Impacts of bud set and lammas phenology on root:shoot biomass partitioning and carbon gain physiology in poplar

Key message

Timing of bud set and occurrence of lammas in trees can alter growth partitioning (i.e., root:shoot ratios), while only bud set effectively modifies carbon gain by increasing photosynthesis-related physiological traits.

Abstract

Bud set and lammas (second bud flushing) phenology may strongly influence growth, physiology, and biomass in trees. To test effects of these phenological events, 54 individuals from 16 genotypes of black cottonwood poplar (Populus trichocarpa) were grown in a potted trial under greenhouse conditions (with extended daylengths promoting growth), followed by open-air cultivation (with natural daylengths promoting bud set and/or lammas). Trees were monitored for phenology, repeatedly measured for photosynthesis-related traits, harvested for biomass, and assessed for growth partitioning (separating above- and belowground parts). We grouped trees by phenology for comparisons: (1) trees with early summer bud set, (2) trees with early summer bud set that underwent lammas, (3) trees with late summer bud set (August), and (4) trees with bud set occurring in autumn (September). We found that bud set timing positively affected growth partitioning where earlier bud set resulted in shorter trees with higher root:shoot biomass ratios (by increasing root mass). Lammas growth altered these ratios by significantly increasing shoot growth relative to belowground growth. Trees with bud set occurring in late summer also had higher root:shoot biomass ratios (by increasing root mass) compared to trees setting bud in autumn. Occurrence of bud set coincided with modified physiology of the existing canopy where photosynthesis-related traits were enhanced relative to trees still actively growing. These physiological changes were unaltered by occurrence of lammas. This suggests that bud set prompts a significant, coordinated mechanism of higher carbon gain physiology and belowground biomass accumulation in plants within a “post-bud set” phase.

     

The effect of summer shading on flower bud morphogenesis in apricot (Prunus armeniaca L.)

The aim of this investigation was to assess whether imposed summer shading treatments in apricot (Prunus armeniaca L.) can affect the main phenological phases related to the floral morphogenesis (floral differentiation, xylogenesis), flower bud growth and quality in terms of bud capacity to set fruit. Experimental trials were carried out on fully-grown trees of ‘San Castrese’ and ‘Stark Early Orange’ cultivars characterized by different biological and agronomical traits to which shadings were imposed in July and August. Histological analysis was carried out from summer onwards in order to determine the evolution of floral bud differentiation, and the acropetal progression of primary xylem differentiation along the flower bud axis. Periodical recordings to evaluate the bud drop, blooming time, flowering and fruit set rates were performed also. These shade treatments determined a temporary shutdown of floral differentiation, slowed xylem progression up to the resumption of flower bud growth and a reduced entity of flowering and fruit set. These events were particularly marked in ‘San Castrese’ cultivar, which is well known for its adaptability to different climatic conditions. These findings suggest that adequate light penetration within the canopy during the summer season could be the determining factor when defining the qualitative traits of flower buds and their regular growth, and ultimately to obtain good and constant crops.     

Differences in leaf phenology between juvenile and adult individuals of two tree species in a seasonally dry tropical woodland

In temperate forests, juvenile trees anticipate leaf phenology compared to adults, thus avoiding shading and herbivory. This is also expected to occur in seasonal tropical forests due to intense herbivory and shading during the rainy season; however, the anticipation of leaf phenology by juveniles in seasonal tropical forests has yet to be demonstrated. Stem‐succulent species are expected to be prone to juvenile phenological anticipation because these species are able to use water stored in their stems for leaf flushing in the dry season. We investigated this hypothesis by comparing leaf phenology (bud break, leaf expansion) of juveniles and adults of two species with contrasting wood densities in the transition between dry and rainy seasons in a tropical dry woodland. We also investigated the level of light limitation that juveniles experience in the rainy season. Both species exhibited bud break during the dry season, but only expanded their leaves with the occurrence of the first rains. In general, the stem‐succulent species had a more precocious bud break; however, anticipation by juveniles occurred only in the species with more dense wood. Canopy openness was lower than in temperate deciduous forests, but the fact that the full expansion of leaves occurred only with rainfall indicates that bud break in anticipation of canopy closure contributes only to keeping leaf photosynthetic balance from going negative, and not to higher carbon gain. The importance of anticipated budding for escaping herbivory remains an alternative explanation in need of investigation.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Evolutionary dynamics of the leaf phenological cycle in an oak metapopulation along an elevation gradient

