also inhibited the capacity of the light reactions to dissipate excess energy as heat, via non-photochemical quenching, whereas photochemical quenching of excitation energy decreased gradually with temperature in agreement with the gas exchange data., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Emissions of biogenic volatile organic compounds (BVOC) by boreal evergreen trees have strong seasonality, with low emission rates during photosynthetically inactive winter and increasing rates towards summer. Yet, the regulation of this seasonality remains unclear. We measured in situ monoterpene emissions from Scots pine shoots during several spring periods and analysed their dynamics in connection with the spring recovery of photosynthesis. We found high emission peaks caused by enhanced monoterpene synthesis consistently during every spring period (monoterpene emission bursts, MEB). The timing of the MEBs varied relatively little between the spring periods. The timing of the MEBs showed good agreement with the photosynthetic spring recovery, which was studied with simultaneous measurements of chlorophyll fluorescence, CO2 exchange and a simple, temperature history-based proxy for state of photosynthetic acclimation, S. We conclude that the MEBs were related to the early stages of photosynthetic recovery, when the efficiency of photosynthetic carbon reactions is still low whereas the light harvesting machinery actively absorbs light energy. This suggests that the MEBs may serve a protective functional role for the foliage during this critical transitory state and that these high emission peaks may contribute to atmospheric chemistry in the boreal forest in springtime., (© 2015 John Wiley & Sons Ltd.)

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., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Understanding the seasonality of photosynthesis in boreal evergreen trees and its control by the environment requires separation of the instantaneous and slow responses, as well as the dynamics of light reactions, carbon reactions, and respiration. We determined the seasonality of photosynthetic light response and respiration parameters of Scots pine (Pinus sylvestris L.) in the field in southern Finland and in controlled laboratory conditions. CO2 exchange and chlorophyll fluorescence were measured in the field using a continuously operated automated chamber setup and fluorescence monitoring systems. We also carried out monthly measurements of photosynthetic light, CO2 and temperature responses in standard conditions with a portable IRGA and fluorometer instrument. The field and response measurements indicated strong seasonal variability in the state of the photosynthetic machinery with a deep downregulation during winter. Despite the downregulation, the photosynthetic machinery retained a significant capacity during winter, which was not visible in the field measurements. Light-saturated photosynthesis (P sat) and the initial slope of the photosynthetic light response (α) obtained in standard conditions were up to 20% of their respective summertime values. Respiration also showed seasonal acclimation with peak values of respiration in standard temperature in spring and decline in autumn. Spring recovery of all photosynthetic parameters could be predicted with temperature history. On the other hand, the operating quantum yield of photosystem II and the initial slope of photosynthetic light response stayed almost at the summertime level until late autumn while at the same time P sat decreased following the prevailing temperature. Comparison of photosynthetic parameters with the environmental drivers suggests that light and minimum temperature are also decisive factors in the seasonal acclimation of photosynthesis in boreal evergreen trees.

Background and Aims: Woody plants can suffer from winter embolism as gas bubbles are formed in the water-conducting conduits when freezing occurs: gases are not soluble in ice, and the bubbles may expand and fill the conduits with air during thawing. A major assumption usually made in studies of winter embolism formation is that all of the gas dissolved in the xylem sap is trapped within the conduits and forms bubbles during freezing. The current study tested whether this assumption is actually valid, or whether efflux of gases from the stem during freezing reduces the occurrence of embolism., Methods: CO2 efflux measurements were conducted during freezing experiments for saplings of three Scots pine (Pinus sylvestris) and three Norway spruce (Picea abies) trees under laboratory conditions, and the magnitudes of the freezing-related bursts of CO2 released from the stems were analysed using a previously published mechanistic model of CO2 production, storage, diffusion and efflux from a tree stem. The freezing-related bursts of CO2 released from a mature Scots pine tree growing in field conditions were also measured and analysed., Key Results: Substantial freezing-related bursts of CO2 released from the stem were found to occur during both the laboratory experiments and under field conditions. In the laboratory, the fraction of CO2 released from the stem ranged between 27 and 96 % of the total CO2 content within the stem., Conclusions: All gases dissolved in the xylem sap are not trapped within the ice in the stem during freezing, as has previously been assumed, thus adding a new dimension to the understanding of winter embolism formation. The conduit water volume not only determines the volume of bubbles formed during freezing, but also the efficiency of gas efflux out of the conduit during the freezing process., (© The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Background and Aims: Tree models simulate productivity using general gas exchange responses and structural relationships, but they rarely check whether leaf gas exchange and resulting water and assimilate transport and driving pressure gradients remain within acceptable physical boundaries. This study presents an implementation of the cohesion-tension theory of xylem transport and the Münch hypothesis of phloem transport in a realistic 3-D tree structure and assesses the gas exchange and transport dynamics., Methods: A mechanistic model of xylem and phloem transport was used, together with a tested leaf assimilation and transpiration model in a realistic tree architecture to simulate leaf gas exchange and water and carbohydrate transport within an 8-year-old Scots pine tree. The model solved the dynamics of the amounts of water and sucrose solute in the xylem, cambium and phloem using a fine-grained mesh with a system of coupled ordinary differential equations., Key Results: The simulations predicted the observed patterns of pressure gradients and sugar concentration. Diurnal variation of environmental conditions influenced tree-level gradients in turgor pressure and sugar concentration, which are important drivers of carbon allocation. The results and between-shoot variation were sensitive to structural and functional parameters such as tree-level scaling of conduit size and phloem unloading., Conclusions: Linking whole-tree-level water and assimilate transport, gas exchange and sink activity opens a new avenue for plant studies, as features that are difficult to measure can be studied dynamically with the model. Tree-level responses to local and external conditions can be tested, thus making the approach described here a good test-bench for studies of whole-tree physiology.

