Cambial Phenology Research Articles

Cambial phenology in Juniperus przewalskii along different altitudinal gradients in a cold and arid region.

Knowing more precisely the cambial phenology and wood formation dynamics of trees can lead to a better understanding on how trees react to short-term changes in environmental conditions. Such an understanding could also shed light on the physiological foundation of climate-growth interactions at a regional scale. Although it has been documented that temperature is an important factor determining the cambial phenology in cold and humid climates, there is less agreement on the driver(s) that trigger the onset and end of wood formation in cold and arid climates. Here, the phenological traits of cambial activity and xylem formation were analyzed biweekly along an altitudinal transect ranging from 3580 to 3980 m above sea level, a transect that covers the distribution of Qilian juniper (Juniperus przewalskii Kom.) along a slope of the Tibetan Plateau. Cambial phenology and the duration and rate of wood formation were assessed from anatomical observations during the growing season of the developing xylem obtained from microcores collected from the stem of 10 trees total in 2012 (five at two altitudes each) and 25 trees (five at five altitudes each) in 2013. We found that the onset of wood formation was significantly correlated with altitude in 2013, with onset beginning 8.2 days earlier with every 100 m decrease in elevation. The change in onset with elevation corresponds to a change of 14.1 days °C-1 when adjusted for the monitored altitudinal lapse rate of -0.58 °C per 100 m. The duration of wood formation lasted from mid-May to mid-August, with the length of the 2013 growing season decreasing from 97 to 65 days from low to high elevation. Although the end of growing season appeared minimally related to altitude during both growing seasons, differences in end of wood production and wood formation between the two growing seasons were significant. It appears that summer drought conditions constricted the end of growing season across all elevations along our transect in 2013. Sensitivity analysis found xylem growth was positively correlated with rate and duration of wood production, with the former explaining most variability in growth. Our findings provide new data on the timing and duration of wood formation and help quantify the potential impacts of global warming on tree growth and productivity in cold and arid regions.

Xylem and Leaf Functional Adjustments to Drought in Pinus sylvestris and Quercus pyrenaica at Their Elevational Boundary.

Climatic scenarios for the Mediterranean region forecast increasing frequency and intensity of drought events. Consequently, a reduction in Pinus sylvestris L. distribution range is projected within the region, with this species being outcompeted at lower elevations by more drought-tolerant taxa such as Quercus pyrenaica Willd. The functional response of these species to the projected shifts in water availability will partially determine their performance and, thus, their competitive success under these changing climatic conditions. We studied how the cambial and leaf phenology and xylem anatomy of these two species responded to a 3-year rainfall exclusion experiment set at their elevational boundary in Central Spain. Additionally, P. sylvestris leaf gas exchange, water potential and carbon isotope content response to the treatment were measured. Likewise, we assessed inter-annual variability in the studied functional traits under control and rainfall exclusion conditions. Prolonged exposure to drier conditions did not affect the onset of xylogenesis in either of the studied species, whereas xylem formation ceased 1–3 weeks earlier in P. sylvestris. The rainfall exclusion had, however, no effect on leaf phenology on either species, which suggests that cambial phenology is more sensitive to drought than leaf phenology. P. sylvestris formed fewer, but larger tracheids under dry conditions and reduced the proportion of latewood in the tree ring. On the other hand, Q. pyrenaica did not suffer earlywood hydraulic diameter changes under rainfall exclusion, but experienced a cumulative reduction in latewood width, which could ultimately challenge its hydraulic performance. The phenological and anatomical response of the studied species to drought is consistent with a shift in resource allocation under drought stress from xylem to other sinks. Additionally, the tighter stomatal control and higher intrinsic water use efficiency observed in drought-stressed P. sylvestris may eventually limit carbon uptake in this species. Our results suggest that both species are potentially vulnerable to the forecasted increase in drought stress, although P. sylvestris might experience a higher risk of drought-induced decline at its low elevational limit.

Synchronisms between bud and cambium phenology in black spruce: early-flushing provenances exhibit early xylem formation.

Bud and cambial phenology represent the adaptation of species to the local environment that allows the growing season to be maximized while minimizing the risk of frost for the developing tissues. The temporal relationship between the apical and radial meristems can help in the understanding of tree growth as a whole process. The aim of this study was to compare cambial phenology in black spruce (Picea mariana (Mill.) B.S.P.) provenances classified as early and late bud flushing. The different phases of cambial phenology were assessed on wood microcores sampled weekly from April to October in 2014 and 2015 from 61 trees growing in a provenance trial in Quebec, Canada. Trees showing an early bud flush also exhibited early reactivation of xylem differentiation, although an average difference of 12 days for buds corresponded to small although significant differences of 4 days for xylem. Provenances with early bud flush had an early bud set and completed xylem formation earlier than late bud flush provenances. No significant difference in the period of xylem formation and total growth was observed between the flushing classes. Our results demonstrate that the ecotype differentiation of black spruce provenances represented by the phenological adaptation of buds to the local climate corresponds to specific growth dynamics of the xylem.

