What role for photoperiod in the bud burst phenology of European beech

A considerable number of studies have investigated the phenology of European beech using models, experimental controlled conditions, or descriptive surveys of patterns in situ. In spite of this interest, there is no consensus about the environmental factors controlling bud burst in beech, especially about the role of photoperiod and chilling temperature (cold temperature effective to release bud dormancy). However, recent experimental and modelling studies provide new insights into the means by which these environmental factors control beech phenology. This present contribution aims to reconcile contradictory hypotheses about the main environmental factors controlling bud burst date of European beech. First, we review the main published results on the environmental control of beech phenology both in controlled and in natural conditions. Second, supported by the findings of recent studies, we propose a new theory for the role of photoperiod during the chilling phase for explaining spatial and temporal variations in bud burst phenology of European beech. Examples using long-term data from the Swiss Alps and Germany are presented to support this theory. The possible impacts of future and ongoing climate warming on beech phenology are discussed. Finally, due to interactions between chilling, forcing temperature, and photoperiod, we assert that beech phenology follows a nonlinear trend across biogeographical gradients such as changes in elevation or latitude and that the bud burst date of beech is expected not to undergo significant changes in response to global warming, especially in warmer climate

Fast acclimation of freezing resistance suggests no influence of winter minimum temperature on the range limit of European beech

Low temperature extremes drive species distribution at a global scale. Here, we assessed the acclimation potential of freezing resistance in European beech ( Fagus sylvatica L.) during winter. We specifically asked (i) how do beech populations growing in contrasting climates differ in their maximum freezing resistance, (ii) do differences result from genetic differentiation or phenotypic plasticity to preceding temperatures and (iii) is beech at risk of freezing damage in winter across its distribution range. We investigated the genetic and environmental components of freezing resistance in buds of adult beech trees from three different populations along a natural large temperature gradient in north-western Switzerland, including the site holding the cold temperature record in Switzerland. Freezing resistance of leaf primordia in buds varied significantly among populations, with LT 50 values (lethal temperature for 50% of samples) ranging from −25 to −40 °C, correlating with midwinter temperatures of the site of origin. Cambial meristems and the pith of shoots showed high freezing resistance in all three populations, with only a trend to lower freezing resistance at the warmer site. After hardening samples at −6 °C for 5 days, freezing resistance of leaf primordia increased in all provenances by up to 4.5 K. After additional hardening at −15 °C for 3 days, all leaf primordia were freezing resistant to −40 °C. We demonstrate that freezing resistance of F. sylvatica has a high ability to acclimate to temperature changes in winter, whereas the genetic differentiation of freezing resistance among populations seems negligible over this small geographic scale but large climatic gradient. In contrast to the assumption made in most of the species distribution models, we suggest that absolute minimum temperature in winter is unlikely to shape the cold range limit of beech. We conclude that the rapid acclimation of freezing resistance to winter temperatures allows beech to track changing climatic conditions, especially during unusually warm winters interrupted by very cold weather

Daily Maximum Temperatures Induce Lagged Effects on Leaf Unfolding in Temperate Woody Species Across Large Elevational Gradients

The timing of leaf unfolding in temperate woody species is predominantly controlled by the seasonal course of temperature in late winter and early spring. However, quantifying lagged temperature effects on spring phenology is still challenging. Here, we aimed at investigating lagged and potentially non-linear effects of daily maximum temperatures on the probability of leaf unfolding in temperate woody species growing across large elevational gradients. We analyzed 5280 observations of leaf-out time of four tree species (European beech, horse chestnut, European larch, Norway spruce) and one shrub species (common hazel) that were recorded by volunteers over 40 years at 42 locations in Switzerland. We used a case-crossover sampling design to match leaf-out dates with control dates (i.e., dates before or after leaf-out), and analyzed these data with conditional logistic regression accounting for lagged temperature effects over 60 days. Multivariate meta-analyses were used to synthesize lagged temperature and elevational effects on leaf unfolding across multiple phenological stations. Temperature effects on the probability of leaf unfolding were largest at relatively short lags (i.e., within ca. 10 days) and decreased with increasing lags. Short- to mid-term effects (i.e., within ca. 10 to 20 days) were larger for late-leafing species known to be photoperiod-sensitive (beech, Norway spruce). Temperature effects increased for the broadleaved species (horse chestnut, hazel, beech) with decreasing elevation, particularly within ca. 10 to 40 days, i.e., leaf unfolding occurs more rapidly at low elevations for a given daily maximum temperature. Our novel findings provide evidence of cumulative and long-term temperature effects on leaf unfolding, whereby the efficiency of relatively high temperatures to trigger leaf-out becomes higher shortly before bud burst. These lagged associations between temperature and leaf unfolding improve our understanding of phenological responses across temperate woody species with differing ecological requirements that occur along elevational gradients

