Identifying the main drivers for the production and maturation of Scots pine tracheids along a temperature gradient

Even though studies monitoring the phenology and seasonal dynamics of the wood formation have accumulated for several conifer species across the Northern Hemisphere, the environmental control of tracheid production and differentiation is still fragmentary. With microcore and environmental data from six stands in Finland and France, we built auto-calibrated data-driven black box models for analyzing the most important factors controlling the tracheid production and maturation in Scots pine stem. In the best models, estimation was accurate to within a fraction of a tracheid per week. We compared the relative results of models built using different predictors, and found that the rate of tracheid production was partly regular but current and previous air temperature had influence on the sites in the middle of the temperature range and photosynthetic production in the coldest ones. The rate of mature cell production was more difficult to relate to the predictors but recent photosynthetic production was included in all successful models.Peer reviewe

A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere

Background and Aims Ongoing global warming has been implicated in shifting phenological patterns such as the timing and duration of the growing season across a wide variety of ecosystems. Linear models are routinely used to extrapolate these observed shifts in phenology into the future and to estimate changes in associated ecosystem properties such as net primary productivity. Yet, in nature, linear relationships may be special cases. Biological processes frequently follow more complex, non-linear patterns according to limiting factors that generate shifts and discontinuities, or contain thresholds beyond which responses change abruptly. This study investigates to what extent cambium phenology is associated with xylem growth and differentiation across conifer species of the northern hemisphere. Methods Xylem cell production is compared with the periods of cambial activity and cell differentiation assessed on a weekly time scale on histological sections of cambium and wood tissue collected from the stems of nine species in Canada and Europe over 1-9 years per site from 1998 to 2011. Key Results The dynamics of xylogenesis were surprisingly homogeneous among conifer species, although dispersions from the average were obviously observed. Within the range analysed, the relationships between the phenological timings were linear, with several slopes showing values close to or not statistically different from 1. The relationships between the phenological timings and cell production were distinctly non-linear, and involved an exponential pattern Conclusions The trees adjust their phenological timings according to linear patterns. Thus, shifts of one phenological phase are associated with synchronous and comparable shifts of the successive phases. However, small increases in the duration of xylogenesis could correspond to a substantial increase in cell production. The findings suggest that the length of the growing season and the resulting amount of growth could respond differently to changes in environmental condition

Partial asynchrony of coniferous forest carbon sources and sinks at the intra-annual time scale

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.O

Partial asynchrony of coniferous forest carbon sources and sinks at the intra-annual time scale.

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

Woody biomass production lags stem-girth increase by over one month in coniferous forests

Wood is the main terrestrial biotic reservoir for long-term carbon sequestration1, and its formation in trees consumes around 15% of anthropogenic carbon dioxide emissions each year2. However, the seasonal dynamics of woody biomass production cannot be quantified from eddy covariance or satellite observations. As such, our understanding of this key carbon cycle component, and its sensitivity to climate, remains limited. Here, we present high-resolution cellular based measurements of wood formation dynamics in three coniferous forest sites in northeastern France, performed over a period of 3 years. We show that stem woody biomass production lags behind stem-girth increase by over 1 month. We also analyse more general phenological observations of xylem tissue formation in Northern Hemisphere forests and find similar time lags in boreal, temperate, subalpine and Mediterranean forests. These time lags question the extension of the equivalence between stem size increase and woody biomass production to intra-annual time scales3–6. They also suggest that these two growth processes exhibit differential sensitivities to local environmental conditions. Indeed, in the wellwatered French sites the seasonal dynamics of stem-girth increase matched the photoperiod cycle, whereas those of woody biomass production closely followed the seasonal course of temperature. We suggest that forecasted changes in the annual cycle of climatic factors7 may shift the phase timing of stem size increase and woody biomass production in the future

Intra-annual wood formation dynamics of three conifer species (silver fir, Norway spruce, and Scots pine) in northeast France : From the description of the growth seasonal patterns to the study of the environmental influence on the kinetics of cell development and the anatomical features of the xylem