Intra-annual stem growth dynamics of Lebanon Cedar along climatic gradients

Onset of cambial activity in Lebanon Cedar is triggered by stem temperature but may be delayed by high site-LAI. Higher growth rates and tree-ring widths were observed under better water and light availability. Daily stem radius variations were sensitive to humid conditions. Studies on intra-annual dynamics of stem growth provide useful information on tree growth responses to environmental conditions, but are fragmentary for species from Mediterranean Mountain ranges. Lebanon Cedar is a frost and drought tolerant species, growing between 1000 and 2000 m a.s.l in the Taurus Mountains (Turkey). Foresters see it as a potential candidate for plantation in Central European forests facing global warming. To describe the natural variability of Lebanon Cedar growth dynamics, five study sites were established: four along an altitudinal gradient at a natural site in SW-Turkey and one in a Lebanon Cedar plantation in Central Germany. Two stem growth monitoring methods were used: (1) bi-weekly microcoring during 2013 growing season and (2) point dendrometers during 2013 and 2014. Histological analyses were used to calibrate dendrometer records and to describe cambium phenology. Seasonal dynamics of xylem and stem radial increments were modelled by Gompertz functions. Onset of cambial activity was observed 1–2 weeks after stem temperatures reached a threshold of 5 °C but could be delayed by high site-LAI. Cedars growing under better light and water availability showed higher growth rates and wider tree rings. Daily stem radius variations (dSRV) extracted from dendrometer records were negatively related to vapor pressure deficit and global radiation; multiple linear regressions explained 30–52% of dSRV variance being dominated by relative humidity, precipitation, and soil water content. Best growth performance was observed at the German site, likely for a continuous water supply throughout the year, underlining the potential of Lebanon Cedar for Central European Forestry.

Deep Soil Conditions Make Mediterranean Cork Oak Stem Growth Vulnerable to Autumnal Rainfall Decline in Tunisia

Tree rings provide fruitful information on climate features driving annual forest growth through statistical correlations between annual tree growth and climate features. Indices built upon tree growth limitation by carbon sequestration (source hypothesis) or drought-driven cambial phenology (sink hypothesis) can be used to better identify underlying processes. We used both analytical frameworks on Quercus suber, a sparsely studied species due to tree ring methodological issues, and growing on a favorable sub-humid Mediterranean climate and deep soil conditions in Tunisia (North Africa). Statistical analysis revealed the major role of autumnal rainfall before the growing season on annual tree growth over the 1918–2008 time series. Using a water budget model, we were able to explain the critical role of the deep soil water refill during the wet season in affecting both the drought onset controlling growth phenology and the summer drought intensity affecting carbon assimilation. Analysis of recent climate changes in the region additionally illustrated an increase in temperatures enhancing the evaporative demand and advancing growth start, and a decline in rainfalls in autumn, two key variables driving stem growth. We concluded on the benefits of using process-based indices in dendrochronological analysis and identified the main vulnerability of this Mediterranean forest to autumnal rainfall decline, a peculiar aspect of climate change under summer-dry climates.

Growth dynamics of Centrolobium robustum (Vell.) Mart. ex Benth. (Leguminosae-Papilionoideae) in the Atlantic Forest

Knowledge about growth rhythm and longevity of tropical trees is essential to understand life strategies of native species, and consequently, of forest remnants. This study aims to investigate the growth dynamics of Centrolobium robustum, a representative species from Atlantic Forest. For this purpose, radial growth was examined through tree rings and cambium analysis, from macroscopical, anatomical, and ultrastructural perspectives. The apical and radial stem growth was evaluated through leaf and cambium phenology, respectively. Leaf and cambium behaviors were associated with environmental seasonality. The results showed that the period of cell production and cambial development occurred in association with leaf expansion and during the wet, warm, and long-day season. Seasonality of cambial and leaf phenology in response to the environment determined the annual formation of tree rings, and it allowed to evaluate C. robustum growth rates and longevity. Species seasonality also responded to abnormal climatic events during the study period, which shows its sensitivity to environmental changes and represents a factor to be considered in the forest management programs.