It has been predicted that environmental changes will radically alter the selective pressures on phenological traits. Long‐lived species, such as trees, will be particularly affected, as they may need to undergo major adaptive change over only one or a few generations. The traits describing the annual life cycle of trees are generally highly evolvable, but nothing is known about the strength of their genetic correlations. Tight correlations can impose strong evolutionary constraints, potentially hampering the adaptation of multivariate phenological phenotypes. In this study, we investigated the evolutionary, genetic and environmental components of the timing of leaf unfolding and senescence within an oak metapopulation along an elevation gradient. Population divergence, estimated from in situ and common‐garden data, was compared to expectations under neutral evolution, based on microsatellite markers. This approach made it possible (1) to evaluate the influence of genetic correlation on multivariate local adaptation to elevation and (2) to identify traits probably exposed to past selective pressures due to the colder climate at high elevation. The genetic correlation was positive but very weak, indicating that genetic constraints did not shape the local adaptation pattern for leaf phenology. Both spring and fall (leaf unfolding and senescence, respectively) phenology timings were involved in local adaptation, but leaf unfolding was probably the trait most exposed to climate change‐induced selection. Our data indicated that genetic variation makes a much smaller contribution to adaptation than the considerable plastic variation displayed by a tree during its lifetime. The evolutionary potential of leaf phenology is, therefore, probably not the most critical aspect for short‐term population survival in a changing climate.     

Comparing phenocam color indices with phenological observations of black spruce in the boreal forest

Bud phenology identifies the growing period of trees and determines the pattern of mass and energy exchanges between forest and atmosphere over time and space. Canopy color metrics derived from phenocams have been widely used to investigate tree phenology. However, it remains unclear which color-based index better tracks the seasonal variations of tree phenology in evergreen forest ecosystems. Herein, we compared four color metrics (red chromatic coordinate (RCC), green chromatic coordinate (GCC), vegetation contrast index (VCI) and excess green index (ExG)) derived from phenocam images with bud phenological phases recorded in black spruce [Picea mariana (Mill.) B.S·P] during 2017–2020 at a boreal forest site in Quebec, Canada. Canopy redness (RCC) and greenness (GCC, ExG, and VCI) showed a bimodal and bell-shaped seasonal pattern, respectively. The phases of bud burst and bud set lasted from end-May to end-June and from mid-July to end-September, respectively. The neural network model indicated that GCC had the best predictive ability in capturing the sequential phases of bud phenology. Bud phenological phases predicted by GCC showed the highest correlation with actual bud phenological phases among four indices, with R² above 0.9 and RMSE lower than 0.5. Overall, color indices performed better when representing bud burst than bud set. Our findings improve the efficiency and confidence of the phenocam greenness index to characterize the growing season of evergreen forests.     

