High preseason temperature variability drives convergence of xylem phenology in the Northern Hemisphere conifers.

Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future.

VS-Cambium-Developer: A New Predictive Model of Cambium Functioning under the Influence of Environmental Factors.

Climate changes influence seasonal tree-ring formation. The result is a specific cell structure dependent on internal processes and external environmental factors. One way to investigate and analyze these relationships is to apply diverse simulation models of tree-ring growth. Here, we have proposed a new version of the VS-Cambium-Developer model (VS-CD model), which simulates the cambial activity process in conifers. The VS-CD model does not require the manual year-to-year calibration of parameters over a long-term cell production reconstruction or forecast. Instead, it estimates cell production and simulates the dynamics of radial cell development within the growing seasons. Thus, a new software based on R programming technology, able to efficiently adapt to the VS model online platform, has been developed. The model was tested on indirect observations of the cambium functioning in Larix sibirica trees from southern Siberia, namely on the measured annual cell production from 1963 to 2011. The VS-CD model proves to simulate cell production accurately. The results highlighted the efficiency of the presented model and contributed to filling the gap in the simulations of cambial activity, which is critical to predicting the potential impacts of changing environmental conditions on tree growth.

A critical thermal transition driving spring phenology of Northern Hemisphere conifers.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (-3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°-66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

Plant-Soil-Climate Interaction in Observed and Simulated Tree-Radial Growth Dynamics of Downy Birch in Permafrost.

Climate change projections forecast most significant impacts on high-latitude forest ecosystems. Particularly, climate warming in boreal regions should increase fire severity and shorten its return interval. These processes can change the dynamics of boreal forests as younger stands become more dominating with a shift from gymnosperm to angiosperm. However, despite angiosperm's phenological and physiological traits have a high potential for ecophysiological and dendroclimatological studies in Siberia, they have been rarely investigated due to their short-term lifespan in comparison with gymnosperm. Modeling tree growth is a common way to understand tree growth responses to environmental changes since it allows using available experiment or field data to interpret observed climate-growth relationships based on the biological principles. In our study, we applied the process-based Vaganov-Shashkin (VS) model of tree-ring growth via a parameterization approach VS-oscilloscope for the first time to an angiosperm tree species (Betula pubescens Ehrh.) from continuous permafrost terrain to understand its tree-radial growth dynamic. The parameterization of the VS model provided highly significant positive correlations (p < 0.05) between the simulated growth curve and initial tree-ring chronologies for the period 1971-2011 and displayed the average duration of the growing season and intra-seasonal key limiting factors for xylem formation. Modeled result can be valid at the regional scale for remote birch stands, whereas, justification of the local non-climatic input data of the model provided precise site-specific tree growth dynamic and their substantiated responses to driving factors.

Forward Modeling Reveals Multidecadal Trends in Cambial Kinetics and Phenology at Treeline.

Significant alterations of cambial activity might be expected due to climate warming, leading to growing season extension and higher growth rates especially in cold-limited forests. However, assessment of climate-change-driven trends in intra-annual wood formation suffers from the lack of direct observations with a timespan exceeding a few years. We used the Vaganov-Shashkin process-based model to: (i) simulate daily resolved numbers of cambial and differentiating cells; and (ii) develop chronologies of the onset and termination of specific phases of cambial phenology during 1961-2017. We also determined the dominant climatic factor limiting cambial activity for each day. To asses intra-annual model validity, we used 8 years of direct xylogenesis monitoring from the treeline region of the Krkonose Mts. (Czechia). The model exhibits high validity in case of spring phenological phases and a seasonal dynamics of tracheid production, but its precision declines for estimates of autumn phenological phases and growing season duration. The simulations reveal an increasing trend in the number of tracheids produced by cambium each year by 0.42 cells/year. Spring phenological phases (onset of cambial cell growth and tracheid enlargement) show significant shifts toward earlier occurrence in the year (for 0.28-0.34 days/year). In addition, there is a significant increase in simulated growth rates during entire growing season associated with the intra-annual redistribution of the dominant climatic controls over cambial activity. Results suggest that higher growth rates at treeline are driven by (i) temperature-stimulated intensification of spring cambial kinetics, and (ii) decoupling of summer growth rates from the limiting effect of low summer temperature due to higher frequency of climatically optimal days. Our results highlight that the cambial kinetics stimulation by increasing spring and summer temperatures and shifting spring phenology determine the recent growth trends of treeline ecosystems. Redistribution of individual climatic factors controlling cambial activity during the growing season questions the temporal stability of climatic signal of cold forest chronologies under ongoing climate change.

