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Mixed-species forests are, for multiple reasons, promising options for forest management in Central Europe. However, the extent to which interspecific competition affects tree hydrological processes is not clear. High-resolution dendrometers capture subdaily variations in stem diameter; they can simultaneously monitor stem growth (irreversible changes in diameter) and water status (reversible changes) of individual trees. Using the information on water status, we aimed to assess potential effects of tree species mixture, expressed as local neighborhood identity, on night-time rehydration and water stress. We deployed 112 sensors in pure and mixed forest stands of European beech, Norway spruce and Douglas fir on four sites in the northwestern Germany, measuring stem diameter in 10-min intervals for a period of four years (2019-2022). In a mixture distribution model, we used environmental variables, namely soil matric potential, atmospheric vapor pressure deficit, temperature, precipitation and neighborhood identity to explain night-time rehydration, measured as the daily minimum tree water deficit (TWDmin). TWDmin was used as a daily indicator of water stress and the daily occurrence of sufficient water supply, allowing for stem growth (potential growth). We found that species and neighborhood identity affected night-time rehydration, but the impacts varied depending on soil water availability. While there was no effect at high water availability, increasing drought revealed species-specific patterns. Beech improved night-time rehydration in mixture with Douglas fir, but not in mixture with spruce. Douglas fir, however, only improved rehydration at a smaller share of beech in the neighborhood, while beech dominance tended to reverse this effect. Spruce was adversely affected when mixed with beech. At species level and under dry conditions, we found that night-time rehydration was reduced in all species, but beech had a greater capacity to rehydrate under high to moderate soil water availability than the conifers, even under high atmospheric water demand. Our study gives new insights into neighborhood effects on tree water status and highlights the importance of species-specific characteristics for tree-water relations in mixed-species forests. It shows that drought stress of European beech can be reduced by admixing Douglas fir, which may point towards a strategy to adapt beech stands to climate change., (© The Author(s) 2024. Published by Oxford University Press.)

Climate change is raising concerns about how forests will respond to extreme droughts, heat waves and their co-occurrence. In this greenhouse study, we tested how carbon and water relations relate to seedling growth and mortality of northeastern US trees during and after extreme drought, warming, and combined drought and warming. We compared the response of our focal species red spruce (Picea rubens Sarg.) with a common associate (paper birch, Betula papyrifera Marsh.) and a species expected to increase abundance in this region with climate change (northern red oak, Quercus rubra L.). We tracked growth and mortality, photosynthesis and water use of 216 seedlings of these species through a treatment and a recovery year. Each red spruce seedling was planted in containers either alone or with another seedling to simulate potential competition, and the seedlings were exposed to combinations of drought (irrigated, 15-d 'short' or 30-d 'long') and temperature (ambient or 16 days at +3.5 °C daily maximum) treatments. We found dominant effects of the drought reducing photosynthesis, midday water potential, and growth of spruce and birch, but that oak showed considerable resistance to drought stress. The effects of planting seedlings together were moderate and likely due to competition for limited water. Despite high temperatures reducing photosynthesis for all species, the warming imposed in this study minorly impacted growth only for oak in the recovery year. Overall, we found that the diverse water-use strategies employed by the species in our study related to their growth and recovery following drought stress. This study provides physiological evidence to support the prediction that native species to this region like red spruce and paper birch are susceptible to future climate extremes that may favor other species like northern red oak, leading to potential impacts on tree community dynamics under climate change., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

The photosynthetic acclimation of boreal evergreen conifers is controlled by regulatory and photoprotective mechanisms that allow conifers to cope with extreme environmental changes. However, the underlying dynamics of photosystem II (PSII) and photosystem I (PSI) remain unresolved. Here, we investigated the dynamics of PSII and PSI during the spring recovery of photosynthesis in Pinus sylvestris and Picea abies using a combination of chlorophyll a fluorescence, P700 difference absorbance measurements, and quantification of key thylakoid protein abundances. In particular, we derived a new set of PSI quantum yield equations, correcting for the effects of PSI photoinhibition. Using the corrected equations, we found that the seasonal dynamics of PSII and PSI photochemical yields remained largely in balance, despite substantial seasonal changes in the stoichiometry of PSII and PSI core complexes driven by PSI photoinhibition. Similarly, the previously reported seasonal up-regulation of cyclic electron flow was no longer evident, after accounting for PSI photoinhibition. Overall, our results emphasize the importance of considering the dynamics of PSII and PSI to elucidate the seasonal acclimation of photosynthesis in overwintering evergreens. Beyond the scope of conifers, our corrected PSI quantum yields expand the toolkit for future studies aimed at elucidating the dynamic regulation of PSI., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Modeling and simulating the growth of the branching of tree species remains a challenge. With existing approaches, we can reconstruct or rebuild the branching architectures of real tree species, but the simulation of the growth process remains unresolved. First, we present a tree growth model to generate branching architectures that resemble real tree species. Secondly, we use a quantitative morphometric approach to infer the shape similarity of the generated simulations and real tree species. Within a functional-structural plant model, we implement a set of biological parameters that affect the branching architecture of trees. By modifying the parameter values, we aim to generate basic shapes of spruce, pine, oak and poplar. Tree shapes are compared using geometric morphometrics of landmarks that capture crown and stem outline shapes. Five biological parameters, namely xylem flow, shedding rate, proprioception, gravitysense and lightsense, most influenced the generated tree branching patterns. Adjusting these five parameters resulted in the different tree shapes of spruce, pine, oak, and poplar. The largest effect was attributed to gravity, as phenotypic responses to this effect resulted in different growth directions of gymnosperm and angiosperm branching architectures. Since we were able to obtain branching architectures that resemble real tree species by adjusting only a few biological parameters, our model is extendable to other tree species. Furthermore, the model will also allow the simulation of structural tree-environment interactions. Our simplifying approach to shape comparison between tree species, landmark geometric morphometrics, showed that even the crown-trunk outlines capture species differences based on their contrasting branching architectures., (© The Author(s) 2024. Published by Oxford University Press.)

To increase the resilience of forests to drought and other hazards, foresters are increasingly planting mixed stands. This requires knowledge about the drought response of tree species in pure and mixed-culture neighborhoods. In addition, drought frequently interacts with continued atmospheric nitrogen (N) deposition. To disentangle these factors for European beech, Norway spruce and Douglas fir, we conducted a replicated 3-factorial sapling growth experiment with three moisture levels, (high, medium, and low), two N levels (high and ambient), and pure and mixed-culture neighborhoods. We measured biomass, stomatal conductance (GS), shoot water potential (at predawn: ΨPD, midday, and turgor loss point: ΨTLP), branch xylem embolism resistance (Ψ50) and minimum epidermal conductance (Gmin). The three species differed most with respect to Gmin (10-fold higher in beech than in the conifers), hydroscape area (larger in beech), and the time elapsed to reach stomatal closure (TΨGS90) and ΨTLP (TTLP; shorter in beech), while Ψ50 and ΨTLP were remarkably similar. Neighborhood (pure vs mixed-culture) influenced biomass production, water status and hydraulic traits, notably GS (higher in Douglas fir, but lower in spruce and beech, in mixtures than pure culture), hydraulic safety margin (smaller for beech in mixtures), and TΨGS90 and TTLP (shorter for spruce in mixture). High N generally increased GS, but no consistent N effects on leaf water status and hydraulic traits were detected, suggesting that neighbor identity had a larger effect on plant water relations than N availability. We conclude that both tree neighborhood and N availability modulate the drought response of beech, spruce, and Douglas fir. Species mixing can alleviate the drought stress of some species, but often by disadvantaging other species. Thus, our study suggests that stabilizing and building resilience of production forests against a drier and warmer climate may depend primarily on the right species choice; species mixing can support the agenda., (© Crown copyright 2024.)

