Non-structural 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 two years, wood formation was monitored in saplings of black spruce submitted to a dry period of about one month in June and exposed to different temperature treatments in a greenhouse. In parallel, NSCs 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 pinitol (conditional R2 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 the most important in explaining the variation in the living cells (conditional R2 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.

Satellite remote sensing is a widely accessible tool to investigate the spatiotemporal variations in the bud phenology of evergreen species, which show limited seasonal changes in canopy greenness. However, there is a need for precise and compatible data to compare remote sensing time series with field observations. In this study, fortnightly MODIS-NDVI was fitted using double-logistic functions and calibrated using ordinal logit models with the sequential phases of bud phenology collected during 2015, 2017 and 2018 in a black spruce stand. Bud break and bud set were spatialized for the period 2009–2018 across 5000 stands in Quebec, Canada. The first phase of bud break and the last phase of bud set were observed in the field in mid-May and at the beginning of September, when NDVI was 80.5% and 92.2% of its maximum amplitude, respectively. The NDVI rate of change was estimated at 0.07 in spring and 0.04 in autumn. When spatialized on the black spruce stands, bud break was detected earlier in the southwestern regions (April–May), and later in the northeastern regions (mid to end of June). No clear trend was observed for bud set, with different patterns being detected among the years. Overall, the process bud break and bud set lasted 51 and 87 days, respectively. Our results demonstrate the potential of satellite remote sensing for providing reliable timings of bud phenological events using calibrated NDVI time series on wide regions that are remote or with limited access.

Intercomparison of satellite-derived vegetation phenology is scarce in remote locations because of the limited coverage area and low temporal resolution of field observations. By their reliable near-ground observations and high-frequency data collection, PhenoCams can be a robust tool for intercomparison of land surface phenology derived from satellites. This study aims to investigate the transition dates of black spruce (Picea mariana (Mill.) B.S.P.) phenology by comparing fortnightly the MODIS normalized difference vegetation index (NDVI) and the enhanced vegetation index (EVI) extracted using the Google Earth Engine (GEE) platform with the daily PhenoCam-based green chromatic coordinate (GCC) index. Data were collected from 2016 to 2019 by PhenoCams installed in six mature stands along a latitudinal gradient of the boreal forests of Quebec, Canada. All time series were fitted by double-logistic functions, and the estimated parameters were compared between NDVI, EVI, and GCC. The onset of GCC occurred in the second week of May, whereas the ending of GCC occurred in the last week of September. We demonstrated that GCC was more correlated with EVI (R² from 0.66 to 0.85) than NDVI (R² from 0.52 to 0.68). In addition, the onset and ending of phenology were shown to differ by 3.5 and 5.4 days between EVI and GCC, respectively. Larger differences were detected between NDVI and GCC, 17.05 and 26.89 days for the onset and ending, respectively. EVI showed better estimations of the phenological dates than NDVI. This better performance is explained by the higher spectral sensitivity of EVI for multiple canopy leaf layers due to the presence of an additional blue band and an optimized soil factor value. Our study demonstrates that the phenological observations derived from PhenoCam are comparable with the EVI index. We conclude that EVI is more suitable than NDVI to assess phenology in evergreen species of the northern boreal region, where PhenoCam data are not available. The EVI index could be used as a reliable proxy of GCC for monitoring evergreen species phenology in areas with reduced access, or where repeated data collection from remote areas are logistically difficult due to the extreme weather.

The variability of tree stem phenology plays a critical role in determining the productivity of forest ecosystems. Therefore, we aim to identify the relationships between the timings of cambium phenology, and forest growth in terms of tree-ring width over a long-term scale. A meta-analysis was performed that combined the timings of xylem formation, which were calculated by a tree-ring formation model of the VS (Vaganov-Shashkin)-oscilloscope during the period 1960–2014, and a tree-ring width series at 20 composite sites on the Tibetan Plateau. Both the start and length of the growing season significantly affected the formation of wood at 70% of the 20 composite sites within the study region. A wider tree ring probably resulted from an earlier start and a longer duration of the growing season. The influence of ending dates on tree-ring width was less evident, and more site-dependent. Weak relationships were identified between the start and end of the growing season at 85% of the composite sites. Compared to the monitoring results, which could only detect the relationships between cambium phenology and xylem cell production from a limited number of trees and years, our long-term relationships deepened such connections, and therefore should be used to improve mechanism models for the accurate evaluating and predicting of wood production and carbon sequestration in forest ecosystems under current and future climate change.

Although cambial reactivation is considered to be strongly dependent on temperature, the importance of water availability at the onset of xylogenesis in semi-arid regions still lacks sufficient evidences. In order to explore how environmental factors influence the initiation of cambial activity and wood formation, we monitored weekly cambial phenology in Qilian juniper (Juniperus przewalskii) from a semi-arid high-elevation region of northwestern China. We collected microcores from 12 trees at two elevations during the growing seasons in 2013 and 2014, testing the hypothesis that rainfall limits cambial reactivation in spring. Cambium was reactivated from late April to mid-May, and completed cell division from late July to early August, lasting 70–100 days. Both sites suffered from severe drought from January to April 2013, receiving < 1 mm of rain in April. In contrast, rainfall from January to April 2014 was 5–6 times higher than that in 2013. However, cambial reactivation in 2014 was delayed by 10 days. In spring, soil moisture gradually increased with warming temperatures, reaching 0.15 m³/m³ before the onset of xylogenesis, which may have ensured water availability for tree growth during the rainless period. We were unable to confirm the hypothesis that rainfall is a limiting factor of cambial reactivation. Our results highlight the importance of soil moisture in semi-arid regions, which better describe the environmental conditions that are favorable for cambial reactivation in water-limited ecosystems.