A number of research groups in various areas of plant biology as well as computer science and applied mathematics have addressed modelling the spatiotemporal dynamics of growth and development of plants. This has resulted in development of functional-structural plant models (FSPMs). In FSPMs, the plant structure is always explicitly represented in terms of a network of elementary units. In this respect, FSPMs are different from more abstract models in which a simplified representation of the plant structure is frequently used (e.g. spatial density of leaves, total biomass, etc.). This key feature makes it possible to build modular models and creates avenues for efficient exchange of model components and experimental data. They are being used to deal with the complex 3-D structure of plants and to simulate growth and development occurring at spatial scales from cells to forest areas, and temporal scales from seconds to decades and many plant generations. The plant types studied also cover a broad spectrum, from algae to trees. This special issue of Annals of Botany features selected papers on FSPM topics such as models of morphological development, models of physical and biological processes, integrated models predicting dynamics of plants and plant communities, modelling platforms, methods for acquiring the 3-D structures of plants using automated measurements, and practical applications for agronomic purposes.

Background and Aims: Automatic acquisition of plant architecture is a major challenge for the construction of quantitative models of plant development. Recently, 3-D laser scanners have made it possible to acquire 3-D images representing a sampling of an object's surface. A number of specific methods have been proposed to reconstruct plausible branching structures from this new type of data, but critical questions remain regarding their suitability and accuracy before they can be fully exploited for use in biological applications., Methods: In this paper, an evaluation framework to assess the accuracy of tree reconstructions is presented. The use of this framework is illustrated on a selection of laser scans of trees. Scanned data were manipulated by experienced researchers to produce reference tree reconstructions against which comparisons could be made. The evaluation framework is given two tree structures and compares both their elements and their topological organization. Similar elements are identified based on geometric criteria using an optimization algorithm. The organization of these elements is then compared and their similarity quantified. From these analyses, two indices of geometrical and structural similarities are defined, and the automatic reconstructions can thus be compared with the reference structures in order to assess their accuracy., Key Results: The evaluation framework that was developed was successful at capturing the variation in similarities between two structures as different levels of noise were introduced. The framework was used to compare three different reconstruction methods taken from the literature, and allowed sensitive parameters of each one to be determined. The framework was also generalized for the evaluation of root reconstruction from 2-D images and demonstrated its sensitivity to higher architectural complexity of structure which was not detected with a global evaluation criterion., Conclusions: The evaluation framework presented quantifies geometric and structural similarities between two structures. It can be applied to the characterization and comparison of automatic reconstructions of plant structures from laser scanner data and 2-D images. As such, it can be used as a reference test for comparing and assessing reconstruction procedures.