暖温带乔木和灌木物候变化及对气候变化的响应

根据2003-2014年气象数据和暖温带3种乔木(辽东栎、五角枫和核桃楸)和3种灌木(土庄绣线菊、毛叶丁香和六道木)的物候观测数据资料, 采用气候倾向率和回归分析等方法, 观察乔木和灌木物候变化特征的差异, 分析温度、降水以及乔木、灌木的物候变化趋势, 同时对气象因子与乔木和灌木物候期的相关关系进行研究。结果表明: ①研究期间, 北京东灵山平均气温呈不显著的上升趋势, 气候倾向率为0.200℃·10a–1, 春季(3–5月)和夏季(6-8月)温度显著上升; 降水量呈下降趋势, 平均减少71.630 mm·10a–1, 总体呈暖、干的趋势。②3种乔木的生长季长度都缩短, 辽东栎、五角枫和核桃楸平均生长季长度分别缩短50.70 d·10 a–1、29.83 d·10a–1和22.36 d·10a–1。3种灌木的生长季长度也都缩短, 土庄绣线菊、毛叶丁香和六道木的平均生长季长度分别缩短42.55 d·10a–1、42.76 d·10a–1和38.15 d·10a–1。乔木和灌木的物候变化趋势相同, 整体表现为春季物候推迟, 秋季物候提前, 生长季长度都缩短且生长季长度相差不大。乔木和灌木都表现出芽期推迟最明显, 每10年推迟达19天以上。③乔木和灌木各物候期与气温总体表现为负相关, 即气温升高, 物候期提前, 其相关性显示出夏季(6-8月)温度对植被物候期影响较大, 夏季温度与各物候期表现为正相关, 即夏季温度升高, 物候期推迟。同时乔木和灌木与总体降水没有明显的相关关系, 但秋季物候与不同时段降水表现不同的相关性, 由此可知夏季温度变化对木本植物春季物候(出芽期、展叶期和首花期)的影响更大, 而秋季物候(叶变色期和落叶期)受温度和降水共同影响。     

北京地区木本春季物候特征与生物学特性的关系

植物物候是植物生活史中的重要性状,也是指示气候与自然环境变化的重要指标,现已成为全球变化领域的研究热点之一。传统物候研究多假设物候由气候因素决定,如气温、降水、光照等,并主要从植物物候的年际变化角度探讨了气候因素对物候特征的影响。然而,不同物种的物候存在较大差异表明植物物候还与自身生物学特性（如系统发育和功能性状）有关,但植物生物学特性如何影响植物物候仍缺乏深入研究。基于北京地区44种木本植物1965-2018年的展叶始期和开花始期观测资料,以展叶始期和开花始期的3类物候特征（平均物候期、物候对温度的响应敏感度和物候期的积温需求）为例,探究植物物候特征与系统发育和功能性状的关系。首先,利用系统发育信号Blomberg’s K和进化模型检验植物物候特征是否具有系统发育保守性,并通过系统发育信号表征曲线直观表达植物物候特征的进化模式;之后,利用广义估计方程分析植物生活型、传粉型与物候特征的关系,以揭示不同植物的资源利用方式及生存策略的差异。研究发现：（1）除展叶始期的温度敏感度外,其余物候特征的进化均受随机遗传漂变和自然选择力的共同作用,可推断物候特征具有系统发育保守性,即亲缘关系越近的物种物候特征越相似。（2）开花始期的系统发育信号强度比展叶始期更大,表明繁殖物候的系统发育可能比生长物候更保守。（3）植物展叶始期及其积温需求与生活型密切相关。灌木比乔木的展叶时间早、积温需求少。植物开花始期与传粉型相关,风媒植物开花显著早于虫媒植物。研究成果有助于深入理解物候变化的生物学机制,对于丰富物候学的理论研究有重要意义,同时对植物保护也具有重要的指导价值。     

Environmental constraints on phenology and internal nutrient cycling in the Mediterranean winter-deciduous shrub Amelanchier ovalis Medicus

The functional adjustments of winter-deciduous perennials to Mediterranean conditions have received little attention. The objectives of this study were: (i) to determine whether Amelanchier ovalis, a winter-deciduous shrub of Mediterranean and sub-Mediterranean regions, has nutritional and phenological traits in common with temperate zone deciduous trees and shrubs and (ii) to determine the constraints of Mediterranean environmental conditions on these traits. Over two years, phenology and nitrogen, and phosphorus concentrations were monitored monthly in the crown of A. ovalis. Leaf longevity, survival and nutrient resorption from senescing leaves were used to infer nutrient use efficiency and retention times of nutrients within the crown. In A. ovalis, bud burst was much earlier than in temperate deciduous trees and shrubs. Most vegetative and reproductive growth occurred in spring. Limited phenological development took place during the summer drought period. Unexpectedly, leaf shedding was very gradual, which might be related to water shortages in summer. Leaf longevity, nutrient resorption from senescing leaves, and maximum leaf nutrient concentrations indicated that nutrient retention times were short and nutrient use efficiency was low compared to that found in temperate deciduous plants and co-occurring Mediterranean evergreens. A. ovalis exhibited phenological development appropriate for a Mediterranean climate, although its limited ability to retain nutrients likely restricts the types of sites that it can occupy.     