La formation du bois (xylogénèse) produit une large partie de la biomasse de la planète et une ressource essentielle pour l'Homme. Les cellules du bois sont produites par division dans le cambium puis s'élargissent, forment une paroi épaisse lignifiée et meurent. Pendant l'année, ces processus sont définis par des dates, durées et vitesses qui caractérisent la dynamique intra-annuelle de la xylogénèse. Cette dynamique reste peu explorée alors que c'est un aspect clé, car c'est elle qui détermine la quantité et la qualité du bois produit et c'est sur elle que les facteurs de régulation agissent. Ce travail vise à améliorer nos connaissances sur la dynamique intra-annuelle de la xylogénèse. Pendant trois ans (2007-2009), la xylogénèse a été suivie pour 45 arbres de trois espèces de conifères (sapin pectiné, épicéa commun et pin sylvestre) dans les Vosges. Pour ça, des petits échantillons de bois ont été prélevés chaque semaine sur le tronc des arbres sélectionnés. Les échantillons ont été préparés au laboratoire, puis des sections anatomiques ont été réalisées pour observer la xylogénèse au microscope. Cette thèse a permis d'améliorer notre connaissance du fonctionnement de la xylogénèse, un système biologique d'une fascinante complexité. Nous avons caractérisé - grâce à l'innovation d'une méthode statistique performante - les aspects méconnus de la dynamique de différenciation des cellules du bois. Nous avons alors pu dévoiler les mécanismes par lesquels la dynamique de la xylogénèse donne forme à la structure du cerne, établir la dynamique intra-annuelle de l'accumulation du carbone dans le bois et évaluer les mécanismes de l'influence du climat sur la xylogénèseWood formation (xylogenesis) produces a large part of the biomass of this planet and provides a crucial resource to Mankind. Wood cells are produced by division in the cambium, after what they enlarge, build a lignified thick wall and die. During a year, these processes take place at certain dates, last for certain durations and go at certain rates. These dates, durations and rates characterize the intra-annual dynamics of xylogenesis. This dynamics remains poorly explored whereas it is a key aspect as it determines the quantity and quality of the produced wood and conveys the influence of intrinsic (gene, hormone) and extrinsic (environment) regulatory factors. This work aims to improve our knowledge on the intra-annual dynamics of xylogenesis. During three years (2007-2009), xylogenesis was monitored for 45 trees of three conifer species (silver fir, Norway spruce, and Scots pine) in northeast France. For that, small wood samples were collected weekly on tree stem. Samples were prepared at the laboratory, and anatomical sections were cut to observe xylogenesis under a light microscope. This thesis has improved our knowledge on the functioning of xylogenesis, a biological system of a fascinating complexity. We characterized - thanks to the development of an efficient statistical method - the little known aspects of wood cell differentiation dynamics. Based on this characterization, we eluded the mechanisms by which xylogenesis dynamics shapes tree ring structure, we established the intra-annual dynamics of carbon accumulation in wood and we evaluated the mechanisms of the climate influence on xylogenesi

Dynamique intra-annuelle de la formation du bois de trois espèces de conifères (sapin pectiné, épicéa commun et pin sylvestre) dans les Vosges : De la description des patrons saisonniers de la croissance à l'étude de l'influence de l'environnement sur la cinétique du développement cellulaire et les caractéristiques anatomiques du xylène