Tree growth, cambial phenology, and wood anatomy of limber pine at a Great Basin (USA) mountain observatory

Anatomical features of Pinus flexilis under warmer and drier conditions along an altitudinal transect revealed a shorter growing season and shifts in the timing of wood formation. Future climate change driven by greenhouse warming is expected to increase both frequency and severity of drought events and heat waves. Possible consequences for forest ecosystems include changes in foundation species and extended die-off phenomena. We investigated tree growth under the set of biotic and abiotic conditions, and their interactions, that are expected in a drier and warmer world using mountain observatories designed to capture elevation gradients in the Great Basin of North America. Stem cambial activity, wood anatomy, and radial growth of limber pine (Pinus flexilis) were examined at two different elevations using automated dendrometers and repeated histological microcores in 2013–2014. Mean annual temperature was 3.7° cooler at the higher site, which received 170 mm year−1 of precipitation more than the lower site. Mean air temperature thresholds for xylogenesis computed using logistic regression were 7.7 and 12.0 °C at the higher and lower site, respectively. No differences in the onset date of cambial activity were found under such naturally contrasted conditions, with the global change analog provided by the lower site. Growing season was shortened by increasing drought stress at the lower site, thereby reducing xylem production. Stem expansion was only detectable by automated dendrometers at the higher site. Using elevation to simulate climatic changes and their realized ecosystem feedbacks, it was possible to express tree responses in terms of xylem phenology and anatomical adaptations.

Environmental drivers of cambial phenology in Great Basin bristlecone pine.

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.

MAPPING EVENTS: CAMBIUM PHENOLOGY ACROSS THE LATITUDINAL DISTRIBUTION OF BLACK SPRUCE

This study used statistical models for describing the spatial patterns of variation in cambium phenology and xylem cell production across the entire latitudinal distribution of a species. The studied area extends over 600 km from the 48th to the 53rd parallel in the boreal forest of Quebec, Canada. Microcores were collected weekly from April to October 2012 from 50 Black spruce [Picea mariana (Mill.) BSP] trees in five stands. The dates of occurrence of the phases of cambium phenology were identified on histological sections and correlated to the latitude and altitude of the sites by means of linear and non-linear functions. The results were used to estimate the timings of xylem growth and cell production across the sampled region. Phenology was mostly represented by linear functions. The increase in latitude and altitude produced a proportional variation in the beginning and ending of xylem differentiation, thus leading to a shorter length of the period of wood formation. The phase of cell enlargement and cell production changed according to a non-linear pattern represented by a negative exponential curve. Latitude was the factor with the greatest impact on xylem phenology, while altitude had a slight or no effect, especially for nonlinear relationships. Xylem formation is a complex process composed of several phenological phases that change across a species distribution area according to either linear or non-linear patterns. Knowledge and quantification of these patterns are important for modelling the dynamics of tree growth across wide geographical areas and for predicting productivity of forest ecosystems under climate change scenarios.

Synchronisms and correlations of spring phenology between apical and lateral meristems in two boreal conifers

Phenological synchronisms between apical and lateral meristems could clarify some aspects related to the physiological relationships among the different organs of trees. This study correlated the phenological phases of bud development and xylem differentiation during spring 2010-14 in balsam fir (Abies balsamea Mill.) and black spruce [(Picea mariana Mill. (BSP)] of the Monts-Valin National Park (Quebec, Canada) by testing the hypothesis that bud development occurs after the reactivation of xylem growth. From May to September, we conducted weekly monitoring of xylem differentiation using microcores and bud development with direct observations on terminal branches. Synchronism between the beginning of bud development and xylem differentiation was found in both species with significant correlations between the phases of bud and xylem phenology. Degree-day sum was more appropriate in assessing the date of bud growth resumption, while thermal thresholds were more suitable for cambium phenology. Our results provide new knowledge on the dynamics of spring phenology and novel information on the synchronisms between two meristems in coniferous trees. The study demonstrates the importance of precisely defining the phases of bud development in order to correctly analyse the relationships with xylem phenology.

Seasonal variations in phenological traits: leaf shedding and cambial activity in Parkia nitida Miq. and Parkia velutina Benoist (Fabaceae) in tropical rainforest

Studies on the periodicity of wood formation provide essential data on tree age and factors related to the control of tree growth. The aim of this work was to investigate the cambial phenology and its relation with leaf phenology and climatic seasonality in two tropical rainforest species belonging to the genus Parkia known for having a contrasted leaf phenology. From wood samples collected at 15 day intervals from April 2009 to February 2012 in five trees of both species, we quantified cambial activity by counting layers of cells in cambial zone, enlarging cells zone and wall thickening cells zone from wood prepared anatomical micro sections. In the same time, we observed leaf shedding behaviors in the crown of the studied trees. In both species, cambial activity was significantly reduced during leafless period. However, the cambial activity of P. velutina was not affected by seasonality while in P. nitida, we found a significant positive effect of the dry season on cambial activity. This diminution of activity associated to leaf shedding occurred annually during the dry season in P. nitida whereas the trees were not synchronous in time for P. velutina. On the contrary that is often admitted, the rest cambial period can be independent of environmental restriction such as dry season. Leaf phenology remains the best proxy to estimate cambial activity.

Lengthening of the duration of xylogenesis engenders disproportionate increases in xylem production

In cold climates, the expected global warming will lead to earlier cambial resumptions in spring, with a resultant lengthening of the growing season but unknown consequences on forest productivity. The phenological traits of cambium activity and xylem formation were analyzed at a short time scale along a thermal gradient represented by an alti-latitudinal range from the 48th to 53rd parallels and covering the whole closed black-spruce [Picea mariana (Mill.) BSP] forest in Quebec, Canada. A hypothesis was tested that warmer temperatures influence cambium phenology, allowing longer duration and higher intensity of growth, and resulting in proportionally increased xylem production. From April to October 2012, cell division in cambium and post-cambial differentiation of xylem were observed on anatomical sections obtained from microcores collected weekly from the stem of fifty trees. The southern and warmer site was characterized by the highest radial growth, which corresponded to both the highest rates and longest durations of cell production. The differences in terms of xylem phenology and growth were marginal between the other sites. Xylem growth was positively correlated with rate and duration of cell production, with the latter explaining most variability in growth. Within the range analyzed, the relationship between temperature and most phenological phases of xylogenesis was linear. On the contrary, temperature was related with cell production according to an exponential pattern. Periods of xylogenesis of 14 days longer (+13.1%) corresponded to a massive increase in cell production (33 cells, +109%). This disproportionate change occurred at a May-September average temperature of ca. 14 °C and a snow-free period of 210-235 days. At the lower boundary of the distribution of black spruce, small environmental changes allowing marginal lengthening of the period of cell division could potentially lead to disproportionate increases in xylem cell production, with substantial consequences for the productivity of this boreal species.

Intra-annual stem diameter growth of Tamarix ramosissima and association with hydroclimatic factors in the lower reaches of China’s Heihe River

High-resolution observations of cambial phenology and intra-annual growth dynamics are useful approaches for understanding the response of tree growth to climate and environmental change. During the past two decades, rapid socioeconomic development has increased the demand for water resources in the oases of the middle reaches of the Heihe River in northwestern China, and the lower reaches of the Heihe River have changed from a perennial river to an ephemeral stream with a decreased and degraded riparian zone. Tamarisk (Tamarix ramosissima) is the dominant shrub species of the desert riparian forest. In this study, the daily and seasonal patterns of tamarisk stem diameter growth, including the main period of tree ring formation, were examined. Observations concerning the driving forces of growth changes, along with implications for the ecology of the dendrohydrological area and management of desert riparian forests in similar arid regions, are also presented. The diurnal-seasonal activity of stem diameter and the dynamics of growth ring formation were studied using a point dendrometer and micro-coring methods during the 2012 growing season in shrub tamarisk in a desert riparian forest stand in the lower reaches of the Heihe River in Ejin Banner, Inner Mongolia of northwestern China. Generally, the variation in diurnal diameter of tamarisk was characterized by an unstable multi-peak pattern, with the cumulative stem diameter growth over the growing season following an S-shaped curve. The period from late May to early August was the main period of stem diameter growth and growth-ring formation. Among all of the hydroclimatic factors considered in this study, only groundwater depth was significantly correlated with stem diameter increment during this period. Therefore, for the dendrochronological study, the annual rings of the tamarisk can be used to reconstruct processes that determine the regional water regime, such as river runoff and fluctuations in groundwater depth. For the management of desert riparian forests, suitable groundwater depths must be maintained in the spring and summer to sustain tree health and a suitable stand structure.

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

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. 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. 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. 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 conditions.

Xylogenesis of Pinus pinaster under a Mediterranean climate

Context The knowledge on cambial activity in water-limited environments, such as the Mediterranean, is still fragmentary. Dendrochronological studies have determined that spring precipitation plays an important part in determining tree-ring width and the properties of tracheids. However, the complex relation between cambial phenology and climate is still far from understood.

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.