Growth-limiting factors and climate response variability in Norway spruce (Picea abies L.) along an elevation and precipitation gradients in Slovenia.

Norway spruce (Picea abies L.) is among the most sensitive coniferous species to ongoing climate change. However, previous studies on its growth response to increasing temperatures have yielded contrasting results (from stimulation to suppression), suggesting highly site-specific responses. Here, we present the first study that applies two independent approaches, i.e. the nonlinear, process-based Vaganov-Shashkin (VS) model and linear daily response functions. Data were collected at twelve sites in Slovenia differing in climate regimes and ranging elevation between 170 and 1300 m a.s.l. VS model results revealed that drier Norway spruce sites at lower elevations are mostly moisture limited, while moist high-elevation sites are generally more temperature limited. Daily response functions match well the pattern of growth-limiting factors from the VS model and further explain the effect of climate on radial growth: prevailing growth-limiting factors correspond to the climate variable with higher correlations. Radial growth correlates negatively with rising summer temperature and positively with higher spring precipitation. The opposite response was observed for the wettest site at the highest elevation, which positively reacts to increased summer temperature and will most likely benefit from a warming climate. For all other sites, the future radial growth of Norway spruce largely depends on the balance between spring precipitation and summer temperature.

Ecological and conceptual consequences of Arctic pollution.

Although the effect of pollution on forest health and decline received much attention in the 1980s, it has not been considered to explain the 'Divergence Problem' in dendroclimatology; a decoupling of tree growth from rising air temperatures since the 1970s. Here we use physical and biogeochemical measurements of hundreds of living and dead conifers to reconstruct the impact of heavy industrialisation around Norilsk in northern Siberia. Moreover, we develop a forward model with surface irradiance forcing to quantify long-distance effects of anthropogenic emissions on the functioning and productivity of Siberia's taiga. Downwind from the world's most polluted Arctic region, tree mortality rates of up to 100% have destroyed 24,000 km2 boreal forest since the 1960s, coincident with dramatic increases in atmospheric sulphur, copper, and nickel concentrations. In addition to regional ecosystem devastation, we demonstrate how 'Arctic Dimming' can explain the circumpolar 'Divergence Problem', and discuss implications on the terrestrial carbon cycle.

Modeled Tracheidograms Disclose Drought Influence on Pinus sylvestris Tree-Rings Structure From Siberian Forest-Steppe.

Wood formation allows trees to adjust in a changing climate. Understanding what determine its adjustment is crucial to evaluate impacts of climatic changes on trees and forests growth. Despite efforts to characterize wood formation, little is known on its impact on the xylem cellular structure. In this study we apply the Vaganov-Shashkin model to generate synthetic tracheidograms and verify its use to investigate the formation of intra-annual density fluctuations (IADF), one of the most frequent climate tree-ring markers in drought-exposed sites. Results indicate that the model can produce realistic tracheidograms, except for narrow rings (<1 mm), when cambial activity stops due to an excess of drought or a lack of growth vigor. These observations suggest that IADFs are caused by a release of drought limitation to cells formation in the first half of the growing season, but that narrow rings are indicators of an even more extreme and persistent water stress. Taking the example of IADFs formation, this study demonstrated that the Vaganov-Shashkin model is a useful tool to study the climatic impact on tree-ring structures. The ability to produce synthetic tracheidogram represents an unavoidable step to link climate to tree growth and xylem functioning under future scenarios.

Trends in recent temperature and radial tree growth spanning 2000 years across northwest Eurasia.

This paper describes variability in trends of annual tree growth at several locations in the high latitudes of Eurasia, providing a wide regional comparison over a 2000-year period. The study focuses on the nature of local and widespread tree-growth responses to recent warming seen in instrumental observations, available in northern regions for periods ranging from decades to a century. Instrumental temperature data demonstrate differences in seasonal scale of Eurasian warming and the complexity and spatial diversity of tree-growing-season trends in recent decades. A set of long tree-ring chronologies provides empirical evidence of association between inter-annual tree growth and local, primarily summer, temperature variability at each location. These data show no evidence of a recent breakdown in this association as has been found at other high-latitude Northern Hemisphere locations. Using Kendall's concordance, we quantify the time-dependent relationship between growth trends of the long chronologies as a group. This provides strong evidence that the extent of recent widespread warming across northwest Eurasia, with respect to 100- to 200-year trends, is unprecedented in the last 2000 years. An equivalent analysis of simulated temperatures using the HadCM3 model fails to show a similar increase in concordance expected as a consequence of anthropogenic forcing.