• Attempts to combine biometric and eddy-covariance (EC) quantifications of carbon allocation to different storage pools in forests have been inconsistent and variably successful in the past. • We assessed above-ground biomass changes at five long-term EC forest stations based on tree-ring width and wood density measurements, together with multiple allometric models. Measurements were validated with site-specific biomass estimates and compared with the sum of monthly CO₂ fluxes between 1997 and 2009. • Biometric measurements and seasonal net ecosystem productivity (NEP) proved largely compatible and suggested that carbon sequestered between January and July is mainly used for volume increase, whereas that taken up between August and September supports a combination of cell wall thickening and storage. The inter-annual variability in above-ground woody carbon uptake was significantly linked with wood production at the sites, ranging between 110 and 370 g C m(-2) yr(-1) , thereby accounting for 10-25% of gross primary productivity (GPP), 15-32% of terrestrial ecosystem respiration (TER) and 25-80% of NEP. • The observed seasonal partitioning of carbon used to support different wood formation processes refines our knowledge on the dynamics and magnitude of carbon allocation in forests across the major European climatic zones. It may thus contribute, for example, to improved vegetation model parameterization and provides an enhanced framework to link tree-ring parameters with EC measurements., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Xylem and phloem need to maintain steady transport rates of water and carbohydrates to match the exchange rates of these compounds at the leaves. A major proportion of the carbon and nitrogen assimilated by a tree is allocated to the construction and maintenance of the xylem and phloem long distance transport tissues. This proportion can be expected to increase with increasing tree size due to the growing transport distances between the assimilating tissues, i.e., leaves and fine roots, at the expense of their growth. We formulated whole tree level scaling relations to estimate how xylem and phloem volume, nitrogen content and hydraulic conductance scale with tree size, and how these properties are distributed along a tree height. Xylem and phloem thicknesses and nitrogen contents were measured within varying positions in four tree species from Southern Finland. Phloem volume, nitrogen amount and hydraulic conductance were found to be concentrated toward the branch and stem apices, in contrast to the xylem where these properties were more concentrated toward the tree base. All of the species under study demonstrated very similar trends. Total nitrogen amount allocated to xylem and phloem was predicted to be comparable to the nitrogen amount allocated to the leaves in small and medium size trees, and to increase significantly above the nitrogen content of the leaves in larger trees. Total volume, hydraulic conductance and nitrogen content of the xylem were predicted to increase faster than that of the phloem with increasing tree height in small trees (<~10 m in height). In larger trees, xylem sapwood turnover to heartwood, if present, would maintain phloem conductance at the same level with xylem conductance with further increases in tree height. Further simulations with a previously published xylem-phloem transport model demonstrated that the Münch pressure flow hypothesis could explain phloem transport with increasing tree height even for the tallest trees.

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.

· Currently, phloem transport in plants under field conditions is not well understood. This is largely the result of the lack of techniques suitable for the measurement of the physiological properties of phloem. · We present a model that interprets the changes in xylem diameter and live bark thickness and separates the components responsible for such changes. We test the predictions from this model on data from three mature Scots pine trees in Finland. The model separates the live bark thickness variations caused by bark water capacitance from a residual signal interpreted to indicate the turgor changes in the bark. · The predictions from the model are consistent with processes related to phloem transport. At the diurnal scale, this signal is related to patterns of photosynthetic activity and phloem loading. At the seasonal scale, bark turgor showed rapid changes during two droughts and after two rainfall events, consistent with physiological predictions. Daily cumulative totals of this turgor term were related to daily cumulative totals of canopy photosynthesis. Finally, the model parameter representing radial hydraulic conductance between phloem and xylem showed a temperature dependence consistent with the temperature-driven changes in water viscosity. · We propose that this model has potential for the continuous field monitoring of tree phloem function., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

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.)

The photochemical reflectance index (PRI) is regarded as a promising proxy to track the dynamics of photosynthetic light use efficiency (LUE) via remote sensing. The implementation of this approach requires the relationship between PRI and LUE to scale not only in space but also in time. The short-term relationship between PRI and LUE is well known and is based on the regulative process of non-photochemical quenching (NPQ), but at the seasonal timescale the mechanisms behind the relationship remain unclear. We examined to what extent sustained forms of NPQ, photoinhibition of reaction centres, seasonal changes in leaf pigment concentrations, or adjustments in the capacity of alternative energy sinks affect the seasonal relationship between PRI and LUE during the year in needles of boreal Scots pine. PRI and NPQ were highly correlated during most of the year but decoupled in early spring when the foliage was deeply downregulated. This phenomenon was attributed to differences in the physiological mechanisms controlling the seasonal dynamics of PRI and NPQ. Seasonal adjustments in the pool size of the xanthophyll cycle pigments, on a chlorophyll basis, controlled the dynamics of PRI, whereas the xanthophyll de-epoxidation status and other xanthophyll-independent mechanisms controlled the dynamics of NPQ at the seasonal timescale. We conclude that the PRI leads to an underestimation of NPQ, and consequently overestimation of LUE, under conditions of severe stress in overwintering Scots pine, and most likely also in species experiencing severe drought. This severe stress-induced decoupling may challenge the implementation of the PRI approach.