Elevational adaptation and plasticity in seedling phenology of temperate deciduous tree species

Phenological events, such as the initiation and the end of seasonal growth, are thought to be under strong evolutionary control because of their influence on tree fitness. Although numerous studies highlighted genetic differentiation in phenology among populations from contrasting climates, it remains unclear whether local adaptation could restrict phenological plasticity in response to current warming. Seedling populations of seven deciduous tree species from high and low elevations in the Swiss Alps were investigated in eight common gardens located along two elevational gradients from 400 to 1,700m. We addressed the following questions: are there genetic differentiations in phenology between populations from low and high elevations, and are populations from the upper elevational limit of a species' distribution able to respond to increasing temperature to the same extent as low-elevation populations? Genetic variation of leaf unfolding date between seedlings from low and high populations was detected in six out of seven tree species. Except for beech, populations from high elevations tended to flush later than populations from low elevations, emphasizing that phenology is likely to be under evolutionary pressure. Furthermore, seedlings from high elevation exhibited lower phenological plasticity to temperature than low-elevation provenances. This difference in phenological plasticity may reflect the opposing selective forces involved (i.e. a trade-off between maximizing growing season length and avoiding frost damages). Nevertheless, environmental effects were much stronger than genetic effects, suggesting a high phenological plasticity to enable tree populations to track ongoing climate change, which includes the risk of tracking unusually warm springs followed by fros

Coordination between growth, phenology and carbon storage in three coexisting deciduous tree species in a temperate forest

In deciduous trees growing in temperate forests, bud break and growth in spring must rely on intrinsic carbon (C) reserves. Yet it is unclear whether growth and C storage occur simultaneously, and whether starch C in branches is sufficient for refoliation. To test in situ the relationships between growth, phenology and C utilization, we monitored stem growth, leaf phenology and stem and branch nonstructural carbohydrate (NSC) dynamics in three deciduous species: Carpinus betulus L., Fagus sylvatica L. and Quercus petraea (Matt.) Liebl. To quantify the role of NSC in C investment into growth, a C balance approach was applied. Across the three species, >95% of branchlet starch was consumed during bud break, confirming the importance of C reserves for refoliation in spring. The C balance calculation showed that 90% of the C investment in foliage (7.0–10.5 kg tree −1 and 5–17 times the C needed for annual stem growth) was explained by simultaneous branchlet starch degradation. Carbon reserves were recovered sooner than expected, after leaf expansion, in parallel with stem growth. Carpinus had earlier leaf phenology (by ∼25 days) but delayed cambial growth (by ∼15 days) than Fagus and Quercus , the result of a competitive strategy to flush early, while having lower NSC levels

Vapor-pressure deficit and extreme climatic variables limit tree growth

Assessing the effect of global warming on forest growth requires a better understanding of species-specific responses to climate change conditions. Norway spruce and European beech are among the dominant tree species in Europe and are largely used by the timber industry. Their sensitivity to changes in climate and extreme climatic events, however, endangers their future sustainability. Identifying the key climatic factors limiting their growth and survival is therefore crucial for assessing the responses of these two species to ongoing climate change. We studied the vulnerability of beech and spruce to warmer and drier conditions by transplanting saplings from the top to the bottom of an elevational gradient in the Jura Mountains in Switzerland. We (1) demonstrated that a longer growing season due to warming could not fully account for the positive growth responses, and the positive effect on sapling productivity was species-dependent, (2) demonstrated that the contrasting growth responses of beech and spruce were mainly due to different sensitivities to elevated vapor-pressure deficits (VPD), (3) determined the species-specific limits to VPD above which growth rate began to decline, and (4) demonstrated that models incorporating extreme climatic events could account for the response of growth to warming better than models using only average values. These results support that the sustainability of forest trees in the coming decades will depend on how extreme climatic events will change, irrespective of the overall warming trend

