Isometric scaling to model water transport in conifer tree rings across time and environments.

The hydraulic properties of xylem determine the ability of plants to efficiently and safely provide water to their leaves. These properties are key to understanding plant responses to environmental conditions and evaluating their fate under a rapidly changing climate. However, their assessment is hindered by the challenges of quantifying basic hydraulic components such as bordered pits and tracheids. Here, we use isometric scaling between tracheids and pit morphology to merge partial hydraulic models of the tracheid component and to upscale these properties to the tree-ring level in conifers. Our new model output is first cross-validated with the literature and then applied to cell anatomical measurements from Larix sibirica tree rings formed under harsh conditions in southern Siberia to quantify the intra- and inter-annual variability in hydraulic properties. The model provides a means of assessing how different-sized tracheid components contribute to the hydraulic properties of the ring. Upscaled results indicate that natural inter- and intra-ring anatomical variations have a substantial impact on the tree's hydraulic properties. Our model facilitates the assessment of important xylem functional attributes because it requires only the more accessible measures of cross-sectional tracheid size. This approach, if applied to dated tree rings, provides a novel way to investigate xylem structure-function relationships across time and environmental conditions.

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.

What prevails in climatic response of Pinus sylvestris in-between its range limits in mountains: slope aspect or elevation?

The roles of slope orientation and elevational temperature gradient were investigated for Scots pine (Pinus sylvestris L.) growth in the middle of its growth range, where these factors can significantly modulate microclimate and thus plant growth. We assumed that slope orientation causes more complex and severe effects than elevation because it influences all three main factors of plant growth: light, heat, and moisture. In addition to the total ring width, the earlywood and latewood width and latewood ratio were considered variables that contain information about tree ring growth during the season and wood structure over all tree life span on three sampling sites at different elevations and opposite slopes. Despite the observed dependence of pine growth rate on temperature and solar radiation, the mean latewood ratio is stable and similar between all sampling sites, being presumably defined by the genotype of individual trees. The seasonality of the climatic response of tree growth is bound to spatiotemporal variation of the vegetative season timing due to the elevational temperature lapse and local warming. However, its direction is primarily defined by slope orientation, where southern slope is moisture-limited, even at adjacent sites, and divergent climatic reactions of earlywood (weak moisture-limited in the last decades) and latewood growth (temperature-limited) were revealed on the northern slope.

New Methods in Digital Wood Anatomy: The Use of Pixel-Contrast Densitometry with Example of Angiosperm Shrubs in Southern Siberia.

This methodological study describes the adaptation of a new method in digital wood anatomy, pixel-contrast densitometry, for angiosperm species. The new method was tested on eight species of shrubs and small trees in Southern Siberia, whose wood structure varies from ring-porous to diffuse-porous, with different spatial organizations of vessels. A two-step transformation of wood cross-section photographs by smoothing and Otsu's classification algorithm was proposed to separate images into cell wall areas and empty spaces within (lumen) and between cells. Good synchronicity between measurements within the ring allowed us to create profiles of wood porosity (proportion of empty spaces) describing the growth ring structure and capturing inter-annual differences between rings. For longer-lived species, 14-32-year series from at least ten specimens were measured. Their analysis revealed that maximum (for all wood types), mean, and minimum porosity (for diffuse-porous wood) in the ring have common external signals, mostly independent of ring width, i.e., they can be used as ecological indicators. Further research directions include a comparison of this method with other approaches in densitometry, clarification of sample processing, and the extraction of ecologically meaningful data from wood structures.

Temperature-induced responses of xylem structure of Larix sibirica (Pinaceae) from the Russian Altay.

PREMISE OF THE STUDY: Xylem structure determines the hydraulic and mechanical properties of a stem, and its plasticity is fundamental for maintaining tree performance under changing conditions. Unveiling the mechanism and the range of xylem adjustment is thus necessary to anticipate climate change impacts on vegetation. METHODS: To understand the mechanistic process and the functional impact of xylem responses to warming in a cold-limited environment, we investigated the relationship between temperature and tracheid anatomy along a 312-yr tree-ring chronology of Larix sibirica trees from the Altay Mountains in Russia. KEY RESULTS: Climate-growth analyses indicated that warming favors wider earlywood cell lumen, thicker latewood walls, denser maximum latewood, and wider rings. The temperature signal of the latewood was stronger (r > 0.7) and covered a longer and more stable period (from June to August) than that of earlywood and tree-ring width. Long-term analyses indicated a diverging trend between lumen and cell wall of early- and latewood. CONCLUSIONS: Xylem anatomy appears to respond to warming temperatures. A warmer early-growing season raises water conduction capacity by increasing the number and size of earlywood tracheids. The higher-performing earlywood tracheids promote more carbon fixation of the latewood cells by incrementing the rate of assimilation when summer conditions are favorable for growth. The diverging long-term variation of lumen and cell wall in earlywood vs. latewood suggests that xylem adjustments in latewood increase mechanical integrity and support increasing tree size under the ameliorated growing conditions.

Modeling of the Statistical Distribution of Tracheids in Conifer Rings: Finding Universal Criterion for Earlywood-Latewood Distinction.