胡杨枝芽生长特征及其展叶物候特征

以5个不同发育阶段的胡杨(Populus euphratica Oliv.)个体为研究对象,观测记录了枝芽展叶物候、枝芽生长特征和叶形变化的空间分布规律。结果表明:不同发育阶段的胡杨个体以及同一个体树冠的不同层次,其枝芽生长及其展叶物候期表现出不同的时空特征。随着树龄的增加和树冠层次的增高(由基向顶),当年新生枝条长度、枝条叶片数和叶形指数逐渐减小,但叶面积和叶片干重逐渐增大。5个不同发育阶段胡杨个体均表现出展叶物候始于树冠顶层,依次向下结束于树冠基部;展叶物候期共性表现在枝芽萌动期均在4月上旬,起始展叶期集中在4月中旬,展叶终期则在5月上旬到下旬;树龄较大的个体其枝芽萌动期、起始展叶期、展叶终期较树龄较小的个体早;其枝芽萌动期到展叶终期的时间进程较树龄较小的个体短;不同发育阶段的个体枝芽萌动期出现的时间较为离散,起始展叶期和展叶终期出现的时间较为集中。相关分析表明,出叶周期与枝条长度、枝条叶片数量和叶形指数呈极显著正相关,与叶面积和叶片干重呈显著负相关。     

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 ± 0.3 to 6.6 ± 0.4 days °C⁻¹ (mean ± SD) and a 0 to 5.6 ± 0.6 days °C⁻¹ delay in leaf senescence. Together both changes resulted in a 6.9 ± 1.0 to 13.0 ± 0.7 days °C⁻¹ lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.     

Larger temperature response of autumn leaf senescence than spring leaf‐out phenology

Climate warming is substantially shifting the leaf phenological events of plants, and thereby impacting on their individual fitness and also on the structure and functioning of ecosystems. Previous studies have largely focused on the climate impact on spring phenology, and to date the processes underlying leaf senescence and their associated environmental drivers remain poorly understood. In this study, experiments with temperature gradients imposed during the summer and autumn were conducted on saplings of European beech to explore the temperature responses of leaf senescence. An additional warming experiment during winter enabled us to assess the differences in temperature responses of spring leaf‐out and autumn leaf senescence. We found that warming significantly delayed the dates of leaf senescence both during summer and autumn warming, with similar temperature sensitivities (6–8 days delay per °C warming), suggesting that, in the absence of water and nutrient limitation, temperature may be a dominant factor controlling the leaf senescence in European beech. Interestingly, we found a significantly larger temperature response of autumn leaf senescence than of spring leaf‐out. This suggests a possible larger contribution of delays in autumn senescence, than of the advancement in spring leaf‐out, to extending the growing season under future warmer conditions.     

Timing of photoperiodic competency causes phenological mismatch in balsam poplar (Populus balsamifera L.)

Plant phenology is expected to be sensitive to climate warming. In boreal trees, spring flush is primarily temperature driven, whereas height growth cessation and autumn leaf senescence are predominantly controlled by photoperiod. Cuttings of 525 genotypes from the full range of balsam poplar were planted into two common gardens (Vancouver and Indian Head, Canada) at similar latitudes, but with differing winter temperatures and growing seasons. There was clinal variation in spring and, particularly, summer and fall phenology. Bud flush and, despite milder climate, bud set and leaf drop were earlier at Vancouver than at Indian Head by 44, 28 and 7 d, respectively. Although newly flushed growth is insensitive to photoperiod, many genotypes at both sites became competent before the summer solstice. At Vancouver, high‐latitude genotypes set dormant terminal buds in mid‐spring. Most other genotypes grew until midsummer or set bud temporarily and then experienced a second flush. In both gardens and in a growth chamber experiment, earlier bud set was associated with reduced height growth and higher root/shoot ratios. Shoots attained competency ～5 weeks after flushing, which would normally prevent dormancy induction before the solstice, but may be insufficient if spring advances by more than a few weeks.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Changes in Leaf Phenology are Dependent on Tree Height inAcer mono,a Deciduous Broad-leaved Tree