Daylength helps temperate deciduous trees to leaf-out at the optimal time

Global warming has led to substantially earlier spring leaf-out in temperate-zone deciduous trees. The interactive effects of temperature and daylength underlying this warming response remain unclear. However, they need to be accurately represented by earth system models to improve projections of the carbon and energy balances of temperate forests and the associated feedbacks to the Earth's climate system. We studied the control of leaf-out by daylength and temperature using data from six tree species across 2,377 European phenological network (www.pep725.eu), each with at least 30 years of observations. We found that, in addition to and independent of the known effect of chilling, daylength correlates negatively with the heat requirement for leaf-out in all studied species. In warm springs when leaf-out is early, days are short and the heat requirement is higher than in an average spring, which mitigates the warming-induced advancement of leaf-out and protects the tree against precocious leaf-out and the associated risks of late frosts. In contrast, longer-than-average daylength (in cold springs when leaf-out is late) reduces the heat requirement for leaf-out, ensuring that trees do not leaf-out too late and miss out on large amounts of solar energy. These results provide the first large-scale empirical evidence of a widespread daylength effect on the temperature sensitivity of leaf-out phenology in temperate deciduous trees

How Do Climate Change Experiments Alter Plot-Scale Climate?

To understand and forecast biological responses to climate change, scientists frequently use field experiments that alter temperature and precipitation. Climate manipulations can manifest in complex ways, however, challenging interpretations of biological responses. We reviewed publications to compile a database of daily plot-scale climate data from 15 active-warming experiments. We find that the common practices of analysing treatments as mean or categorical changes (e.g. warmed vs.unwarmed) masks important variation in treatment effects over space and time. Our synthesis showed that measured mean warming, in plots with the same target warming within a study, differed by up to 1.6 Celsius degrees (63% of target), on average, across six studies with blocked designs. Variation was high across sites and designs: for example, plots differed by 1.1Celsius degrees (47% of target) on average, for infrared studies with feedback control (n = 3) vs. by 2.2 Celsius degrees (80% of target) on average for infrared with constant wattage designs (n = 2). Warming treatments produce non-temperature effects as well, such as soil drying. The combination of these direct and indirect effects is complex and can have important biological consequences. With a case study of plant phenology across five experiments in our database, we show how accounting for drier soils with warming tripled the estimated sensitivity of budburst to temperature. We provide recommendations for future analyses, experimental design,and data sharing to improve our mechanistic understanding from climate change experiments, and thus their utility to accurately forecast species' responses

Long-term soil water limitation and previous tree vigor drive local variability of drought-induced crown dieback in Fagus sylvatica.

Ongoing climate warming is increasing evapotranspiration, a process that reduces plant-available water and aggravates the impact of extreme droughts during the growing season. Such an exceptional hot drought occurred in Central Europe in 2018 and caused widespread defoliation in mid-summer in European beech (Fagus sylvatica L.) forests. Here, we recorded crown damage in 2021 in nine mature even-aged beech-dominated stands in northwestern Switzerland along a crown damage severity gradient (low, medium, high) and analyzed tree-ring widths of 21 mature trees per stand. We aimed at identifying predisposing factors responsible for differences in crown damage across and within stands such as tree growth characteristics (average growth rates and year-to-year variability) and site-level variables (mean canopy height, soil properties). We found that stand-level crown damage severity was strongly related to soil water availability, inferred from tree canopy height and plant available soil water storage capacity (AWC). Trees were shorter in drier stands, had higher year-to-year variability in radial growth, and showed higher growth sensitivity to moisture conditions of previous late summer than trees growing on soils with sufficient AWC, indicating that radial growth in these forests is principally limited by soil water availability. Within-stand variation of post-drought crown damage corresponded to growth rate and tree size (diameter at breast height, DBH), i.e., smaller and slower-growing trees that face more competition, were associated with increased crown damage after the 2018 drought. These findings point to tree vigor before the extreme 2018 drought (long-term relative growth rate) as an important driver of damage severity within and across stands. Our results suggest that European beech is less likely to be able to cope with future climate change-induced extreme droughts on shallow soils with limited water retention capacity