Beneficial and negative effects of species interactions can strongly influence water fluxes in forest ecosystems. However, little is known about how trees dynamically adjust their water use when growing with interspecific neighbours. Therefore, we investigated the interaction effects between Fagus sylvatica (European beech) and Picea abies (Norway spruce) on water-use strategies and aboveground structural characteristics. We used continuous in situ isotope spectroscopy of xylem and soil water to investigate source water dynamics and root water uptake depths. Picea abies exhibited a reduced sun-exposed crown area in equally mixed compared with spruce-dominated sites, which was further correlated to a reduction in sap flow of -14.5 ± 8.2%. Contrarily, F. sylvatica trees showed +13.3 ± 33.3% higher water fluxes in equally mixed compared with beech-dominated forest sites. Although a significantly higher crown interference by neighbouring trees was observed, no correlation of water fluxes and crown structure was found. High time-resolved xylem δ2H values showed a large plasticity of tree water use (-74.1 to -28.5‰), reflecting the δ2H dynamics of soil and especially precipitation water sources. Fagus sylvatica in equally mixed sites shifted water uptake to deeper soil layers, while uptake of fresh precipitation was faster in beech-dominated sites. Our continuous in situ water stable isotope measurements traced root water uptake dynamics at unprecedented temporal resolution, indicating highly dynamic use of water sources in response to precipitation and to neighbouring species competition. Understanding this plasticity may be highly relevant in the context of increasing water scarcity and precipitation variability under climate change., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

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., (© The Author(s) 2023. Published by Oxford University Press.)

Conifers growing in temperate forests utilize sustained forms of thermal dissipation during winter to protect the photosynthetic apparatus from damage, which can be monitored via pronounced reductions in photochemical efficiency (Fv/Fm) during winter. Eastern white pine (Pinus strobus L.) and white spruce (Picea  glauca (Moench) Voss) are known to recover from winter stress at different rates, with pine recovering more slowly than spruce, suggesting different mechanisms for sustained dissipation in these species. Our objectives were to monitor pine and spruce throughout spring recovery in order to provide insights into key mechanisms for sustained dissipation in both species. We measured chlorophyll fluorescence, pigments, and abundance and phosphorylation status of key photosynthetic proteins. We found that both species rely on two forms of sustained dissipation involving retention of high amounts of antheraxanthin (A) + zeaxanthin (Z), one that is very slowly reversible and temperature independent and one that is more dynamic and occurs only on subzero days. Differences in protein abundance suggest that spruce, but not pine, likely upregulates cyclic or alternative pathways of electron transport involving the cytochrome b6f complex and photosystem I (PSI). Both species show an increased sustained phosphorylation of the D1 protein on subzero days, and spruce additionally shows dramatic increases in the sustained phosphorylation of light-harvesting complex II (LHCII) and other PSII core proteins on subzero days only, suggesting that a mechanism of sustained dissipation that is temperature dependent requires sustained phosphorylation of photosynthetic proteins in spruce, possibly allowing for direct energy transfer from PSII to PSI as a mechanism of photoprotection. The data suggest differences in strategy among conifers in mechanisms of sustained thermal dissipation in response to winter stress. Additionally, the flexible induction of sustained A + Z and phosphorylation of photosynthetic proteins in response to subzero temperatures during spring recovery seem to be important in providing photoprotection during transitional periods with high temperature fluctuation., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Abiotic factors such as water and nutrient availability can exert a dominant influence on the susceptibility of plants to various pathogens. Effects of abiotic environmental factors on phenolic compound concentrations in the plant tissue may represent one of the major underlying mechanisms, as these compounds are known to play a substantial role in plant resistance to pests. In particular, this applies to conifer trees, in which a large range of phenolic compounds are produced constitutively and/or induced by pathogen attack. We subjected Norway spruce saplings to water limitation and elevated nutrient supply over 2 years and subsequently controlled infection with the needle rust Chrysomyxa rhododendri (DC.) de Bary and analysed both constitutive and inducible phenolic compound concentrations in the needles as well as the degree of infection. Compared with the control group, both drought and fertilization profoundly modified the constitutive and pathogen-induced profiles of phenolic compounds, but had little impact on the total phenolic content. Fertilization predominantly affected the inducible phenolic response and led to higher infection rates by C. rhododendri. Drought stress, in contrast, mainly shaped the phenolic profiles in healthy plant parts and had no consequences on the plant susceptibility. The results show that specific abiotic effects on individual compounds seem to be decisive for the infection success of C. rhododendri, whereby the impaired induced response in saplings subjected to nutrient supplementation was most critical. Although drought effects were minor, they varied depending on the time and length of water limitation. The results indicate that prolonged drought periods in the future may not significantly alter the foliar defence of Norway spruce against C. rhododendri, but fertilization, often propagated to increase tree growth and forest productivity, can be counterproductive in areas with high pathogen pressure., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

It is assumed that the stimulatory effects of elevated CO2 concentration ([CO2]) on photosynthesis and growth may be substantially reduced by co-occurring environmental factors and the length of CO2 treatment. Here, we present the study exploring the interactive effects of three manipulated factors ([CO2], nitrogen supply and water availability) on physiological (gas-exchange and chlorophyll fluorescence), morphological and stoichiometric traits of Norway spruce (Picea abies) saplings after 2 and 3 years of the treatment under natural field conditions. Such multifactorial studies, going beyond two-way interactions, have received only limited attention until now. Our findings imply a significant reduction of [CO2]-enhanced rate of CO2 assimilation under reduced water availability which deepens with the severity of water depletion. Similarly, insufficient nitrogen availability leads to a down-regulation of photosynthesis under elevated [CO2] being particularly associated with reduced carboxylation efficiency of the Rubisco enzyme. Such adjustments in the photosynthesis machinery result in the stimulation of water-use efficiency under elevated [CO2] only when it is combined with a high nitrogen supply and reduced water availability. These findings indicate limited effects of elevated [CO2] on carbon uptake in temperate coniferous forests when combined with naturally low nitrogen availability and intensifying droughts during the summer periods. Such interactions have to be incorporated into the mechanistic models predicting changes in terrestrial carbon sequestration and forest growth in the future., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (gs) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (udaily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, udaily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and gs recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and gs. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of udaily at the crown base compared with DBH level upon watering. However, udaily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g., udaily, ΨPD and gs, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Norway spruce (Picea abies (L.) Karst) trees are affected by ongoing climate change, including warming and exposure to phytotoxic levels of ozone. Non-volatile terpenoids and volatile terpenoids (biogenic organic volatile compounds, BVOCs) protect spruce against biotic and abiotic stresses. BVOCs also affect the atmosphere's oxidative capacity. Four-year-old Norway spruce were exposed to elevated ozone (EO) (1.4 × ambient) and warming (1.1 °C + ambient air) alone and in combination on an open-field exposure site in Central Finland. Net photosynthesis, needle terpenoid concentrations and BVOC emissions were measured four times during the experiment's second growing season: after bud opening in May, during the mid-growing season in June, and after needle maturation in August and September. Warming increased terpene concentrations in May due to advanced phenology and decreased them at the end of the growing season in matured current-year needles. Ozone enhanced these effects of warming on several compounds. Warming decreased concentrations of oxygenated sesquiterpenes in previous-year needles. Decreased emissions of oxygenated monoterpenes by warming and ozone alone in May were less prominent when ozone and warming were combined. A similar interactive treatment response in isoprene, camphene, tricyclene and α-pinene was observed in August when the temperature and ozone concentration was high. The results suggest long-term warming may reduce the terpenoid-based defence capacity of young spruce, but the defence capacity can be increased during the most sensitive growth phase (after bud break), and when high temperatures or ozone concentrations co-occur. Reduced BVOC emissions from young spruce may decrease the atmosphere's oxidative capacity in the warmer future, but the effect of EO may be marginal because less reactive minor compounds are affected., (© The Author(s) 2022. Published by Oxford University Press.)

