From xylogenesis to tree rings: wood traits to investigate tree response to environmental changes.

International audienceIt is noteworthy that the largest part of global vegetation biomass depends on a thin layer of cells: the vascular cambium. Understanding the wood formation processes and relationships with environmental factors is a crucial and timely research question requiring interdisciplinary efforts, also to upscale the information gained and to evaluate implications for tree growth and forest productivity. We provide an overview of wood formation processes up to tree-ring development, bearing in mind that the combined action of intrinsic factors and environmental drivers determines the anatomical traits of a tree ring formed at a specific time and position within the tree’s architecture. After briefly reviewing intrinsic factors, we focus attention on environmental drivers highlighting how a correct interpretation of environmental signals in tree rings must be grounded in a deep knowledge of xylogenesis and consequent wood anatomical traits. We provide guidelines on novel methods and approaches recently developed to study xylem formation. We refer to existing literature on established techniques for retrospective analyses in tree-ring series of anatomical and isotopic traits, to assess long-term ecophysiological responses to environmental variations, also giving advice on possible bias because of interand within-tree variability. Finally, we highlight that, once the temporal axis of intra-annual tree-ring variability of xylem traits is established by xylogenesis analysis, a multidisciplinary approach linking classical dendro-ecology, wood functional traits (dendro-anatomy) and eco-physiology (here focusing on dendro-isotopes) allows a better interpretation of past environmental events hidden in tree rings, and more reliable forecasts of wood growth in response to climate change

Critical temperatures for xylogenesis in conifers of cold climates

Aim To identify temperatures at which cell division and differentiation are active in order to verify the existence of a common critical temperature determining growth in conifers of cold climates. Location Ten European and Canadian sites at different latitudes and altitudes. Methods The periods of cambial activity and cell differentiation were assessed on a weekly time-scale on histological sections of cambium and wood tissue collected over 2 to 5 years per site from 1998 to 2005 from the stems of seven conifer species. All data were compared with daily air temperatures recorded from weather stations located close to the sites. Logistic regressions were used to calculate the probability of xylogenesis and of cambium being active at a given temperature. Results Xylogenesis lasted from May to October, with a growing period varying from 3 to 5 months depending on location and elevation. Despite the wide geographical range of the monitored sites, temperatures for onset and ending of xylogenesis converged towards narrow ranges with average values around 4-5, 8-9 and 13-14 degrees C for daily minimum, mean and maximum temperature, respectively. On the contrary, cell division in the cambium stopped in July-August, when temperatures were still high. Main conclusions Wood formation in conifers occurred when specific critical temperatures were reached. Although the timing and duration of xylogenesis varied among species, sites and years, the estimated temperatures were stable for all trees studied. These results provide biologically based evidence that temperature is a critical factor limiting production and differentiation of xylem cells in cold climates. Although daily temperatures below 4-5 degrees C are still favourable for photosynthesis, thermal conditions below these values could inhibit the allocation of assimilated carbon to structural investment, i.e. xylem growth

Are neighboring trees in tune? Wood formation in Pinus pinaster

Neighboring trees growing under identical environmental conditions can exhibit different dynamics and periods of growth. Despite the recent advances in cambial biology, the exogenous and endogenous factors generating asynchronous xylem growths still remain undetermined. This study investigated timings and duration of xylem formation in maritime pine (Pinus pinaster Ait.) from an even-aged plantation in Portugal growing under Mediterranean climate. Cambial phenology and stem diameter were monitored weekly, from March to December 2010, on two classes of trees divided according to the tree ring widths of the last 15 years, but similar age and size: fast- and slow-growing trees. We tested the hypothesis that differences in tree ring widths result from cell production which in turn affects timings of xylogenesis and that the bimodal growth pattern, typical of the Mediterranean, originates from a double reactivation of the cambium: in spring and autumn. Cambial activity started earlier and ended later in fast-growing trees, confirming that cell production is a key factor determining the duration of xylogenesis. Intra-annual variations in stem diameter recorded by band dendrometers revealed two peaks of increment occurring in spring and late summer. However, the number of cambial cells did not increase in late summer, which suggested that the second peak of increment was caused by stem rehydration, rather than by a reactivation of cell division. These results demonstrated that the variability in the timings of xylem phenology observed among trees of the same age and size and growing under similar environmental conditions was closely related to cell production and not to age or size per se.This study was supported by the Fundac¸a˜o para a Cieˆncia e a Tecnologia, Ministe´rio da Educac¸a˜o e Cieˆncia (FCT) cofinanced by Compete, through the project PTDC/AAC-AMB/111675/ 2009. Joana Vieira was supported by a Ph.D. Grant (SFRH/BD/ 48089/2008) and Filipe Campelo by a postdoctoral research grant (SFRH/BPD/47822/2008), both grants from FCT with funds from POPH (Portuguese Operational Human Potential Program) and QREN Portugal (Portuguese National Strategic Reference Framework)

Photoperiod and temperature as dominant environmental drivers triggering secondary growth resumption in Northern Hemisphere conifers

Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes