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Intra-annual density fluctuations (IADFs) are triggered by environmental cues, but whether they are distributed uniformly throughout the stem is not well documented. The spatial distribution of IADFs could help us understand variations in cambial sensitivity to environmental cues throughout the tree. We investigate how IADF distribution varies radially, longitudinally, and circumferentially within white pine (Pinus strobus L.) stems. We took wood samples at breast height, near branches, and at the top of the trees. We identified IADFs visually and measured their radial position within a ring as well as their circumferential arc in cross-sections. Intra-annual density fluctuations occurred in 22.2% of rings. The radial position of IADFs within a ring was remarkably consistent at roughly 80% of the total annual radial increment across heights, trees, and years of formation. The main factors affecting the likelihood of IADF occurrence were ring width, year of formation, and the interaction between the two. Being near branches or at the top of the tree slightly increased the probability of occurrence. Though the sample size was not large enough to provide conclusive results about the circumferential distribution of IADFs, our data suggest that the circumferential arc of the IADFs might be conserved throughout the stem. 

ABSTRACT Stomata control plant water loss and photosynthetic carbon gain. Developing more generalized and accurate stomatal models is essential for earth system models and predicting responses under novel environmental conditions associated with global change. Plant optimality theories offer one promising approach, but most such theories assume that stomatal conductance maximizes photosynthetic net carbon assimilation subject to some cost orconstraintof water. We move beyond this approach by developing a new, generalized optimality theory of stomatal conductance, optimizing any non‐foliar proxy that requires water and carbon reserves, like growth, survival, and reproduction. We overcome two prior limitations. First, we reconcile the computational efficiency ofinstantaneousoptimization with a more biologically meaningfuldynamic feedbackoptimization over plant lifespans. Second, we incorporatenon‐steady‐statephysics in the optimization to account for the temporal changes in the water, carbon, and energy storage within a plant and its environment that occur over the timescales that stomata act, contrary to previous theories. Our optimal stomatal conductance compares well to observations from seedlings, saplings, and mature trees from field and greenhouse experiments. Our model predicts predispositions to mortality during the 2018 European drought and captures realistic responses to environmental cues, including the partial alleviation of heat stress by evaporative cooling and the negative effect of accumulating foliar soluble carbohydrates, promoting closure under elevated CO₂. We advance stomatal optimality theory by incorporating generalized evolutionary fitness proxies and enhance its utility without compromising its realism, offering promise for future models to more realistically and accurately predict global carbon and water fluxes. 

Abstract The Sun sporadically produces eruptive events leading to intense fluxes of solar energetic particles (SEPs) that dramatically disrupt the near-Earth radiation environment. Such events have been directly studied for the last decades but little is known about the occurrence and magnitude of rare, extreme SEP events. Presently, a few events that produced measurable signals in cosmogenic radionuclides such as 14 C, 10 Be and 36 Cl have been found. Analyzing annual 14 C concentrations in tree-rings from Switzerland, Germany, Ireland, Russia, and the USA we discovered two spikes in atmospheric 14 C occurring in 7176 and 5259 BCE. The ~2% increases of atmospheric 14 C recorded for both events exceed all previously known 14 C peaks but after correction for the geomagnetic field, they are comparable to the largest event of this type discovered so far at 775 CE. These strong events serve as accurate time markers for the synchronization with floating tree-ring and ice core records and provide critical information on the previous occurrence of extreme solar events which may threaten modern infrastructure. 

Wood formation consumes around 15% of the anthropogenic CO 2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes. 

Summary Wood formation determines major long‐term carbon (C) accumulation in trees and therefore provides a crucial ecosystem service in mitigating climate change. Nevertheless, we lack understanding of how species with contrasting wood anatomical types differ with respect to phenology and environmental controls on wood formation.In this study, we investigated the seasonality and rates of radial growth and their relationships with climatic factors, and the seasonal variations of stem nonstructural carbohydrates (NSC) in three species with contrasting wood anatomical types (red oak: ring‐porous; red maple: diffuse‐porous; white pine: coniferous) in a temperate mixed forest during 2017–2019.We found that the high ring width variability observed in both red oak and red maple was caused more by changes in growth duration than growth rate. Seasonal radial growth patterns did not vary following transient environmental factors for all three species. Both angiosperm species showed higher concentrations and lower inter‐annual fluctuations of NSC than the coniferous species.Inter‐annual variability of ring width varied by species with contrasting wood anatomical types. Due to the high dependence of annual ring width on growth duration, our study highlights the critical importance of xylem formation phenology for understanding and modelling the dynamics of wood formation. 

Abstract How variations in carbon supply affect wood formation remains poorly understood in particular in mature forest trees. To elucidate how carbon supply affects carbon allocation and wood formation, we attempted to manipulate carbon supply to the cambial region by phloem girdling and compression during the mid‐ and late‐growing season and measured effects on structural development, CO₂efflux and nonstructural carbon reserves in stems of mature white pines. Wood formation and stem CO₂efflux varied with a location relative to treatment (i.e., above or below the restriction). We observed up to twice as many tracheids formed above versus below the treatment after the phloem transport manipulation, whereas the cell‐wall area decreased only slightly below the treatments, and cell size did not change relative to the control. Nonstructural carbon reserves in the xylem, needles and roots were largely unaffected by the treatments. Our results suggest that low and high carbon supply affects wood formation, primarily through a strong effect on cell proliferation, and respiration, but local nonstructural carbon concentrations appear to be maintained homeostatically. This contrasts with reports of decoupling of source activity and wood formation at the whole‐tree or ecosystem level, highlighting the need to better understand organ‐specific responses, within‐tree feedbacks, as well as phenological and ontogenetic effects on sink‐source dynamics.