Recent climate projections predict a more rapid increase of winter temperature than summer and global temperature averages in temperate and cold environments. As there is relatively little experimental knowledge on the effect of winter warming on cambium phenology and stem growth in species growing in cold environments, the setting of manipulative experiments is considered of primary importance, and they can help to decipher the effect of reduced winter chilling and increased forcing temperatures on cambium reactivation, growth and xylem traits. In this study, localized stem heating was applied to investigate the effect of warming from the rest to the growth phase on cambium phenology, intra-annual stem growth dynamics and ring wood features in Picea abies (L.) H.Karst. We hypothesized that reduced winter chilling induces a postponed cambium dormancy release and decrease of stem growth, while high temperature during cell wall lignification determines an enrichment of latewood-like cells. The heating device was designed to maintain a +5 °C temperature delta with respect to air temperature, thus allowing an authentic scenario of warming. Continuous stem heating from the rest (November) to the growing phase determined, at the beginning of radial growth, a reduction of the number of cell layers in the cambium, higher number of cell layers in the wall thickening phase and an asynchronous stem radial growth when comparing heated and ambient saplings. Nevertheless, heating did not induce changes in the number of produced cell layers at the end of the growing season. The analyses of two-photon fluorescence images showed that woody rings formed during heating were enriched with latewood-like cells. Our results showed that an increase of 5 °C of temperature applied to the stem from the rest to growth might not influence, as generally reported, onset of cambial activity, but it could affect xylem morphology of Norway spruce in mountain environments., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Increasing seawater exposure is causing mortality of coastal forests, yet the physiological response associated with seawater-induced tree mortality, particularly in non-halophytes, is poorly understood. We investigated the shifts in carbon and nitrogen (N) metabolism of mature Sitka-spruce trees that were dying after an ecosystem-scale manipulation of tidal seawater exposure. Soil porewater salinity and foliar ion concentrations increased after seawater exposure and were strongly correlated with the percentage of live foliated crown (PLFC; e.g., crown 'greenness', a measure of progression to death). Co-occurring with decreasing PLFC was decreasing photosynthetic capacity, N-investment into photosynthesis, N-resorption efficiency and non-structural carbohydrate (soluble sugars and starch) concentrations, with the starch reserves depleted to near zero when PLFC dropped below 5%. Combined with declining PLFC, these changes subsequently decreased total carbon gain and thus exacerbated the carbon starvation process. This study suggests that an impairment in carbon and N metabolism during the mortality process after seawater exposure is associated with the process of carbon starvation, and provides critical knowledge necessary to predict sea-level rise impacts on coastal forests., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

This study presents results from continuous measurements of stem CO2 efflux carried out for seven growing seasons in a young Norway spruce forest. The objective of the study was to determine differences in temperature sensitivity of stem CO2 efflux (Q10) during night (when sap flow is zero or nearly zero), during early afternoon (when the maximum rate of sap flow occurs) and during two transition periods between the aforementioned periods. The highest Q10 was recorded during the period of zero sap flow, while the lowest Q10 was observed in period of the highest sap flow. Calculating Q10 using only data from the period of zero sap flow resulted in a Q10 that was higher by as much as 19% compared with Q10 calculated using 24 h data. On the other hand, basing the calculation on data from the period of the highest sap flow yielded 5.6% lower Q10 than if 24 h data were used. Considering that change in CO2 efflux lagged in time behind changing stem temperature, there was only a small effect on calculated Q10 for periods with zero and the highest sap flow. A larger effect of the time lag (by as much as 15%) was observed for the two transition periods. Stem CO2 efflux was modelled based on the night CO2 efflux response to temperature. This model had a tendency to overestimate CO2 efflux during daytime, thus indicating potential daytime depression of stem CO2 efflux compared with the values predicated on the basis of temperature caused by CO2 transport upward in the sap flow. This view was supported by our results inasmuch as the overestimation grew with sap flow that was modelled on the basis of photosynthetically active radiation and vapour pressure deficit., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Winter has long been considered a dormant season in boreal forests regarding plant physiological activity such as nutrient acquisition. However, biogeochemical data clearly show that soil can remain unfrozen with substantial rates of nutrient transformation for several weeks following autumn snowfall. Here we examined nitrate (NO3--N) assimilation by black spruce (Picea mariana (Mill.) Britton, Sterns and Poggenb.) during summer and winter in Interior Alaska to test our hypothesis that this boreal species is able to assimilate NO3--N, even at the very low temperatures typical of early winter. Nitrate reductase activity (NRA) was measured in current year needles and fine roots of black spruce as an indicator of NO3--N assimilation in the summer and winter at two boreal forest sites. Nitrate concentration in the needles and roots were also measured to determine whether NO3--N was available in plant tissue for the enzyme. Nitrate reductase activity and NO3--N were detected in needles and roots in the winter as well as the summer. The results of a generalized linear mixed model showed that season had minimal effects on NRA and NO3--N concentration in this species. Additionally, the effect of incubation temperature for the NRA assays was tested at 30 °C and -3 °C for samples collected in the winter. Substantial enzyme activity was detected in winter-collected samples, even in incubations conducted at -3 °C. These results indicate that this dominant tree species in the boreal forests of Interior Alaska, black spruce, has the capacity to assimilate NO3--N below freezing temperatures, suggesting that the physiological activity required for nitrogen (N) resource acquisition may extend beyond the typical growing season. Our findings coupled to biogeochemical evidence for high microbial activity under the snow also indicate that winter N acquisition should be taken into account when estimating the annual N budgets of boreal forest ecosystems., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Synchrony between host budburst and insect emergence greatly influences the time window for insect development and survival. A few alterations of bud phenology have been reported under defoliation without clear consensus regarding the direction of effects, i.e., advance or delay. Here, we compared budburst phenology between conifers in defoliation and control treatments, and measured carbon allocation as a potential mechanistic explanation of changes in phenology. In a 2-year greenhouse experiment, saplings of balsam fir, black spruce and white spruce of two different provenances (north and south) were subjected to either control (no larvae) or natural defoliation treatment (larvae added) by spruce budworm. Bud and instar phenology, primary and secondary growth, defoliation and non-structural carbohydrates were studied during the growing season. No differences were observed in bud phenology during the first year of defoliation. After 1 year of defoliation, bud phenology advanced by 6-7 days in black spruce and balsam fir and by 3.5 days in white spruce compared with the control. Because of this earlier bud break, apical and shoot growth exceeded 50% of its final length before mature instar defoliation occurred, which decreased the overall level of damage. A sugar-mediated response, via earlier starch breakdown, and higher sugar availability to buds explains the advanced budburst in defoliated saplings. The advanced phenological response to defoliation was consistent across the conifer species and provenances except for one species × provenance combination. Allocation of carbon to buds and shoots growth at the expense of wood growth in the stem and reserve accumulation represents a shift in the physiological resources priorities to ensure tree survival. This advancement in bud phenology could be considered as a physiological response to defoliation based on carbohydrate needs for primary growth, rather than a resistance trait to spruce budworm., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We quantified seasonal CO2 assimilation capacities for seven dominant vascular species in a wet boreal forest peatland then applied data to a land surface model parametrized to the site (ELM-SPRUCE) to test if seasonality in photosynthetic parameters results in differences in simulated plant responses to elevated CO2 and temperature. We collected seasonal leaf-level gas exchange, nutrient content and stand allometric data from the field-layer community (i.e., Maianthemum trifolium (L.) Sloboda), understory shrubs (Rhododendron groenlandicum (Oeder) Kron and Judd, Chamaedaphne calyculata (L.) Moench., Kalmia polifolia Wangenh. and Vaccinium angustifolium Alton.) and overstory trees (Picea mariana (Mill.) B.S.P. and Larix laricina (Du Roi) K. Koch). We found significant interspecific seasonal differences in specific leaf area, nitrogen content (by area; Na) and photosynthetic parameters (i.e., maximum rates of Rubisco carboxylation (Vcmax25°C), electron transport (Jmax25°C) and dark respiration (Rd25°C)), but minimal correlation between foliar Na and Vcmax25°C, Jmax25°C or Rd25°C, which illustrates that nitrogen alone is not a good correlate for physiological processes such as Rubisco activity that can change seasonally in this system. ELM-SPRUCE was sensitive to the introduction of observed interspecific seasonality in Vcmax25°C, Jmax25°C and Rd25°C, leading to simulated enhancement of net primary production (NPP) using seasonally dynamic parameters as compared with use of static parameters. This pattern was particularly pronounced under simulations with higher temperature and elevated CO2, suggesting a key hypothesis to address with future empirical or observational studies as climate changes. Inclusion of species-specific seasonal photosynthetic parameters should improve estimates of boreal ecosystem-level NPP, especially if impacts of seasonal physiological ontogeny can be separated from seasonal thermal acclimation., (Published by Oxford University Press 2019.)

This study presents results from continuous measurements of stem CO2 efflux carried out for seven experimental seasons (from May to October) in a young Norway spruce forest. The objectives of the study were to determine variability in the response of stem CO2 efflux to stem temperature over the season and to observe differences in the calculated relationship between stem temperature and CO2 efflux based on full growing season data or on data divided into periods according to stem growth rate. Temperature sensitivity of stem CO2 efflux (Q10) calculated for the established periods ranged between 1.61 and 3.46 and varied over the season, with the lowest values occurring in July and August. Q10 calculated using data from the full growing seasons ranged between 2.30 and 2.94 and was often significantly higher than Q10 calculated for the individual periods. Temperature-normalized stem CO2 efflux (R10) determined using Q10 from growing season data was overestimated when the temperature was below 10 °C and underestimated when the temperature was above 10 °C, compared with R10 calculated using Q10 established for the individual periods. The differences in daily mean R10 calculated by these two approaches ranged between -0.9 and 0.2 μmol CO2 m-2 s-1. The results of this study confirm that long periods for determining the temperature dependence of stem CO2 efflux encompass different statuses of the wood (especially in relation to stem growth). This may cause bias in models using this relationship for estimating stem CO2 efflux as a function of temperature.

As a tree grows taller, the increase in gravitational pressure and path length resistance results in lower water potentials at a given flow rate and higher carbon construction costs to transport a given amount of water to the leaves. We investigated how hydraulic safety and efficiency are coordinated under the constraints of higher cavitation risks and higher carbon construction costs with increasing tree height. We combined measurements of xylem tracheid anatomical traits with the vulnerability to drought-induced embolism and hydraulic conductivity of the apical shoots of 2- to 37-m tall Picea abies trees growing at two sites in the Dolomites (Italian Eastern Alps). We found that the theoretical hydraulic conductivity of the apical shoots increased with tree height at both sites (P < 0.001) as a result of an increase in either total tracheid number or mean hydraulic diameter. The xylem water potential inducing 50% loss of apical conductance significantly increased from small (-4.45 ± 0.20 MPa) to tall trees (-3.65 ± 0.03 MPa) (P = 0.007). The more conductive xylem at the treetop of taller trees allows the full compensation for the height-related hydraulic constraints and minimizes the additional carbon costs of transporting water over a longer path length. The corresponding increase in vulnerability to cavitation shows that hydraulic efficiency is prioritized over safety during height growth.

Water stress has been identified as a key mechanism of the contemporary increase in tree mortality rates in northwestern North America. However, a detailed analysis of boreal tree hydrodynamics and their interspecific differences is still lacking. Here we examine the hydraulic behaviour of co-occurring larch (Larix laricina) and black spruce (Picea mariana), two characteristic boreal tree species, near the southern limit of the boreal ecozone in central Canada. Sap flux density (Js), concurrently recorded stem radius fluctuations and meteorological conditions are used to quantify tree hydraulic functioning and to scrutinize tree water-use strategies. Our analysis revealed asynchrony in the diel hydrodynamics of the two species with the initial rise in Js occurring 2 h earlier in larch than in black spruce. Interspecific differences in larch and black spruce crown architecture explained the observed asynchrony in their hydraulic functioning. Furthermore, the two species exhibited diverging stomatal regulation strategies with larch and black spruce employing relatively isohydric and anisohydric behaviour, respectively. Such asynchronous and diverging tree-level hydrodynamics provide new insights into the ecosystem-level complementarity in tree form and function, with implications for understanding boreal forests' water and carbon dynamics and their resilience to environmental stress.

Background and Aims: Form and function relationships in plant reproductive structures have long fascinated biologists. Although the intricate associations between specific pollinators and reproductive morphology have been widely explored among animal-pollinated plants, the evolutionary processes underlying the diverse morphologies of wind-pollinated plants remain less well understood. Here we study how this diversity may have arisen by focusing on two conifer species in the pine family that have divergent reproductive cone morphologies at pollination., Methods: Standard histology methods, artificial wind pollination assays and phylogenetic analyses were used in this study., Key Results: A detailed study of cone ontogeny in these species reveals that variation in the rate at which their cone scales mature means that pollination occurs at different stages in their development, and thus in association with different specific morphologies. Pollination experiments nevertheless indicate that both species effectively capture pollen., Conclusions: In wind-pollinated plants, morphological diversity may result from simple variation in development among lineages rather than selective pressures for any major differences in function or performance. This work also illustrates the broader importance of developmental context in understanding plant form and function relationships; because plant reproductive structures perform many different functions over their lifetime, subtle differences in development may dramatically alter the specific morphologies that they use to meet these demands.

The dormancy and the growth of trees in temperate climates are synchronized with seasons. Preparation for dormancy and its proper progression are key for survival and development in the next season. Using a unique approach that combined microscopy and proteomic methods, we investigated changes in Norway spruce (Picea abies (L.) H. Karst.) embryonic shoots during four distinct stages of dormancy in natural weather conditions. We identified 13 proteins that varied among dormancy stages, and were linked to regulation of protein level; functioning of chloroplasts and other plastids; DNA and RNA regulation; and oxidative stress. We also found a group of five proteins, related to cold hardiness, that did not differ in expression among stages of dormancy, but had the highest abundancy level. Ultrastructure of organelles is tightly linked to their metabolic activity, and hence may indicate dormancy status. The observed ultrastructure during endodormancy was stable, whereas during ecodormancy, the structural changes were dynamic and related mainly to nucleus, plastids and mitochondria. At the ultrastructural level, the lack of starch and the presence of callose in plasmodesmata in all regions of embryonic shoot were indicators of full endodormancy. At the initiation of ecodormancy, we noted an increase in metabolic activity of organelles, tissue-specific starch hyperaccumulation and degradation. However, in proteomic analysis, we did not find variation in expression of proteins related to starch degradation or to symplastic isolation of cells. The combination of ultrastructural and proteomic methods gave a more complete picture of vegetative bud dormancy than either of them applied separately. We found some changes at the structural level, but not their analogues in the proteome. Our study suggests a very important role of plastids' organization and metabolism, and their protection in the course of dormancy and during the shift from endo- to ecodormancy and the acquisition of growth competence.

Bud primordia of Picea abies (L.) H. Karst. remain ice free at subzero temperatures by supercooling. Once ice forms inside the primordium, it is immediately injured. Supercooling capacity increases seasonally from ~-5 °C to as much as -50 °C by currently unknown mechanisms. Among other prerequisites, dehydration of tissues over the winter months has been considered to play a key role in freezing tolerance. In this regard, the water content of bud primordia may be crucial, especially in reference to supercooling. In order to assess the role of dehydration in supercooling capacity, seasonal changes in supercooling capacity and the water potential of bud primordia of Picea abies (L.) H. Karst were measured at two sites that differed by 1298 m in elevation, after artificial frost hardening and dehardening treatments and after controlled bench drying. The extent of supercooling of bud primordia varied from -7 °C in summer to -24.6 °C in winter, a difference of 17.6 -19.3 K. Total actual water potential (Ψtact) of bud primordia was -2 MPa in summer and decreased to a mean of -3.8 MPa in midwinter. The decline involved dehydration, and to a lesser extent, osmoregulation. At decreased Ψtact values (<3.0 MPa), supercooling capacity significantly increased <-19.5 °C, however, the correlation between actual water potential and supercooling capacity was poor. Frost-hardening treatments increased the supercooling capacity of bud primordia (-0.6 K day-1) and lowered Ψtact (-0.2 MPa day-1). Frost-dehardening treatments reduced supercooling capacity (+1.1 K day-1), and at the same time, increased Ψtact (+0.3 MPa day-1). In contrast, artificial drying of bud primordia in the range observed seasonally (-2.0 MPa) had no effect on supercooling capacity. These results suggest that there is no causal relationship between desiccation and the supercooling capacity of bud primordia in P. abies, but rather it involves other compounds within the cells of the bud primordium that reduce the water potential.

Norway spruce ( Picea abies) is by far the most important timber species in Europe, but its outstanding role in future forests is jeopardized by its high sensitivity to drought. We analyzed drought response of Norway spruce at the warmest fringe of its natural range. Based on a 35-year old provenance experiment we tested for genetic variation among and within seed provenances across consecutively occurring strong drought events using dendroclimatic time series. Moreover, we tested for associations between ≈1,700 variable SNPs and traits related to drought response, wood characteristics and climate-growth relationships. We found significant adaptive genetic variation among provenances originating from the species' Alpine, Central and Southeastern European range. Genetic variation between individuals varied significantly among provenances explaining up to 44% of the phenotypic variation in drought response. Varying phenotypic correlations between drought response and wood traits confirmed differences in selection intensity among seed provenances. Significant associations were found between 29 SNPs and traits related to drought, climate-growth relationships and wood properties which explained between 11 and 43% of trait variation, though 12 of them were due to single individuals having extreme phenotypes of the respective trait. The majority of these SNPs are located within exons of genes and the most important ones are preferentially expressed in cambium and xylem expansion layers. Phenotype-genotype associations were stronger if only provenances with significant quantitative genetic variation in drought response were considered. The present study confirms the high adaptive variation of Norway spruce in Central and Southeastern Europe and demonstrates how quantitative genetic, dendroclimatic and genomic data can be linked to understand the genetic basis of adaptation to climate extremes in trees., (Copyright © 2018 Trujillo-Moya et al.)

