Higher latewood to earlywood ratio increases resistance of radial growth to severe droughts in larch.

As drought has caused great losses of tree growth across the world, the mechanism of how trees adapt to drought has been extensively investigated. However, how trees change their late- to earlywood ratio (LER) to adapt to severe drought events remains poorly understood. We used a network of Larix principis-rupprechtii earlywood and latewood width data from 1979 to 2018, covering most of the distribution of planted larch across North China, to investigate how latewood proportion affected trees' resistance to drought. The interactions among LER, minimum temperature, vapor pressure deficit (VPD), growing season length, and their contributions to drought resistant (Rt) were estimated using structural equation models. The results show a significant increase in LER of the juvenile wood throughout the first 15 growth rings after which it stabilizes. The LER decreased significantly with elevation for the juvenile wood. March-May temperature and VPD were the main determinant in the LER of mature wood. The sensitivity of radial growth to droughts was positively changed with LER when LER was below 0.50, but negatively changed with LER when LER is above 0.50. We confirmed that high LER increases resistance of tree growth to severe droughts in L. principis-rupprechtii. Our results highlight that a higher proportion of latewood is formed in dry years, and this high drought sensitivity of LER in turn led to an increased resistance to drought. This combination of reduced radial growth during dry years, while the latewood proportion remains increases maybe an adaptive strategy of larch trees to cope with severe droughts.

Fading regulation of diurnal temperature ranges on drought-induced growth loss for drought-tolerant tree species.

Warming-induced droughts caused tree growth loss across the globe, leading to substantial carbon loss to the atmosphere. Drought-induced growth loss, however, can be regulated by changes in diurnal temperature ranges. Here, we investigated long term radial growth responses of 23 widespread distributed tree species from 2327 sites over the world and found that species' drought tolerances were significantly and positively correlated with diurnal temperature range-growth loss relationships for the period 1901-1940. Since 1940, this relationship has continued to fade, likely due to asymmetric day and night warming trends and the species' ability to deal with them. The alleviation of reduced diurnal temperature ranges on drought-induced growth loss was mainly found for drought resistant tree species. Overall, our results highlight the need to carefully consider diurnal temperature ranges and species-specific responses to daytime and nighttime warming to explore tree growth responses to current and future warmer and drier climates.

Reassessment of growth-climate relations indicates the potential for decline across Eurasian boreal larch forests.

Larch, a widely distributed tree in boreal Eurasia, is experiencing rapid warming across much of its distribution. A comprehensive assessment of growth on warming is needed to comprehend the potential impact of climate change. Most studies, relying on rigid calendar-based temperature series, have detected monotonic responses at the margins of boreal Eurasia, but not across the region. Here, we developed a method for constructing temporally flexible and physiologically relevant temperature series to reassess growth-temperature relations of larch across boreal Eurasia. Our method appears more effective in assessing the impact of warming on growth than previous methods. Our approach indicates widespread and spatially heterogeneous growth-temperature responses that are driven by local climate. Models quantifying these results project that the negative responses of growth to temperature will spread northward and upward throughout this century. If true, the risks of warming to boreal Eurasia could be more widespread than conveyed from previous works.

Insights into source/sink controls on wood formation and photosynthesis from a stem chilling experiment in mature red maple.

Whether sources or sinks control wood growth remains debated with a paucity of evidence from mature trees in natural settings. Here, we altered carbon supply rate in stems of mature red maples (Acer rubrum) within the growing season by restricting phloem transport using stem chilling; thereby increasing carbon supply above and decreasing carbon supply below the restrictions, respectively. Chilling successfully altered nonstructural carbon (NSC) concentrations in the phloem without detectable repercussions on bulk NSC in stems and roots. Ring width responded strongly to local variations in carbon supply with up to seven-fold differences along the stem of chilled trees; however, concurrent changes in the structural carbon were inconclusive at high carbon supply due to large local variability of wood growth. Above chilling-induced bottlenecks, we also observed higher leaf NSC concentrations, reduced photosynthetic capacity, and earlier leaf coloration and fall. Our results indicate that the cambial sink is affected by carbon supply, but within-tree feedbacks can downregulate source activity, when carbon supply exceeds demand. Such feedbacks have only been hypothesized in mature trees. Consequently, these findings constitute an important advance in understanding source-sink dynamics, suggesting that mature red maples operate close to both source- and sink-limitation in the early growing season.

Reduced diurnal temperature range mitigates drought impacts on larch tree growth in North China.

Forests are facing climate changes such as warmer temperatures, accelerated snowmelt, increased drought, as well as changing diurnal temperature ranges (DTR) and cloud cover regimes. How tree growth is influenced by the changes in daily to monthly temperatures and its associations with droughts has been extensively investigated, however, few studies have focused on how changes in sub-daily temperatures i.e., DTR, influence tree growth during drought events. Here, we used a network of Larix principis-rupprechtii tree-ring data from 1989 to 2018, covering most of the distribution of planted larch across North China, to investigate how DTR, cloud cover and their interactions influence the relationship between drought stress and tree growth. DTR showed a negative correlation with larch growth in 95 % of sites (rmean = -0.30, significant in 42 % of sites). Cloud cover was positively correlated with growth in 87 % of sites (rmean = 0.13, significant in 5 % of sites). Enhanced tree growth was found at lower DTR in the absence of severe drought. Our findings highlight that in the absence of severe droughts, reduced DTR benefits tree growth, while increased cloud cover tended to benefit tree growth only during severe drought periods. Given how DTR influences drought impacts on tree growth, net tree growth was found to be larger in regions with smaller DTR.

Inter-annual and inter-species tree growth explained by phenology of xylogenesis.

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.

Manipulating phloem transport affects wood formation but not local nonstructural carbon reserves in an evergreen conifer.

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, CO2 efflux and nonstructural carbon reserves in stems of mature white pines. Wood formation and stem CO2 efflux 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.

Evidence of unprecedented rise in growth synchrony from global tree ring records.

Changes in the temporal coherence between populations, which can influence their stability, resilience and persistence, remain a critical uncertainty of climate change. Recent studies have documented increasing spatial synchrony between populations at continental scales and linked it to anthropogenic climate change. However, the lack of long-term and global baseline perspectives on spatial synchrony presents a challenge to understanding the importance of these trends. Here, we show a steady rise in the spatial synchrony of annual tree growth from a global tree ring database over the past 50 years that is consistent across continents, species and environmental conditions and is unprecedented for the past millennium. Increasing growth synchrony coincided with warming trends and potentially rising synchrony in the temperature records. We discuss the potential driving mechanisms and the limitations in the interpretation of this trend, and we propose that increasing mutual dependency on external factors (also known as Moran's effect) linked to rising global temperatures is the most likely driver of more homogeneous global growth dynamics.

Snowmelt and early to mid-growing season water availability augment tree growth during rapid warming in southern Asian boreal forests.

Boreal forests are facing profound changes in their growth environment, including warming-induced water deficits, extended growing seasons, accelerated snowmelt, and permafrost thaw. The influence of warming on trees varies regionally, but in most boreal forests studied to date, tree growth has been found to be negatively affected by increasing temperatures. Here, we used a network of Pinus sylvestris tree-ring collections spanning a wide climate gradient the southern end of the boreal forest in Asia to assess their response to climate change for the period 1958-2014. Contrary to findings in other boreal regions, we found that previously negative effects of temperature on tree growth turned positive in the northern portion of the study network after the onset of rapid warming. Trees in the drier portion did not show this reversal in their climatic response during the period of rapid warming. Abundant water availability during the growing season, particularly in the early to mid-growing season (May-July), is key to the reversal of tree sensitivity to climate. Advancement in the onset of growth appears to allow trees to take advantage of snowmelt water, such that tree growth increases with increasing temperatures during the rapidly warming period. The region's monsoonal climate delivers limited precipitation during the early growing season, and thus snowmelt likely covers the water deficit so trees are less stressed from the onset of earlier growth. Our results indicate that the growth response of P. sylvestris to increasing temperatures strongly related to increased early season water availability. Hence, boreal forests with sufficient water available during crucial parts of the growing season might be more able to withstand or even increase growth during periods of rising temperatures. We suspect that other regions of the boreal forest may be affected by similar dynamics.

Stem Compression: A Means to Reversibly Reduce Phloem Transport in Tree Stems.

Stem compression reduces or terminates the phloem-mediated transport of carbohydrates and other solutes in tree stems, without causing permanent damage to phloem functioning (Henriksson et al. Tree Physiol. 35:1075-1085, 2015). This has been tested on two species of pine trees, with diameters ranging from 3 to 26 cm in a forest in northern Sweden (Henriksson et al. Tree Physiol. 35:1075-1085, 2015) and in Harvard Forest, USA. Halting the phloem transport of trees in a forest is useful for studying tree physiological processes related to, or dependent on, phloem-transported compounds as well as downstream processes, in particular interactions with soil microbes. Phloem compression can be deployed in the lab and field on single trees, subsets, or over larger areas, depending on what is relevant for a particular research question.

Ten-year assessment of the 100 priority questions for global biodiversity conservation.