Background and Aims: Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ε)., Methods: A functional-structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI(ind)) together with a genetic algorithm to find distributions of leaf angle (L(A)) and leaf photosynthetic capacity (A(max)) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A(max) either unconstrained or constrained to an upper value consistent with reported values for A(max) in A. saccharum., Key Results: It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ε were simultaneously maximized. Maximization of ε required simultaneous adjustments in L(A) and A(max) along gradients of PPFD in the plants. When A(max) was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ε because PPFD incident on leaves was higher than the PPFD at which ε(max) was attainable. Average leaf ε in constrained plants nonetheless improved with increasing LAI(ind) because of an increase in self-shading., Conclusions: It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ε at the scale of leaves, which requires a highly integrated response between L(A), A(max) and LAI(ind). The results also suggest that to maximize ε plants have evolved mechanisms that co-ordinate the L(A) and A(max) of individual leaves with PPFD availability.

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.)

Cambial growth was modelled as a function of detailed levelled physiological processes for cell enlargement and water and sugar transport to the cambium. Cambial growth was described at the cell level where local sugar concentration and turgor pressure induce irreversible cell expansion and cell wall synthesis. It was demonstrated how transpiration and photosynthesis rates, metabolic and physiological processes and structural features of a tree mediate their effects directly on the local water and sugar status and influence cambial growth. Large trees were predicted to be less sensitive to changes in the transient water and sugar status, compared with smaller ones, as they have more water and sugar storage and were, therefore, less coupled to short-term changes in the environment. Modelling the cambial dynamics at the individual cell level turned out to be a complex task as the radial short-distance transport of water and sugars and control signals determining cell division and cessation of cell enlargement and cell wall synthesis had to be described simultaneously.

* The variability of branch-level hydraulic properties was assessed across 12 Scots pine populations covering a wide range of environmental conditions, including some of the southernmost populations of the species. The aims were to relate this variability to differences in climate, and to study the potential tradeoffs between traits. * Traits measured included wood density, radial growth, xylem anatomy, sapwood- and leaf-specific hydraulic conductivity (K(S) and K(L)), vulnerability to embolism, leaf-to-sapwood area ratio (A(L) : A(S)), needle carbon isotope discrimination (Delta13C) and nitrogen content, and specific leaf area. * Between-population variability was high for most of the hydraulic traits studied, but it was directly associated with climate dryness (defined as a combination of atmospheric moisture demand and availability) only for A(L) : A(S), K(L) and Delta13C. Shoot radial growth and A(L) : A(S) declined with stand development, which is consistent with a strategy to avoid exceedingly low water potentials as tree size increases. In addition, we did not find evidence at the intraspecific level of some associations between hydraulic traits that have been commonly reported across species. * The adjustment of Scots pine's hydraulic system to local climatic conditions occurred primarily through modifications of A(L) : A(S) and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited.

We carried out a theoretical analysis of phloem transport based on Münch hypothesis by developing a coupled xylem-phloem transport model. Results showed that the maximum sugar transport rate of the phloem was limited by solution viscosity and that transport requirements were strongly affected by prevailing xylem water potential. The minimum number of xylem and phloem conduits required to sustain transpiration and assimilation, respectively, were calculated. At its maximum sugar transport rate, the phloem functioned with a high turgor pressure difference between the sugar sources and sinks but the turgor pressure difference was reduced if additional parallel conduits were added or solute relays were introduced. Solute relays were shown to decrease the number of parallel sieve tubes needed for phloem transport, leading to a more uniform turgor pressure and allowing faster information transmission within the phloem. Because xylem water potential affected both xylem and phloem transport, the conductance of the two systems was found to be coupled such that large structural investments in the xylem reduced the need for investment in the phloem and vice versa.

Gross primary production (GPP) is the primary source of all carbon fluxes in the ecosystem. Understanding variation in this flux is vital to understanding variation in the carbon sink of forest ecosystems, and this would serve as input to forest production models. Using GPP derived from eddy-covariance (EC) measurements, it is now possible to determine the most important factor to scale GPP across sites. We use long-term EC measurements for six coniferous forest stands in Europe, for a total of 25 site-years, located on a gradient between southern France and northern Finland. Eddy-derived GPP varied threefold across the six sites, peak ecosystem leaf area index (LAI) (all-sided) varied from 4 to 22 m(2) m(-2) and mean annual temperature varied from -1 to 13 degrees C. A process-based model operating at a half-hourly time-step was parameterized with available information for each site, and explained 71-96% in variation between daily totals of GPP within site-years and 62% of annual total GPP across site-years. Using the parameterized model, we performed two simulation experiments: weather datasets were interchanged between sites, so that the model was used to predict GPP at some site using data from either a different year or a different site. The resulting bias in GPP prediction was related to several aggregated weather variables and was found to be closely related to the change in the effective temperature sum or mean annual temperature. High R(2)s resulted even when using weather datasets from unrelated sites, providing a cautionary note on the interpretation of R(2) in model comparisons. A second experiment interchanged stand-structure information between sites, and the resulting bias was strongly related to the difference in LAI, or the difference in integrated absorbed light. Across the six sites, variation in mean annual temperature had more effect on simulated GPP than the variation in LAI, but both were important determinants of GPP. A sensitivity analysis of leaf physiology parameters showed that the quantum yield was the most influential parameter on annual GPP, followed by a parameter controlling the seasonality of photosynthesis and photosynthetic capacity. Overall, the results are promising for the development of a parsimonious model of GPP.