The quantitative description of growth rings is yet incomplete, including the functional division into earlywood and latewood. Methods developed to date, such as the Mork criterion for conifers, can be biased and arbitrary depending on species and growth conditions. We proposed the use of modeling of the statistical distribution of tracheids to determine a universal criterion applicable to all conifer species. Thisstudy was based on 50-year anatomical measurements of Pinus sylvestris L., Pinus sibirica Du Tour, and Picea obovata Ledeb. near the upper tree line in the Western Sayan Mountains (South Siberia). Statistical distributions of the cell wall thickness (CWT)-to-radial-diameter (D) ratio and its slope were investigated for raw and standardized data (divided by the mean). The bimodal distribution of the slope for standardized CWT and D was modeled with beta distributions for earlywood and latewood tracheids and a generalized normal distribution for transition wood to account for the gradual shift in cell traits. The modelcan describe with high accuracy the growth ring structure for species characterized by various proportions of latewood, histometric traits, and gradual or abrupt transition. The proportion of two (or three, including transition wood) zones in the modeled distribution is proposed as a desired criterion.

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.

Factors Limiting Radial Growth of Conifers on Their Semiarid Borders across Kazakhstan.

The forests of Central Asia are biodiversity hotspots at risk from rapid climate change, but they are understudied in terms of the climate-growth relationships of trees. This classical dendroclimatic case study was performed for six conifer forest stands near their semiarid boundaries across Kazakhstan: (1-3) Pinus sylvestris L., temperate forest steppes; (4-5) Picea schrenkiana Fisch. & C.A. Mey, foothills, the Western Tien Shan, southeast; (6) Juniperus seravschanica Kom., montane zone, the Western Tien Shan, southern subtropics. Due to large distances, correlations between local tree-ring width (TRW) chronologies are significant only within species (pine, 0.19-0.50; spruce, 0.55). The most stable climatic response is negative correlations of TRW with maximum temperatures of the previous (from -0.37 to -0.50) and current (from -0.17 to -0.44) growing season. The strength of the positive response to annual precipitation (0.10-0.48) and Standardized Precipitation Evapotranspiration Index (0.15-0.49) depends on local aridity. The timeframe of climatic responses shifts to earlier months north-to-south. For years with maximum and minimum TRW, differences in seasonal maximal temperatures (by ~1-3 °C) and precipitation (by ~12-83%) were also found. Heat stress being the primary factor limiting conifer growth across Kazakhstan, we suggest experiments there on heat protection measures in plantations and for urban trees, alongside broadening the coverage of the dendroclimatic net with accents on the impact of habitat conditions and climate-induced long-term growth dynamics.

Climate impacts on tree-ring stable isotopes across the Northern Hemispheric boreal zone.

Boreal regions are changing rapidly with anthropogenic global warming. In order to assess risks and impacts of this process, it is crucial to put these observed changes into a long-term perspective. Summer air temperature variability can be well reconstructed from conifer tree rings. While the application of stable isotopes can potentially provide complementary climatic information over different seasons. In this study, we developed new triple stable isotope chronologies in tree-ring cellulose (δ13Ctrc, δ18Otrc, δ2Htrc) from a study site in Canada. Additionally, we performed regional aggregated analysis of available stable isotope chronologies from 6 conifers' tree species across high-latitudinal (HL) and - altitudinal (HA) as well as Siberian (SIB) transects of the Northern Hemispheric boreal zone. Our results show that summer air temperature still plays an important role in determining tree-ring isotope variability at 11 out of 24 sites for δ13Ctrc, 6 out of 18 sites for δ18Otrc and 1 out of 6 sites for δ2Htrc. Precipitation, relative humidity and vapor pressure deficit are significantly and consistently recorded in both δ13Ctrc and δ18Otrc along HL. Summer sunshine duration is captured by all isotopes, mainly for HL and HA transects, indicating an indirect link with an increase in air and leaf temperature. A mixed temperature-precipitation signal is preserved in δ13Ctrc and δ18Otrc along SIB transect. The δ2Htrc data obtained for HL-transect provide information not only about growing seasonal moisture and temperature, but also capture autumn, winter and spring sunshine duration signals. We conclude that a combination of triple stable isotopes in tree-ring studies can provide a comprehensive description of climate variability across the boreal forest zone and improve ecohydrological reconstructions.

Modern aridity in the Altai-Sayan mountain range derived from multiple millennial proxies.

Temperature and precipitation changes are crucial for larch trees growing at high-elevation sites covered by permafrost in the Altai-Sayan mountain range (ASMR). To contextualize the amplitude of recent climate fluctuations, we have to look into the past by analyzing millennial paleoclimatic archives recording both temperature and precipitation. We developed annually resolved 1500-year tree-ring cellulose chronologies (δ13Ccell, δ18Ocell), and used these new records to reconstruct the variability in local summer precipitation and air temperature. We combined our new local reconstructions with existing paleoclimatic archives available for the Altai. The data show a strong decreasing trend by ca. 49% in regional summer precipitation, along with a regional summer temperature increase towards the twenty-first century, relative to the preceding 1500 years. Modern dry conditions (1966-2016 CE) in the ASMR are the result of simultaneous summer warming and decreased precipitation. Our new reconstructions also demonstrate that climate change in the ASMR is much stronger compared to the global average.