In 2008, a group of conservation scientists compiled a list of 100 priority questions for the conservation of the world's biodiversity. However, now almost a decade later, no one has yet published a study gauging how much progress has been made in addressing these 100 high-priority questions in the peer-reviewed literature. We took a first step toward reexamining the 100 questions to identify key knowledge gaps that remain. Through a combination of a questionnaire and a literature review, we evaluated each question on the basis of 2 criteria: relevance and effort. We defined highly relevant questions as those that - if answered - would have the greatest impact on global biodiversity conservation and quantified effort based on the number of review publications addressing a particular question, which we used as a proxy for research effort. Using this approach, we identified a set of questions that, despite being perceived as highly relevant, have been the focus of relatively few review publications over the past 10 years. These questions covered a broad range of topics but predominantly tackled 3 major themes: conservation and management of freshwater ecosystems, role of societal structures in shaping interactions between people and the environment, and impacts of conservation interventions. We believe these questions represent important knowledge gaps that have received insufficient attention and may need to be prioritized in future research.

Modeling Tree Growth Taking into Account Carbon Source and Sink Limitations.

Increasing CO2 concentrations are strongly controlled by the behavior of established forests, which are believed to be a major current sink of atmospheric CO2. There are many models which predict forest responses to environmental changes but they are almost exclusively carbon source (i.e., photosynthesis) driven. Here we present a model for an individual tree that takes into account the intrinsic limits of meristems and cellular growth rates, as well as control mechanisms within the tree that influence its diameter and height growth over time. This new framework is built on process-based understanding combined with differential equations solved by numerical method. Our aim is to construct a model framework of tree growth for replacing current formulations in Dynamic Global Vegetation Models, and so address the issue of the terrestrial carbon sink. Our approach was successfully tested for stands of beech trees in two different sites representing part of a long-term forest yield experiment in Germany. This model provides new insights into tree growth and limits to tree height, and addresses limitations of previous models with respect to sink-limited growth.

Evaluation of climate-related carbon turnover processes in global vegetation models for boreal and temperate forests.

Turnover concepts in state-of-the-art global vegetation models (GVMs) account for various processes, but are often highly simplified and may not include an adequate representation of the dominant processes that shape vegetation carbon turnover rates in real forest ecosystems at a large spatial scale. Here, we evaluate vegetation carbon turnover processes in GVMs participating in the Inter-Sectoral Impact Model Intercomparison Project (ISI-MIP, including HYBRID4, JeDi, JULES, LPJml, ORCHIDEE, SDGVM, and VISIT) using estimates of vegetation carbon turnover rate (k) derived from a combination of remote sensing based products of biomass and net primary production (NPP). We find that current model limitations lead to considerable biases in the simulated biomass and in k (severe underestimations by all models except JeDi and VISIT compared to observation-based average k), likely contributing to underestimation of positive feedbacks of the northern forest carbon balance to climate change caused by changes in forest mortality. A need for improved turnover concepts related to frost damage, drought, and insect outbreaks to better reproduce observation-based spatial patterns in k is identified. As direct frost damage effects on mortality are usually not accounted for in these GVMs, simulated relationships between k and winter length in boreal forests are not consistent between different regions and strongly biased compared to the observation-based relationships. Some models show a response of k to drought in temperate forests as a result of impacts of water availability on NPP, growth efficiency or carbon balance dependent mortality as well as soil or litter moisture effects on leaf turnover or fire. However, further direct drought effects such as carbon starvation (only in HYBRID4) or hydraulic failure are usually not taken into account by the investigated GVMs. While they are considered dominant large-scale mortality agents, mortality mechanisms related to insects and pathogens are not explicitly treated in these models.

Carbon residence time dominates uncertainty in terrestrial vegetation responses to future climate and atmospheric CO2.

Future climate change and increasing atmospheric CO2 are expected to cause major changes in vegetation structure and function over large fractions of the global land surface. Seven global vegetation models are used to analyze possible responses to future climate simulated by a range of general circulation models run under all four representative concentration pathway scenarios of changing concentrations of greenhouse gases. All 110 simulations predict an increase in global vegetation carbon to 2100, but with substantial variation between vegetation models. For example, at 4 °C of global land surface warming (510-758 ppm of CO2), vegetation carbon increases by 52-477 Pg C (224 Pg C mean), mainly due to CO2 fertilization of photosynthesis. Simulations agree on large regional increases across much of the boreal forest, western Amazonia, central Africa, western China, and southeast Asia, with reductions across southwestern North America, central South America, southern Mediterranean areas, southwestern Africa, and southwestern Australia. Four vegetation models display discontinuities across 4 °C of warming, indicating global thresholds in the balance of positive and negative influences on productivity and biomass. In contrast to previous global vegetation model studies, we emphasize the importance of uncertainties in projected changes in carbon residence times. We find, when all seven models are considered for one representative concentration pathway × general circulation model combination, such uncertainties explain 30% more variation in modeled vegetation carbon change than responses of net primary productivity alone, increasing to 151% for non-HYBRID4 models. A change in research priorities away from production and toward structural dynamics and demographic processes is recommended.