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.

Functional-structural plant growth models (FSPMs) combine the description of the structure of plants and the resource acquisition and partitioning at a detailed architectural level. They offer a means to study tree and stand development on the basis of a structurally accurate description that combines resource capture at the same level of detail. We describe here how a 'shoot-based' individual tree model, LIGNUM of Scots pine (Pinus sylvestris L.) has been applied to a group of identical trees (forest). The model has been applied to isolated trees and saplings growing in forest gaps. First, we present the LIGNUM model and the changes necessary for simulation of a forest instead of individual trees. LIGNUM derives tree growth on the basis of a process-based model of tree carbon balance and the architectural development of the 3-D tree crown. The time step is 1 year. We realised the forest as consisting of individual Scots pine trees on a plot 17 × 17 m, but simplified the stand description by simulating the growth of only one tree in the middle of the plot and assumed that the other trees were identical to it at all times. The model produced results that are comparable with observations made in real Scots pine trees and tree stands in Finland. The simulations with variable values of the parameters controlling the foliage-sapwood relationship, amount of sapwood required below a point in a branch or a stem, and the senescence of sapwood showed how growth declines when the sapwood requirement in the branches and stem was high. In this case, the proportion of resources allocated to the needles became small and the needle mass was low.

Acclimation of the partitioning of absorbed light energy in Photosystem II (PSII) between photochemical and non-photochemical processes includes short-term adjustments that are rapidly reversed in the dark and seasonal acclimation processes that are unaffected by dark acclimation. Thus, by using dark-acclimated leaves to study the seasonal acclimation of PSII, the confounding effect of short-term adjustments is eliminated. The maximum quantum yield of photochemistry, estimated by chlorophyll fluorescence analysis as F(v)/F(m), where F(v) = (F(m) - F(o)), and F(m) and F(o) are maximum and minimum chlorophyll fluorescence, respectively, has been widely used to follow the seasonal acclimation of PSII, because it is measured in dark-acclimated leaves. Seasonal changes in F(v)/F(m) can be caused by adjustments in either the photochemical capacity in PSII, or the capacity of thermal dissipation in PSII, or both. However, there is a lack of chlorophyll fluorescence parameters that can distinguish between these processes. In this study, we introduce two new parameters: the rate constants of sustained thermal energy dissipation (k(NPQ)) and of photochemistry (k(P)). We estimated k(NPQ) and k(P) from dark-acclimated F(o) and F(m) measured during spring recovery of photosynthesis in Scots pine (Pinus sylvestris L.) trees. We suggest that k(NPQ) and k(P) be used to study the mechanisms underlying the observed seasonal acclimation in PSII, because these parameters provide quantitative data that complement and extend F(v)/F(m) measurements.

Photosynthesis in evergreen conifers is characterized by down-regulation in autumn and rapid up-regulation in spring. This seasonal pattern is largely driven by temperature, but the light environment also plays a role. In overwintering Scots pine (Pinus sylvestris L.) trees, PSII is less down-regulated and recovers faster from winter stress in shaded needles than in needles exposed to full sunlight. Because the effect of light on the seasonal acclimation of PSII has not been quantitatively studied under field conditions, we used the rate constants for sustained thermal energy dissipation and photochemistry to investigate the dynamics and kinetics of the seasonal acclimation of PSII in needles exposed to different light environments. We monitored chlorophyll fluorescence and needle pigment concentration during the winter and spring in Scots pine seedlings growing in the field in different shading treatments, and within the crowns of mature trees. The results indicated that differences in acclimation of PSII in overwintering Scots pine among needles exposed to different light environments can be chiefly attributed to sustained thermal dissipation. We also present field evidence that zeaxanthin-facilitated thermal dissipation and aggregation of thylakoid membrane proteins are key mechanisms in the regulation of sustained thermal dissipation in Scots pine trees in the field.