Intra-annual dynamics of non-structural carbohydrates in the cambium of mature conifer trees reflects radial growth demands

The presence of soluble carbohydrates in the cambial zone, either from sugars recently produced during photosynthesis or from starch remobilized from storage organs, is necessary for radial tree growth. However, considerable uncertainties on carbohydrate dynamics and the consequences on tree productivity exist. This study aims to better understand the variation in different carbon pools at intra-annual resolution by quantifying how cambial zone sugar and starch concentrations fluctuate over the season and in relation to cambial phenology. A comparison between two physiologically different species growing at the same site, i.e., the evergreen Picea abies Karst. and the deciduous Larix decidua Mill., and between L. decidua from two contrasting elevations, is presented to identify mechanisms of growth limitation. Results indicate that the annual cycle of sugar concentration within the cambial zone is coupled to the process of wood formation. The highest sugar concentration is observed when the number of cells in secondary wall formation and lignification stages is at a maximum, subsequent to most radial growth. Starch disappears in winter, while other freeze-resistant non-structural carbohydrates (NSCs) increase. Slight differences in NSC concentration between species are consistent with the differing climate sensitivity of the evergreen and deciduous species investigated. The general absence of differences between elevations suggests that the cambial activity of trees growing at the treeline was not limited by the availability of carbohydrates at the cambial zone but instead by environmental controls on the growing season duration.

Climatic control of tracheid production of black spruce in dense mesic stands of eastern Canada

Inferences on climate change effects are reliable only if they are based on a causal relationship rather than simple statistical predictive capacity. To assess for causal links between climate and mature black spruce (Picea mariana (Mills.) BSP) radial growth, we combined the use of wood anatomy, cambium phenology, climate and soil measurements (air temperature and humidity, precipitations, soil temperature and water content, photosynthetically active radiation), and a model selection approach proceeding backwards from a full model. Results show that the number of tracheids is responsible for 88% of the variation in ring width whereas mean tracheid diameter accounts for the remaining 12%. The number of tracheids produced depends on factors related to photosynthesis during tracheid production, i.e., daily light intensity and maximum temperature between the day of initiation and the day of cessation of tracheid production, plus soil temperature during August of the previous year which is an important period for determining the number of new needles produced. It is also important to consider duration of the period for tracheid production. These results imply that short-term climate change should increase black spruce radial growth. They also suggest that the typical use of post-growth ring width sampling individually linked to air temperature and precipitations is not sufficient to infer climate change effects accurately on radial growth where there is no strong single climatic limitation but multiple limitations instead.

Impact of Drought on the Intra-Annual Growth of Pinus sylvestris L.

Climatic scenarios for the Mediterranean Region forecast increasing frequency and intensity of drought events. We analyzed how Pinus sylvestris modifies its cambial phenology and intra-annual growth under enhanced drought conditions. For this purpose, a rainfall exclusion experiment was applied in 2012 to trees located in the lower elevation limit of a mountain forest in Central Spain. Preliminary results show a significantly longer growing season in small trees than in big trees, regardless of the treatment. The tracheid enlargement period was longer in control trees than in trees subjected to rainfall exclusion regardless the size class. Cambial activity and gas exchange rates of Pinus sylvestris responded immediately to drier conditions, which could indicate vulnerability to drought enhancement. Stronger differences between treatments are expected for the second year of the experiment due to the cumulative effect of drought in the rainfall exclusion plots.

Duration of xylogenesis in black spruce lengthened between 1950 and 2010

Reconstructions have identified the 20th century as being uniquely warm in the last 1000 years. Changes in the phenology of primary meristems converged toward increases in length of the growing season. Has the phenology of secondary meristem changed during the last century, and to what extent? Timings of wood formation in black spruce, Picea mariana, were monitored for 9 years on a weekly timescale at four sites in the boreal forest of Quebec, Canada. Models for assessing xylem phenology were defined and applied to reconstruct onset, ending and duration of xylogenesis between 1950 and 2010 using thermal thresholds on chronologies of maximum and minimum temperatures. All sites exhibited increasing trends of both annual and May-September temperatures, with the greatest changes observed at the higher latitudes. Phenological events in spring were more affected than those occurring in autumn, with cambial resumptions occurring 0·5-0·8 d decade(-1) earlier. The duration of xylogenesis has lengthened significantly since 1950, although the models supplied wide ranges of variations, between 0·07 and 1·5 d decade(-1), respectively. The estimated changes in past cambial phenology demonstrated the marked effects of the recent increase in temperature on the phenological traits of secondary meristems. In the long run, the advancement of cambial activity could modify the short time window for growth of boreal species and dramatically affect the dynamics and productivity of trees in these temperature-limited ecosystems.