Ontogenetic changes in leaf phenology of a hardwood tree,Acer mono, were investigated in individuals in different size classes in a temperate forest. Leaf emergence was earliest in current-year seedlings, and was increasingly delayed with increasing height of the individual. The shorter the tree, the longer the duration of leaf emergence. Timing of leaf emergence of the dominant heterospecific canopy trees was almost identical to that of conspecific adults; understorey light then gradually decreased with expansion of canopy leaves. These traits indicate that smaller individuals that receive the least light in summer can acquire favourable light for a longer period in spring than taller plants even in a forest understorey, but the advantage decreases with increasing plant height. Changes in the duration of leaf emergence and leaf longevity in response to environmental light regime [sun (forest edge)vs. shade (forest understorey)], were greatest for current year seedlings but decreased with increasing plant height. These results suggest that the plastic response of leaf phenology in juvenile stages may reduce the risk of losing an entire cohort in spatially heterogenous environments in the understorey of temperate forests.     

Leaf flushing and shedding,bud and flower production,and stem elongation in tall birch trees subjected to increases in aboveground temperature

Key message

Tall birch trees allocate extra resource due to aboveground temperature elevation to bud and male flower production rather than to plant growth. Saplings increased only plant growth under warming. Size-dependent response should be considered.

Abstract

We experimentally heated canopy organs of tall birch trees (Betula ermanii Cham.; 18–20 m high) growing at a high latitude to determine how leaf phenology, plant growth, and bud and male flower production might shift in response to increases in aboveground temperature during global climate change. We warmed the canopies with infrared heat lamps fixed to steel pipe scaffolds built around the trees. The temperature of the warmed canopies increased by approximately 1 °C. Warming extended the length of the growing season of canopy leaves (by accelerating leaf flush and delaying leaf fall), and significantly increased the numbers of buds and male flowers per shoot. Bud production and shoot length were positively correlated in both warmed and control branches. However, warming did not increase canopy shoot lengths. The intercept value of the positive regression slope between bud production and shoot length for warmed branches was higher than that for control branches. Thus, canopy warming had a direct positive effect on the bud production but had no indirect effect via increases in shoot length. Our experiment showed that tall birch trees allocated extra resources made available by increased aboveground temperature to bud and male flower production rather than to plant growth.

     

RESPONSES OF PHENOLOGY AND GROWTH OF BETULA UTILIS AND ABIES FAXONIANA IN SUBALPINE TIMBERLINE ECOTONE TO SIMULATED GLOBAL WARMING, WESTERN SICHUAN, CHINA

 采用开顶式生长室(Open-top chamber, OTC)模拟增温对植被影响的研究方法, 研究了川西亚高山林线交错带糙皮桦(Betula utilis) 和岷江冷杉(Abies faxoniana)幼苗物候及生长特性对模拟增温的响应。结果表明, 温度升高使岷江冷杉幼苗芽开放时间显著提前(15.2 d); 糙 皮桦春季芽物候期变化不显著, 而落叶时间明显推迟(19.7 d), 叶寿命延长(22.8 d)。与对照(CK)相比, OTC内糙皮桦叶面积和岷江冷杉叶片长度及两者侧枝生长速率都显著加快。模拟增温对两物种基径相对生长速率都表现为正效应, 增温对两物种枝叶特性及分布格局表现为不同程度 的正效应、负效应或无影响。不同功能型两物种对模拟增温响应方式存在一定程度差异。     

