Significance: Forest trees can live for hundreds to thousands of years, and they play a critical role in mitigating global warming by fixing approximately 15% of anthropogenic CO 2 emissions annually by wood formation. However, the environmental factors triggering wood formation onset in springtime and the cellular mechanisms underlying this onset remain poorly understood, since wood forms beneath the bark and is difficult to monitor. We report that the onset of wood formation in Northern Hemisphere conifers is driven primarily by photoperiod and mean annual temperature. Understanding the unique relationships between exogenous factors and wood formation could aid in predicting how forest ecosystems respond and adapt to climate warming, while improving the assessment of long-term and high-resolution observations of global biogeochemical cycles.
Future seasonal dynamics of wood formation in hyperarid environments are still unclear. Although temperature‐driven extension of the growing season and increased forest productivity are expected for boreal and temperate biomes under global warming, a similar trend remains questionable in water‐limited regions. We monitored cambial activity in a montane stand of ponderosa pine (Pinus ponderosa) from the Mojave Desert for 2 consecutive years (2015–2016) showing opposite‐sign anomalies between warm‐ and cold‐season precipitation. After the wet winter/spring of 2016, xylogenesis started 2 months earlier compared to 2015, characterized by abundant monsoonal (July–August) rainfall and hyperarid spring. Tree size did not influence the onset and ending of wood formation, highlighting a predominant climatic control over xylem phenological processes. Moisture conditions in the previous month, in particular soil water content and dew point, were the main drivers of cambial phenology. Latewood formation started roughly at the same time in both years; however, monsoonal precipitation triggered the formation of more false rings and density fluctuations in 2015. Because of uncertainties in future precipitation patterns simulated by global change models for the Southwestern United States, the dependency of P. ponderosa on seasonal moisture implies a greater conservation challenge than for species that respond mostly to temperature conditions.
Dendroclimatic proxies can be generated from the analysis of wood cellular structures, allowing for a more complete understanding of the physiological mechanisms that control the climatic response of tree species. Century-long (1870-2013) time series of anatomical parameters were developed for Great Basin bristlecone pine (Pinus longaeva D.K. Bailey) by capturing strongly contrasted microscopic images through a Confocal Laser Scanning Microscope. Environmental information embedded in wood anatomical series was analyzed in comparison with ring-width series using measures of empirical signal strength. Response functions were calculated against monthly climatic variables to evaluate climate sensitivity of cellular features (e.g. lumen area; lumen diameter) for the period 1950-2013. Calibration-verification tests were used to determine the potential to generate long climate reconstructions from these anatomical proxies. A total of eight tree-ring parameters (two ring-width and six chronologies of xylem anatomical parameters) were analyzed. Synchronous variability among samples varied among tree-ring parameters, usually decreasing from ring width to anatomical features. Cellular parameters linked to plant hydraulic performance (e.g. tracheid lumen area and radial lumen diameter) showed empirical signal strength similar to ring-width series, while noise was predominant in chronologies of lumen tangential width and cell-wall thickness. Climatic signals were different between anatomical and ring-width chronologies, revealing a positive and temporally stable correlation of tracheid size (i.e. lumen and cell diameter) with monthly (i.e. March) and seasonal precipitation. In particular, tracheid lumen diameter emerged as a reliable moisture indicator and was then used to reconstruct total March-August precipitation from 1870 to 2013. Wood anatomy holds great potential to refine and expand dendroclimatic records by allowing estimates of plant physiological adaptations to external stressors. Integrating xylem cellular features with ring-width chronologies can widen our understanding of past climatic variability (including annual extreme events) and improve the evaluation of long-term plant response to drought, especially in connection with future warming scenarios.
The timing of wood formation is crucial to determine how environmental factors affect tree growth. The long-lived bristlecone pine (Pinus longaeva D. K. Bailey) is a foundation treeline species in the Great Basin of North America reaching stem ages of about 5000 years. We investigated stem cambial phenology and radial size variability to quantify the relative influence of environmental variables on bristlecone pine growth. Repeated cellular measurements and half-hourly dendrometer records were obtained during 2013 and 2014 for two high-elevation stands included in the Nevada Climate-ecohydrological Assessment Network. Daily time series of stem radial variations showed rehydration and expansion starting in late April–early May, prior to the onset of wood formation at breast height. Formation of new xylem started in June and lasted until mid-September. There were no differences in phenological timing between the two stands, or in the air and soil temperature thresholds for the onset of xylogenesis. A multiple logistic regression model highlighted a separate effect of air and soil temperature on xylogenesis, the relevance of which was modulated by the interaction with vapor pressure and soil water content. While air temperature plays a key role in cambial resumption after winter dormancy, soil thermal conditions coupled with snowpack dynamics also influence the onset of wood formation by regulating plant–soil water exchanges. Our results help build a physiological understanding of climate–growth relationships in P. longaeva, the importance of which for dendroclimatic reconstructions can hardly be overstated. In addition, environmental drivers of xylogenesis at the treeline ecotone, by controlling the growth of dominant species, ultimately determine ecosystem responses to climatic change.
As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels.