Jian‐Guo Huang

Abstract The interaction between xylem phenology and climate assesses forest growth and productivity and carbon storage across biomes under changing environmental conditions. We tested the hypothesis that patterns of wood formation are maintained unaltered despite the temperature changes across cold ecosystems. Wood microcores were collected weekly or biweekly throughout the growing season for periods varying between 1 and 13 years during 1998–2014 and cut in transverse sections for assessing the onset and ending of the phases of xylem differentiation. The data set represented 1321 trees belonging to 10 conifer species from 39 sites in the Northern Hemisphere and covering an interval of mean annual temperature exceeding 14 K. The phenological events and mean annual temperature of the sites were related linearly, with spring and autumnal events being separated by constant intervals across the range of temperature analysed. At increasing temperature, first enlarging, wall‐thickening and mature tracheids appeared earlier, and last enlarging and wall‐thickening tracheids occurred later. Overall, the period of wood formation lengthened linearly with the mean annual temperature, from 83.7 days at −2 °C to 178.1 days at 12 °C, at a rate of 6.5 days °C −1 . April–May temperatures produced the best models predicting the dates of wood formation. Our findings demonstrated the uniformity of the process of wood formation and the importance of the environmental conditions occurring at the time of growth resumption. Under warming scenarios, the period of wood formation might lengthen synchronously in the cold biomes of the Northern Hemisphere.

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.

Frost events during the active growth period of plants can cause extensive frost damage with tremendous economic losses and dramatic ecological consequences. A common assumption is that climate warming may bring along a reduction in the frequency and severity of frost damage to vegetation. On the other hand, it has been argued that rising temperature in late winter and early spring might trigger the so called "false spring", that is, early onset of growth that is followed by cold spells, resulting in increased frost damage. By combining daily gridded climate data and 1,489 k in situ phenological observations of 27 tree species from 5,565 phenological observation sites in Europe, we show here that temporal changes in the risk of spring frost damage with recent warming vary largely depending on the species and geographical locations. Species whose phenology was especially sensitive to climate warming tended to have increased risk of frost damage. Geographically, compared with continental areas, maritime and coastal areas in Europe were more exposed to increasing occurrence of frost and these late spring frosts were getting more severe in the maritime and coastal areas. Our results suggest that even though temperatures will be elevated in the future, some phenologically responsive species and many populations of a given species will paradoxically experience more frost damage in the future warming climate. More attention should be paid to the increased frost damage in responsive species and populations in maritime areas when developing strategies to mitigate the potential negative impacts of climate change on ecosystems in the near future.

Nonstructural carbohydrates (NSCs) play a crucial role in xylem formation and represent, with water, the main constraint to plant growth. We assessed the relationships between xylogenesis and NSCs in order to (1) verify the variance explained by NSCs and (2) determine the influence of intrinsic (tissue supplying carbon) and extrinsic (water availability and temperature) factors. During 2 years, wood formation was monitored in saplings of black spruce (Picea mariana) subjected to a dry period of about 1 month in June and exposed to different temperature treatments in a greenhouse. In parallel, NSC concentrations were determined by extracting the sugar compounds from two tissues (cambium and inner xylem), both potentially supplying carbon for wood formation. A mixed-effect model was used to assess and quantify the potential relationships. Total xylem cells, illustrating meristematic activity, were modeled as a function of water, sucrose, and d-pinitol (conditional r(2) of 0.79). Water availability was ranked as the most important factor explaining total xylem cell production, while the contribution of carbon was lower. Cambium stopped dividing under water deficit, probably to limit the number of cells remaining in differentiation without an adequate amount of water. By contrast, carbon factors were ranked as most important in explaining the variation in living cells (conditional r(2) of 0.49), highlighting the functional needs during xylem development, followed by the tissue supplying the NSCs (cambium) and water availability. This study precisely demonstrates the role of carbon and water in structural growth expressed as meristematic activity and tissue formation.