Background and Aims: Knowledge of thermal acclimation of physiological processes of boreal tree species is necessary to determine their ability to adapt to predicted global warming and reduce the uncertainty around the anticipated feedbacks of forest ecosystems and global carbon cycle to climate change. The objective of this work was to examine the extent of thermal acclimation of net photosynthesis (An) and dark respiration (Rd) of two distant white spruce (Picea glauca) seed sources (from south and north of the commerial forest zone in Québec) in response to latitudinal and seasonal variations in growing conditions., Methods: The temperature responses of An, its biochemical and biophysical limitations, and Rd were measured in 1-year-old needles of seedlings from the seed sources growing in eight forest plantations along a regional thermal gradient of 5.5 °C in Québec, Canada., Key Results: The average optimum temperature (Topt) for An was 19 ± 1.2 °C and was similar among seed sources and plantation sites along the thermal gradient. Net photosynthesis at Topt (Aopt) varied significantly among plantation sites and was quadratically related to the mean July temperature (MJT) of plantation sites. Topt for mesophyll conductance, maximum electron transport rate and maximum rate of carboxylation were 28, 22 and 30 °C, respectively. Basal respiration rate (Rd at 10 °C) was linearly and negatively associated with MJT. Q10 of Rd (the rate of change in Rd with a 10 °C increase in temperature) did not show any significant relationship with MJT and averaged 1.5 ± 0.1. The two seed sources were similar in their thermal responses to latitudinal and seasonal variations in growing conditions., Conclusions: The results showed moderate thermal acclimation of respiration and no evidence for thermal acclimation of photosynthesis or local genetic adaptation for traits related to thermal acclimation. Therefore, growth of local white spruces may decline in future climates., (© The Author(s) 2017. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Temperatures are expected to increase over the next century in all terrestrial biomes and particularly in boreal forests, where drought-induced mortality has been predicted to rise. Genomics research is helping to develop hypotheses regarding the molecular basis of drought tolerance and recent work proposed that the osmo-protecting dehydrin proteins have undergone a clade-specific expansion in the Pinaceae, a major group of conifer trees. The objectives of this study were to identify all of the putative members of the gene family, trace their evolutionary origin, examine their structural diversity and test for drought-responsive expression. We identified 41 complete dehydrin coding sequences in Picea glauca, which is four times more than most angiosperms studied to date, and more than in pines. Phylogenetic reconstructions indicated that the family has undergone an expansion in conifers, with parallel evolution implicating the sporadic resurgence of certain amino acid sequence motifs, and a major duplication giving rise to a clade specific to the Pinaceae. A variety of plant dehydrin structures were identified with variable numbers of the A-, E-, S- and K-segments and an N-terminal (N1) amino acid motif including assemblages specific to conifers. The expression of several of the spruce dehydrins was tissue preferential under non-stressful conditions or responded to water stress after 7-18 days without watering, reflecting changes in osmotic potential. We found that dehydrins with N1 K2 and N1 AESK2 sequences were the most responsive to the lack of water. Together, the family expansion, drought-responsive expression and structural diversification involving loss and gain of amino acid motifs suggests that subfunctionalization has driven the diversification seen among dehydrin gene duplicates. Our findings clearly indicate that dehydrins represent a large family of candidate genes for drought tolerance in spruces and in other Pinaceae that may underpin adaptability in spatially and temporally variable environments.

Tetropium fuscum (Fabricius) (Coleoptera: Cerambycidae), a phloem-feeding and wood-boring beetle introduced from Eurasia, attacks spruce in eastern Canada alongside its native congener Tetropium cinnamopterum Kirby. We reared phloem- and wood-feeding insects (and their predators) from bolts of red and Norway spruce (Picea rubens and Picea abies) in Nova Scotia, comparing insect communities between bolts with added eggs of T. fuscum or T. cinnamopterum and bolts without added Tetropium (controls). We tested for impacts of each Tetropium on insect community structure (Simpson's diversity, richness, and evenness). We also asked whether, consistent with Darwin's Naturalization Hypothesis, Tetropium spp. would have greater impacts on emergence of its closer relatives (which might be most likely to compete and/or share natural enemies). Addition of Tetropium eggs (either species) to bolts lowered insect diversity in both host trees. Both richness and evenness components of diversity were always lower in +Tetropium treatments, although different components reached statistical significance in different Tetropium species × host combinations. Addition of Tetropium spp. significantly reduced emergence of some species: Evodinus monticola (Randall) (Coleoptera: Cerambycidae) was reduced by T. fuscum on both hosts and by T. cinnamopterum on Norway spruce; Hylobius congener Dalla Torre, Schenkling, and Marshall was reduced by T. fuscum on red spruce; and Xylophagus sp. (Diptera: Xylophagidae) was reduced by T. cinnamopterum on Norway spruce. However, there was no relationship between Tetropium's impact on a community member and their phylogenetic relatedness, and the overall impacts of Tetropium presence were not very different between T. fuscum and T. cinnamopterum., (© The Author(s) 2017. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Decreasing water availability due to climate change poses the question of whether and to what extent tree species are able to hydraulically acclimate and how hydraulic traits of stems and leaves are coordinated under drought. In a through-fall exclusion experiment, hydraulic acclimation was analyzed in a mixed forest stand of Fagus sylvatica L. and Picea abies (L.) Karst. In drought-stressed (TE, through-fall exclusion over 2 years) and control (CO) trees, hydraulic vulnerability was studied in branches as well as in leaves (F. sylvatica) and end-twigs (P. abies, entirely formed during the drought period) sampled at the same height in sun-exposed portions of the tree crown. In addition, relevant xylem anatomical traits and leaf pressure-volume relations were analyzed. The TE trees reached pre-dawn water potentials down to -1.6 MPa. In both species, water potentials at 50% loss of xylem hydraulic conductivity were ~0.4 MPa more negative in TE than in CO branches. Foliage hydraulic vulnerability (expressed as water potential at 50% loss of leaf/end-twig hydraulic conductance) and water potential at turgor loss point were also, respectively, 0.4 and 0.5 MPa lower in TE trees. Minor differences were observed in conduit mean hydraulic diameter and cell wall reinforcement. Our findings indicate significant and fast hydraulic acclimation under relatively mild drought in both tree species. Acclimation was well coordinated between branches and foliage, which might be essential for survival and productivity of mature trees under future drought periods., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Leaf-level anatomical variation is readily apparent within tall tree crowns, yet the relative importance of water and light availability in controlling this variation remains unclear. Sitka spruce (Picea sitchensis, (Bong.) Carr.) thrives in temperate rainforests of the Pacific Northwest, where it has historically reached heights >100 m, despite rarely living more than 400 years alongside redwoods that are five times older. We examined leaves of trees up to 97 m tall using a combination of transverse sections, longitudinal sections, epidermal imprints and whole-leaf measurements to explore the combined effects of water stress and light availability on leaf development in P. sitchensis. In contrast to the situation in tall Cupressaceae, light availability-not hydraulic limitation-is the primary ecological driver of leaf-level anatomical variation in P. sitchensis. While height-associated decreases in leaf length and mesoporosity are best explained by hydrostatic constraints on leaf elongation, the majority of anatomical traits we measured reflect acclimation to light availability, including increases in leaf width and vascular tissue areas in the brightest parts of the crown. Along with these changes, the appearance of abaxial stomata in the bright upper crown, and the arrangement of mesophyll in uniseriate, transverse plates-with radially arranged apoplastic pathways leading directly to stomata before bridging them with a V-shaped cell-may enhance gas exchange and hydraulic conductivity. This suite of leaf traits suggests an adaptive strategy that maximizes photosynthesis at the expense of water-stress tolerance. Anatomical investigations spanning the height gradient in tall tree crowns build our understanding of mechanisms underlying among-species variation in growth rates, life spans, and potential responses to climate change., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

During winter evergreens maintain a sustained form of thermal energy dissipation that results in reduced photochemical efficiency measured using the chlorophyll fluorescence parameter Fv/Fm. Eastern white pine (Pinus strobus L.) and white spruce [Picea glauca (Moench) Voss] have been shown to differ in their rate of recovery of Fv/Fm from winter stress. The goal of this study was to monitor changes in photosynthetic protein abundance and phosphorylation status during winter recovery that accompany these functional changes. An additional goal was to determine whether light-dependent changes in light harvesting complex II (LHCII) phosphorylation occur during winter conditions. We used a combination of field measurements and recovery experiments to monitor chlorophyll fluorescence and photosynthetic protein content and phosphorylation status. We found that pine recovered three times more slowly than spruce, and that the kinetics of recovery in spruce included a rapid and slow component, while in pine there was only a rapid component to recovery. Both species retained relatively high amounts of the light harvesting protein Lhcb5 (CP26) and the PsbS protein during winter, suggesting a role for these proteins in sustained thermal dissipation. Both species maintained high phosphorylation of LHCII and the D1 protein in darkness during winter. Pine and spruce differed in the kinetics of the dephosphorylation of LHCII and D1 upon warming, suggesting the rate of dephosphorylation of LHCII and D1 may be important in the rapid component of recovery from winter stress. Finally, we demonstrated that light-dependent changes in LHII phosphorylation do not continue to occur on subzero winter days and that needles are maintained in a phosphorylation pattern consistent with the high light conditions to which those needles are exposed. Our results suggest a role for retained phosphorylation of both LHCII and D1 in maintenance of the photosynthetic machinery in a winter conformation that maximizes thermal energy dissipation., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Background: How plants adapt their developmental patterns to regular seasonal changes is an important question in biology. The annual growth cycle in perennial long-lived trees is yet another example of how plants can adapt to seasonal changes. The two main signals that plants rely on to respond to seasonal changes are photoperiod and temperature, and these signals have critical roles in the temporal regulation of the annual growth cycle of trees., Scope: This review presents the latest findings to provide insight into the molecular mechanisms that underlie how photoperiodic and temperature signals regulate seasonal growth in trees., Conclusion: The results point to a high level of conservation in the signalling pathways that mediate photoperiodic control of seasonal growth in trees and flowering in annual plants such as arabidopsis. Furthermore, the data indicate that symplastic communication may mediate certain aspects of seasonal growth. Although considerable insight into the control of phenology in model plants such as poplar and spruce has been obtained, the future challenge is extending these studies to other, non-model trees., (© The Author 2017. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com)

Hydraulic redistribution (HR) of soil water through plant roots is a crucial phenomenon improving the water balance of plants and ecosystems. It is mostly described under severe drought, and not yet studied under moderate drought. We tested the potential of HR under moderate drought, hypothesizing that (H1) tree species redistribute soil water in their roots even under moderate drought and that (H2) neighboring plants are supported with water provided by redistributing plants. Trees were planted in split-root systems with one individual (i.e., split-root plant, SRP) having its roots divided between two pots with one additional tree each. Species were 2- to  4-year-old English oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). A gradient in soil water potential (ψsoil) was established between the two pots (-0.55 ± 0.02 MPa and -0.29 ± 0.03 MPa), and HR was observed by labeling with deuterium-enriched water. Irrespective of species identity, 93% of the SRPs redistributed deuterium enriched water from the moist to the drier side, supporting H1. Eighty-eight percent of the plants in the drier pots were deuterium enriched in their roots, with 61 ± 6% of the root water originating from SRP roots. Differences in HR among species were related to their root anatomy with diffuse-porous xylem structure in both beech and-opposing the stem structure-oak roots. In spruce, we found exclusively tracheids. We conclude that water can be redistributed within roots of different tree species along a moderate ψsoil gradient, accentuating HR as an important water source for drought-stressed plants, with potential implications for ecohydrological and plant physiological sciences. It remains to be shown to what extent HR occurs under field conditions in Central Europe., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Future limited water availability may reduce the potential of tree improvement to increase timber yields. We investigated ecophysiological variation between full-sibling families of Sitka spruce (Picea sitchensis (Bong.) Carr.) growing under contrasting water availability conditions: control (optimal) water availability and limited water availability. One-year-old seedlings of nine improved families plus an unimproved seed lot were grown in pots in a greenhouse and the two water availability treatments imposed via drip irrigation. Whole-plant water use varied between families. Stomatal conductance and the light-saturated quantum yield of photosystem II at times differed between families, but not consistently. Certain families showed considerably greater increases in electron transport rate with increasing photosynthetically active radiation. Limited water availability resulted in reduced branch water potential, leaf stomatal conductance and transpiration per unit leaf area, and increased whole-plant water-use efficiency, in all genetic material. The responses of plant water use and leaf carbon isotope composition to water limitation, were, however, initially influenced by variation in vigour between families-with conservative growth in some material slowing the decline in substrate moisture content. As the duration of water deficit extended, these variables showed a more uniform response across families. Between-family variation in physiological mechanisms of drought tolerance was not detected. Thus, for Sitka spruce, assessing juvenile material may not allow selection to prevent reductions in productivity associated with long-term sub-optimal growing conditions, but screening for conservative growth (within families as well as between families) may be beneficial where survival of relatively short-term water limitation is the primary concern., (© The Author 2016. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Three main hypotheses have been postulated over the past century to explain the outbreaking population dynamics of eastern spruce budworm, Choristoneura fumiferana (Clemens). The Silviculture Hypothesis first arose in the 1920s, with the idea that outbreaks were driven by forestry practices favoring susceptible softwood species. In the 1960s, it was proposed that populations were governed by Multiple Equilibria, with warm weather conditions releasing low-density populations from the regulatory control of natural enemies. Dispersal from outbreak foci, or "epicenters," was seen as causing widespread outbreaks that eventually collapsed following resource depletion. However, in the 1980s, following the re-analysis of data from the 1940s outbreak in New Brunswick, this interpretation was challenged. The alternative Oscillatory Hypothesis proposed that budworm population dynamics were governed by a second-order density-dependent process, with oscillations being driven by natural enemy-victim interactions. Under this hypothesis, weather and resource availability contribute to secondary fluctuations around the main oscillation, and weather and moth dispersal serve to synchronize population cycles regionally. Intensive, independent population studies during the peak and declining phases of the 1980s outbreak supported the principal tenet of the Oscillatory Hypothesis, but concluded that host plant quality played a more important role than this hypothesis proposed. More recent research on the early phase of spruce budworm cycles suggests that mate-finding and natural-enemy-driven Allee effects in low-density populations might be overcome by immigration of moths, which can facilitate the onset of outbreaks. Even more recent research has supported components of all three hypotheses attempting to explain spruce budworm dynamics. In the midst of a new rising outbreak (2006-present), we discuss the evolution of debates surrounding these hypotheses from a historic perspective, examine gaps in current knowledge, and suggest avenues for future research (e.g., intensive studies on low-density populations) to better understand and manage spruce budworm populations., (© The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Temperate forests are expected to be particularly vulnerable to drought and soil drying because they are not adapted to such conditions and perform best in mesic environments. Here we ask (i) how sensitively four common temperate tree species (Fagus sylvatica, Picea abies, Acer pseudoplatanus and Fraxinus excelsior) respond in their water relations to summer soil drying and seek to determine (ii) if species-specific responses to summer soil drying are related to the onset of declining water status across the four species. Throughout 2012 and 2013 we determined tree water deficit (TWD) as a proxy for tree water status from recorded stem radius changes and monitored sap flow rates with sensors on 16 mature trees studied in the field at Lägeren, Switzerland. All tree species responded equally in their relative maximum TWD to the onset of declining soil moisture. This implies that the water supply of all tree species was affected by declining soil moisture and that none of the four species was able to fully maintain its water status, e.g., by access to alternative water sources in the soil. In contrast we found strong and highly species-specific responses of sap flow to declining soil moisture with the strongest decline in P. abies (92%), followed by F. sylvatica (53%) and A. pseudoplatanus (48%). F. excelsior did not significantly reduce sap flow. We hypothesize the species-specific responses in sap flow to declining soil moisture that occur despite a simultaneous increase in relative TWD in all species reflect how fast these species approach critical levels of their water status, which is most likely influenced by species-specific traits determining the hydraulic properties of the species tree., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

Accurate temporal and spatial measurements of leaf optical traits (i.e., absorption, reflectance and transmittance) are paramount to photosynthetic studies. These optical traits are also needed to couple radiative transfer and physiological models to facilitate the interpretation of optical data. However, estimating leaf optical traits in leaves with complex morphologies remains a challenge. Leaf optical traits can be measured using integrating spheres, either by placing the leaf sample in one of the measuring ports (External Method) or by placing the sample inside the sphere (Internal Method). However, in leaves with complex morphology (e.g., needles), the External Method presents limitations associated with gaps between the leaves, and the Internal Method presents uncertainties related to the estimation of total leaf area. We introduce a modified version of the Internal Method, which bypasses the effect of gaps and the need to estimate total leaf area, by painting the leaves black and measuring them before and after painting. We assess and compare the new method with the External Method using a broadleaf and two conifer species. Both methods yielded similar leaf absorption estimates for the broadleaf, but absorption estimates were higher with the External Method for the conifer species. Factors explaining the differences between methods, their trade-offs and their advantages and limitations are also discussed. We suggest that the new method can be used to estimate leaf absorption in any type of leaf independently of its morphology, and be used to study further the impact of gap fraction in the External Method., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Projections of future climates suggest that droughts (Ds) may become more frequent and severe in many regions. Genetic variation, especially within populations in traits related to D resistance, is poorly investigated in forest trees, but this knowledge is necessary to better understand how forests will respond to water shortages. In this study, we investigated variability among seven open-pollinated half-sib families of a single population and two population-level progenies of Norway spruce (Picea abies (L.) H. Karst.) in their gas exchange response to imposed D and xylem vulnerability to embolism. During their third growing season, saplings were subjected to three treatments-control (C), D (for 19 weeks) and broken drought (BD, 54 days without watering starting in mid-July, then well-watered). In response to D, all families reduced their stomatal conductance (gs) and light-saturated rates of photosynthesis (Amax) in a similar way. After rewatering, the xylem water potential (Ψ) recovered in the BD treatment, but gs and Amax remained lower than in C. Needle starch concentration was altered in both D treatments compared with C. Xylem of D-exposed trees was more vulnerable to embolism than in C. The minimum attained safety margin remained positive for all families, indicating that no catastrophic hydraulic failure occurred in stem xylem during D. Significant family variation was found for Ψ early in the D (midday Ψ between -1.2 and -1.8 MPa), and for needle damage, but not for sapling mortality. Family variation found at the initial stages of D, and not afterward, suggests that all families responded similarly to greater D intensity, exhibiting the species-specific response. Limited variation at the family level indicates that the response to D and the traits we examined were conservative within the species. This may limit breeding opportunities for increased D resistance in Norway spruce in light of expected climatic changes., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Unfavourable soil conditions frequently limit tree regeneration in mountain forests on calcareous bedrock. Rocky, shallow organic soils on dolomite pose a particular problem for tree regeneration due to commonly restricted water and nutrient supplies. Moreover, an often dense layer of understorey vegetation competes for the limited resources available. Hence, an array of interacting factors impairs tree seedlings' performance on dolomite, but there is little information on the ecophysiological mechanisms. We studied the effects of substrate, competing vegetation and foliar nutrient concentrations on the photosynthetic rate (A), stomatal conductance (gs) and leaf water potentials (ψ) of sycamore (Acer pseudoplatanus L.), beech (Fagus sylvatica L.), spruce [Picea abies (L.) Karst.] and larch (Larix decidua Mill.) under controlled (well-watered/drought-stressed) conditions and under prevailing field conditions. While A and gs of well-watered spruce in the pot experiment were reduced by the mineral substrate, the organic dolomite substrate with dense competing vegetation reduced gs and ψ of sycamore, spruce and larch under drought-stressed conditions in the field. For sycamore and spruce, A and gs were strongly correlated with foliar nitrogen (N) and potassium (K) concentrations in the pot experiment. In contrast, soil water primarily affected beech and larch. Finally, dense competing vegetation negatively affected A and gs of spruce and A of larch on dolomite. Our results highlight the critical role of N, K and water availability for tree seedlings in shallow soils on calcareous bedrock. On these sites, natural tree regeneration is at particular risk from episodic drought, a likely consequence of climate change., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Climate change can induce substantial modifications in xylem structure and water transport capacity of trees exposed to environmental constraints. To elucidate mechanisms of xylem plasticity in response to climate, we retrospectively analysed different cell anatomical parameters over tree-ring series in Norway spruce (Picea abies L. Karst.). We sampled 24 trees along an altitudinal gradient (1200, 1600 and 2100 m above sea level, a.s.l.) and processed 2335 ± 1809 cells per ring. Time series for median cell lumen area (MCA), cell number (CN), tree-ring width (RW) and tree-ring-specific hydraulic conductivity (Kr) were crossed with daily temperature and precipitation records (1926-2011) to identify climate influence on xylem anatomical traits. Higher Kr at the low elevation site was due to higher MCA and CN. These variables were related to different aspects of intra-seasonal climatic variability under different environmental conditions, with MCA being more sensitive to summer precipitation. Winter precipitation (snow) benefited most parameters in all the sites. Descending the gradient, sensitivity of xylem features to summer climate shifted mostly from temperature to precipitation. In the context of climate change, our results indicate that higher summer temperatures at high elevations will benefit cell production and xylem hydraulic efficiency, whereas reduced water availability at lower elevations could negatively affect tracheids enlargement and thus stem capacity to transport water., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

As sessile organisms, plants have evolved different methods to defend against attacks and have adapted their defense measures to discriminate between mechanical damage and herbivory by insects. One of the ways that plant defenses are triggered is via elicitors from insect oral secretions (OS). In this study, we investigated the ability of second-instar (L2) spruce budworm [SBW; Choristoneura fumiferana (Clemens)] to alter the volatile organic compounds (VOCs) of four conifer species [Abies balsamea (L.) Mill., Picea mariana (Miller) B.S.P., Picea glauca (Moench) Voss, Picea rubens (Sargent)] and found that the emission profiles from all host trees were drastically changed after herbivory. We then investigated whether some of the main elicitors (fatty acid conjugates [FACs], β-glucosidase, and glucose oxidase) studied were present in SBW OS. FACs (glutamine and glutamic acid) based on linolenic, linoleic, oleic, and stearic acids were all observed in varying relative quantities. Hydroxylated FACs, such as volicitin, were not observed. Enzyme activity for β-glucosidase was also measured and found present in SBW OS, whereas glucose oxidase activity was not found in the SBW labial glands. These results demonstrate that SBW L2 larvae have the ability to induce VOC emissions upon herbivory and that SBW OS contain potential elicitors to induce these defensive responses. These data will be useful to further evaluate whether these elicitors can separately induce the production of specific VOCs and to investigate whether and how these emissions benefit the plant., (© Crown copyright 2015.)