Wood formation (xylogenesis) produces a large part of the biomass of this planet and provides a crucial resource to Mankind. Wood cells are produced by division in the cambium, after what they enlarge, build a lignified thick wall and die. During a year, these processes take place at certain dates, last for certain durations and go at certain rates. These dates, durations and rates characterize the intra-annual dynamics of xylogenesis. This dynamics remains poorly explored whereas it is a key aspect as it determines the quantity and quality of the produced wood and conveys the influence of intrinsic (gene, hormone) and extrinsic (environment) regulatory factors. This work aims to improve our knowledge on the intra-annual dynamics of xylogenesis. During three years (2007-2009), xylogenesis was monitored for 45 trees of three conifer species (silver fir, Norway spruce, and Scots pine) in northeast France. For that, small wood samples were collected weekly on tree stem. Samples were prepared at the laboratory, and anatomical sections were cut to observe xylogenesis under a light microscope. This thesis has improved our knowledge on the functioning of xylogenesis, a biological system of a fascinating complexity. We characterized - thanks to the development of an efficient statistical method - the little known aspects of wood cell differentiation dynamics. Based on this characterization, we eluded the mechanisms by which xylogenesis dynamics shapes tree ring structure, we established the intra-annual dynamics of carbon accumulation in wood and we evaluated the mechanisms of the climate influence on xylogenesisLa formation du bois (xylogénèse) produit une large partie de la biomasse de la planète et une ressource essentielle pour l'Homme. Les cellules du bois sont produites par division dans le cambium puis s'élargissent, forment une paroi épaisse lignifiée et meurent. Pendant l'année, ces processus sont définis par des dates, durées et vitesses qui caractérisent la dynamique intra-annuelle de la xylogénèse. Cette dynamique reste peu explorée alors que c'est un aspect clé, car c'est elle qui détermine la quantité et la qualité du bois produit et c'est sur elle que les facteurs de régulation agissent. Ce travail vise à améliorer nos connaissances sur la dynamique intra-annuelle de la xylogénèse. Pendant trois ans (2007-2009), la xylogénèse a été suivie pour 45 arbres de trois espèces de conifères (sapin pectiné, épicéa commun et pin sylvestre) dans les Vosges. Pour ça, des petits échantillons de bois ont été prélevés chaque semaine sur le tronc des arbres sélectionnés. Les échantillons ont été préparés au laboratoire, puis des sections anatomiques ont été réalisées pour observer la xylogénèse au microscope. Cette thèse a permis d'améliorer notre connaissance du fonctionnement de la xylogénèse, un système biologique d'une fascinante complexité. Nous avons caractérisé - grâce à l'innovation d'une méthode statistique performante - les aspects méconnus de la dynamique de différenciation des cellules du bois. Nous avons alors pu dévoiler les mécanismes par lesquels la dynamique de la xylogénèse donne forme à la structure du cerne, établir la dynamique intra-annuelle de l'accumulation du carbone dans le bois et évaluer les mécanismes de l'influence du climat sur la xylogénès

Xylogenesis: Coniferous Trees of Temperate Forests Are Listening to the Climate Tale during the Growing Season But Only Remember the Last Words!

International audienceThe complex inner mechanisms that create typical conifer tree-ring structure (i.e. the transition from large, thin-walled earlywood cells to narrow, thick-walled latewood cells) were recently unraveled. However, what physiological or environmental factors drive xylogenesis key processes remain unclear. Here, we aim to quantify the influence of seasonal variations in climatic factors on the spectacular changes in the kinetics of wood cell differentiation and in the resulting treering structure. Wood formation was monitored in three sites over 3 years for three coniferous species (Norway spruce [Picea abies], Scots pine [Pinus sylvestris], and silver fir [Abies alba]). Cell differentiation rates and durations were calculated and related to tracheid final dimensions and corresponding climatic conditions. On the one hand, we found that the kinetics of cell enlargement and the final size of the tracheids were not explained by the seasonal changes in climatic factors. On the other hand, decreasing temperatures strongly constrained cell wall deposition rates during latewood formation. However, the influence of temperature was permanently written into tree-ring structure only for the very last latewood cells, when the collapse of the rate of wall deposition was no longer counterbalanced by the increase of its duration. Our results show that the formation of the typical conifer tree-ring structure, in normal climatic conditions, is only marginally driven by climate, suggesting strong developmental control of xylogenesis. The late breakage of the compensatory mechanism at work in the wall deposition process appears as a clue to understand the capacity of the maximum latewood density to record past temperature conditions

Discrete triangular associated kernel and bandwidth choices in semiparametric estimation for count data

International audienc