Elevated [CO2] and nutrient status modified leaf phenology and growth rhythm of young Populus trichocarpa trees in a 3-year field study

Young individuals of a single clone of black cottonwood, in Iceland, were exposed for 3 years to elevated atmospheric CO₂ concentrations [CO₂] in whole-tree chambers at natural and high nutrient availability. No treatment effects were found at bud break or the start of shoot extension in spring. Autumn phenology was, however, affected both by elevated [CO₂] and changes in nutrient status. The time of annual growth cessation was linearly related to leaf nitrogen concentration, irrespective of CO₂ treatment. At low (natural) nutrient availability, elevated [CO₂] accelerated growth cessation and bud set, which reduced the period of active growth. An earlier and more pronounced leaf senescence and corresponding loss of photosynthetic capacity further decreased carbon acquisition in elevated [CO₂]. The negative [CO₂] effect on duration of shoot extension and leaf senescence existed, but was not as pronounced, when trees grew at higher nutrient availability. Improved nutrient availability extended the shoot extension period and delayed leaf senescence. It is suggested that trees grown in elevated [CO₂] altered their autumn phenology as an effect of a signal similar to that in trees growing at low nutrient availability, i.e. an imbalance between carbon and nitrogen sources. These alterations in autumn phenology may be important when predicting how trees will grow in a future CO₂ environment.     

Quantitative genetic architecture of adaptive phenology traits in the deciduous tree,Populus trichocarpa (Torr. and Gray)

In a warming climate, the ability to accurately predict and track shifting environmental conditions will be fundamental for plant survival. Environmental cues define the transitions between growth and dormancy as plants synchronise development with favourable environmental conditions, however these cues are predicted to change under future climate projections which may have profound impacts on tree survival and growth. Here, we use a quantitative genetic approach to estimate the genetic basis of spring and autumn phenology in Populus trichocarpa to determine this species capacity for climate adaptation. We measured bud burst, leaf coloration, and leaf senescence traits across two years (2017–2018) and combine these observations with measures of lifetime growth to determine how genetic correlations between phenology and growth may facilitate or constrain adaptation. Timing of transitions differed between years, although we found strong cross year genetic correlations in all traits, suggesting that genotypes respond in consistent ways to seasonal cues. Spring and autumn phenology were correlated with lifetime growth, where genotypes that burst leaves early and shed them late had the highest lifetime growth. We also identified substantial heritable variation in the timing of all phenological transitions (h² = 0.5–0.8) and in lifetime growth (h² = 0.8). The combination of additive variation and favourable genetic correlations in phenology traits suggests that populations of cultivated varieties of P. Trichocarpa may have the capability to adapt their phenology to climatic changes without negative impacts on growth.Subject terms: Plant breeding, Forest ecology, Evolutionary genetics     

Natural phenological variation in aspen (Populus tremula): the SwAsp collection

The genus Populus is currently the main model system for genetic, genomic, and physiological research in trees. Phenotypic variation in aspen (Populus tremula) populations growing in different environments across Sweden is expected to reflect genetic variation that is important for local adaptation. To analyze such natural phenotypic and genetic variation, the Swedish Aspen (SwAsp) Collection was established. Trees were taken from 12 different populations across Sweden, from 56° to 66° latitude north and planted in two common gardens in Ekebo (55.9°N) and Sävar (63.4°N). Data related to phenological and growth traits were collected during the second year of growth. Some traits like the date of bud set and leaf area duration showed strong clinal variation patterns with latitude in both field trials, but the date of bud flush did not change along a latitudinal cline. The phenological traits showed moderate within-populations heritabilities, although growth traits showed weaker clinal patterns and lower heritabilities than the phenological traits. This research forms the starting point for the development of the SwAsp collection, a resource facilitating analysis of the natural genetic variation in aspen, the elucidation of the structure and dynamics of aspen populations, and the future identification of the genes controlling adaptive traits using association mapping of selected candidate genes.