Background and Aims: Plants growing under elevated atmospheric CO2 concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO2 the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase., Methods: The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4-5 years to either ambient (AC; 385 µmol mol(-1)) or elevated (EC; 700 µmol mol(-1)) CO2 concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques., Key Results: Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO2 assimilation (Amax) and a decline in photorespiration (RL) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (Topt) of Amax, Rubisco carboxylation rates (VCmax) and RL were found for EC saplings compared with AC saplings. However, the shifts in Topt of Amax were instantaneous, and disappeared when measured at identical CO2 concentrations. Higher values of Topt at elevated CO2 were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry., Conclusions: Elevated CO2 instantaneously increases temperature optima of Amax due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO2 concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO2 is unlikely. The hypothesis that long-term cultivation at elevated CO2 leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary., (© The Author 2015. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The genome sequence of Tricholoma vaccinum was obtained to predict its secretome in order to elucidate communication of T. vaccinum with its host tree spruce (Picea abies) in interkingdom signaling. The most prominent protein domains within the 206 predicted secreted proteins belong to energy and nutrition (52%), cell wall degradation (19%) and mycorrhiza establishment (9%). Additionally, we found small secreted proteins that show typical features of effectors potentially involved in host communication. From the secretome, 22 proteins could be identified, two of which showed higher protein abundances after spruce root exudate exposure, while five were downregulated in this treatment. The changes in T. vaccinum protein excretion with first recognition of the partner were used to identify small secreted proteins with the potential to act as effectors in the mutually beneficial symbiosis. Our observations support the hypothesis of a complex communication network including a cocktail of communication molecules induced long before physical contact of the partners., (© FEMS 2015. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Many studies have demonstrated linkages between the occurrence of fog and ecophysiological functioning in cloud forests, but few have investigated hydraulic functioning as a determining factor that explains sharp changes in vegetation. The objective of this study was to compare the plant water status during cloud-immersed and non-immersed conditions and hydraulic vulnerability in branches and roots of species across a temperate, mountain fog ecotone. Because cloud forests are often dark, cool and very moist, we expected cloud forest species to have less drought-tolerant characteristics (i.e., lower Pe and P50-the pressures required to induce a 12 and 50% loss in hydraulic conductivity, respectively) relative to non-cloud forest species in adjacent (lower elevation) forests. Additionally, due to the ability of cloud forest species to absorb cloud-fog water, we predicted greater improvements in hydraulic functioning during fog in cloud forest species relative to non-cloud forest species. Across the cloud forest ecotone, most species measured were very resistant to losses in conductivity with branch P50 values from -4.5 to -6.0 MPa, hydraulic safety margins (Ψmin - P50) >1.5 MPa and low calculated hydraulic conductivity losses. Roots had greater vulnerabilities, with P50 values ranging from -1.4 to -2.5 MPa, leading to greater predicted losses in conductivity (∼20%). Calculated values suggested strong losses of midday leaf hydraulic conductance in three of the four species, supporting the hydraulic segmentation hypothesis. In both cloud forest and hardwood species, Ψs were greater on foggy days than sunny days, demonstrating the importance of fog periods to plant water balance across fog regimes. Thus, frequent fog did not result in systemic changes in hydraulic functioning or vulnerability to embolism across our temperate cloud forest ecotone. Finally, roots functioned with lower hydraulic conductivity than branches, suggesting that they may serve as more sensitive indicators of hydraulic functioning in these mesic, foggy ecosystems., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Climate-related variations in functional traits of boreal tree species can result both from physiological acclimation and genetic adaptation of local populations to their biophysical environment. To improve our understanding and prediction of the physiological and growth responses of populations to climate change, we studied the role of climate of seed origin in determining variations in functional traits and its implications for tree improvement programs for a commonly reforested boreal conifer, white spruce (Picea glauca (Moench) Voss). We evaluated growth, root-to-shoot ratio (R/S), specific leaf area (SLA), needle nitrogen (N(mass)), total non-structural carbohydrates (NSC) and photosynthetic traits of 3-year-old seedlings in a greenhouse experiment using seed from six seed orchards (SO) representing the different regions where white spruce is reforested in Québec. Height and total dry mass (TDM) were positively correlated with photosynthetic capacity (A(max)), stomatal conductance (g(s)) and mesophyll conductance (g(m)). Total dry mass, but not height growth, was strongly correlated with latitude of seed origin (SO) and associated climate variables. A(max), g(s), g(m) and more marginally, photosynthetic nitrogen-use efficiency (PNUE) were positively associated with the mean July temperature of the SO, while water use efficiency (WUE) was negatively associated. Maximum rates of carboxylation (V(cmax)), maximum rates of electron transport (J(max)), SLA, N(mass), NSC and R/S showed no pattern. Our results did not demonstrate a higher Amax for northern seed orchards, although this has been previously hypothesized as an adaptation mechanism for maintaining carbon uptake in northern regions. We suggest that gs, gm, WUE and PNUE are the functional traits most associated with fine-scale geographic clines and with the degree of local adaptation of white spruce populations to their biophysical environments. These geographic patterns may reflect in situ adaptive genetic differences in photosynthetic efficiency along the cline., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Non-structural carbohydrates (NSCs) are critical to maintain plant metabolism under stressful environmental conditions, but we do not fully understand how NSC allocation and utilization from storage varies with stress. While it has become established that storage allocation is unlikely to be a mere overflow process, very little empirical evidence has been produced to support this view, at least not for trees. Here we present the results of an intensively monitored experimental manipulation of whole-tree carbon (C) balance (young Picea abies (L.) H Karst.) using reduced atmospheric [CO2] and drought to reduce C sources. We measured specific C storage pools (glucose, fructose, sucrose, starch) over 21 weeks and converted concentration measurement into fluxes into and out of the storage pool. Continuous labeling ((13)C) allowed us to track C allocation to biomass and non-structural C pools. Net C fluxes into the storage pool occurred mainly when the C balance was positive. Storage pools increased during periods of positive C gain and were reduced under negative C gain. (13)C data showed that C was allocated to storage pools independent of the net flux and even under severe C limitation. Allocation to below-ground tissues was strongest in control trees followed by trees experiencing drought followed by those grown under low [CO2]. Our data suggest that NSC storage has, under the conditions of our experimental manipulation (e.g., strong progressive drought, no above-ground growth), a high allocation priority and cannot be considered an overflow process. While these results also suggest active storage allocation, definitive proof of active plant control of storage in woody plants requires studies involving molecular tools., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We developed models to describe the responses of four commonly examined leaf traits (mass per area, weight, area and nitrogen (N) concentration) to gradients of light, soil nutrients and tree height in three conifer species of contrasting shade tolerance. Our observational dataset from the sub-boreal spruce forests of British Columbia included subalpine fir (Abies lasioscarpa [Hook.] Nutt; high shade tolerance), interior spruce (Picea glauca × Picea engelmannii [Moench] Voss; intermediate shade tolerance) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia; low shade tolerance) saplings from 0.18 to 4.87 m tall, in 8-98% of total incident light, from field sites with <17.6 kg ha(-1) to >46.8 kg ha(-1) total dissolved N. Leaf weights and areas showed strong positive responses to light and height, but little or no response to soil nutrients. Parameter estimates indicated that the shape of leaf weight and area responses to light corresponded with shade tolerance ranking for the three species; pine had the most linear response whereas spruce and fir had asymptotic responses. Leaf N concentration responded positively to soil nutrients, negatively to light and idiosyncratically to height. The negative effect of light was only apparent on sites of high soil nutrient availability, and parameter estimates for the shape of the negative response also corresponded to shade tolerance ranking (apine = -0.79, aspruce = -0.15, afir = -0.07). Of the traits we measured, leaf mass per area showed the least response to light, soil nutrient and height gradients. Although it is a common practice in comparisons across many species, characterizing these conifers by mean values of their leaf traits would miss important intraspecific variation across environmental and size gradients. In these forests, parameter estimates representing the intraspecific variability of leaf trait responses can be used to understand relative shade tolerances., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Background and Aims: Bamboo is well known for its fast growth and excellent mechanical performance, but the underlying relationships between its structure and properties are only partially known. Since it lacks secondary thickening, bamboo cannot use adaptive growth in the same way as a tree would in order to modify the geometry of the stem and increase its moment of inertia to cope with bending stresses caused by wind loads. Consequently, mechanical adaptation can only be achieved at the tissue level, and this study aims to examine how this is achieved by comparison with a softwood tree species at the tissue, fibre and cell wall levels., Methods: The mechanical properties of single fibres and tissue slices of stems of mature moso bamboo (Phyllostachys pubescens) and spruce (Picea abies) latewood were investigated in microtensile tests. Cell parameters, cellulose microfibril angles and chemical composition were determined using light and electron microscopy, wide-angle X-ray scattering and confocal Raman microscopy., Key Results: Pronounced differences in tensile stiffness and strength were found at the tissue and fibre levels, but not at the cell wall level. Thus, under tensile loads, the differing wall structures of bamboo (multilayered) and spruce (sandwich-like) appear to be of minor relevance., Conclusions: The superior tensile properties of bamboo fibres and fibre bundles are mainly a result of amplified cell wall formation, leading to a densely packed tissue, rather than being based on specific cell wall properties. The material optimization towards extremely compact fibres with a multi-lamellar cell wall in bamboo might be a result of a plant growth strategy that compensates for the lack of secondary thickening growth at the tissue level, which is not only favourable for the biomechanics of the plant but is also increasingly utilized in terms of engineering products made from bamboo culms., (© The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

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.)