Phenological Plasticity and Adaptive Potential of Sugar Maple Populations

Phenological Plasticity and Adaptive Potential of Sugar Maple Populations | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Phenological Plasticity and Adaptive Potential of Sugar Maple Populations guo xiali, Valentina Buttò, Yann Surget-Groba, Jian-Guo Huang, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-627097/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Global changes affect the growing conditions of terrestrial ecosystems, mismatching the phenological adaptation of plants to local climates at mid and high latitudes. Their long lifespan and slow reproductive cycles prevent trees from tracking the quick shift in their usual climatic conditions, thus endangering the survival of local populations. In this study, we explored the phenological plasticity and adaptive potential of bud burst in sugar maple ( Acer saccharum Marsh.) seedlings from 30 Canadian origins with contrasting climates planted in two common gardens near and at the northern boundary of the species range. Bud development and leafing occurred in April-May, with complete bud burst lasting between 21 and 29 days. On average, bud swelling differed by 12 days between common gardens. However, this difference decreased to 4 days for complete leafing. Both factors site and seed origin affected bud burst, which represented the phenological plasticity and adaptation of sugar maple, respectively. Overall, the former (7.4–88.3%) contributed more than the latter (9.2–25.5%) to the variance in bud burst, despite the wide climatic range among the provenance origins compared with that at the two common gardens. Adaptation to local conditions provide the genetic tools for the survival of species across wide climatic ranges. Plasticity enables physiological responses of individuals to quick environmental changes. Our study demonstrated the major role of plasticity in bud phenology, and revealed the importance of investing resources in mechanisms dealing with the climatic challenges due to inter-annual variations in weather events. Scientific Communication Environmental Engineering Acer saccharum bud burst common garden climate change ecotype temperature Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Climate change is altering the growing conditions of plants in terrestrial ecosystems, and mainly tree growth at mid and high latitudes. In order to survive under such a rapid change, plants can either track their usual conditions through migration (Angert et al. 2011; Chen et al., 2011), or change their growth strategies to match the novel climate in situ (Aitken et al. 2008; Hoffmann and Sgrò 2011). However, given their long lifespan, slow regeneration cycles, the presence of geographical barriers (Scheller and Mladenoff 2008 ), and the rapid rates of climate change (Gray and Hamann 2013 ), trees could be unable to match the new environmental conditions. Under such a context, the ability to deal locally with a changing climate could be a critical factor for the survival of local populations in the medium and long term. Phenology describes the recurring sequence of biological events during the plant cycle, such as bud burst, flowering, and leaf senescence (Richardson et al. 2013 ). Among this sequence, the timings of bud burst in spring and the beginning of the growing season have received a lot attention given the close relationship with carbon sequestration and growth. An earlier bud burst is related to a longer growing season (Leinonen and Kramer 2002 ), allowing plants to produce more carbohydrates to invest in growth and reserves. The accumulation of cold temperatures in autumn and winter (chilling), heat in spring (forcing), and hours of light (photoperiod) are the important factors involved in growth reactivation (Arora et al. 2003 ; Chen et al. 2019 ; Huang et al. 2020 ) and can affect the annual time window when trees can perform photosynthesis. Previous studies have demonstrated that climate warming has strongly advanced bud phenology due to the earlier fulfilment of forcing, resulting in an increased carbon sequestration in trees (Bronson et al. 2009 ). However, the question of the adaptive potential of phenology to climate variations remains unanswered. Local adaptation is a long-term evolutionary process driven by natural selection. In a given site, the genotypes better matching local growing conditions survive and are maintained within the population. Thus, local genotypes show a higher fitness in their home environment (Kawecki and Ebert 2004). Numerous studies have shown the local adaptation regarding a number of functional traits, such as plant diameter (Gárate-Escamilla et al. 2019 ), height (Beaulieu et al. 2004 ), leaf area (Roybal and Butterfield 2018 ), and bud phenology (Salmela et al. 2013 ; Silvestro et al. 2019 ). For example, in a cold environment, where the growing season is limited by low temperatures, trees tend to have a lower forcing requirement, allowing bud burst to start earlier to take advantage of the favorable growing conditions (Guo et al. 2020 ). Phenotypic plasticity is the ability of a genotype to express different phenotypes within a range of environmental conditions (Nicotra et al. 2010 ). Plants invest resources in the synthesis of special compounds for maintenance costs, which allows information to be obtained on the surrounding conditions (Auld et al. 2010 ). Once the signals of environmental change are detected, alternative reaction norms, a series of phenotypes, could be induced to cope with the new conditions. These multiple phenotypes have different fitness, some of them inevitably mismatching the environmental conditions, with consequent maladaptation (Ghalambor et al. 2007 ). As an important component of fitness, phenotypic plasticity enables species to respond quickly, and more quicker/faster than genotypic adaptation, to the environment changes (Chevin et al. 2013 ; Franks et al. 2014 ). However, high phenotypic plasticity is expensive in terms of maintenance costs, developmental instability (Van Buskirk and Steiner 2009 ), and the potential occurrence of maladaptation. Local adaptation and plasticity are two important strategies for plants to deal with climate change. However, no consensus has been reached on the relative importance of them. For example, some studies demonstrated a dominant role of local adaptation in mediating the relationship between functional traits and environment (Leimu & Fischer 2008; Laughlin & Messier 2015). Others have shown that plasticity exceeds adaptation in explaining the variation of tree height (Gárate-Escamilla et al. 2019 ), leaf area (Bresson et al. 2011 ), and bud phenology (Vitasse et al. 2013 ). A precise estimation of the relative contribution of local adaptation and plasticity can quantify the ability of species to perform across wide distribution ranges, and predict the potential changes of this range under global warming. Due to its wide distribution range, strong sensitivity to climate, and the economic importance for eastern North America, sugar maple ( Acer saccharum Marsh.) has been regarded as a model to study the response of species to climate change (Li et al. 2018 ; Putnam and Reich 2017 ). Previous studies showed evidence of ecotypic differentiation of bud phenology, mainly regarding the minimum temperatures occurring in spring at the seed origin (Guo et al. 2020 ; Ren et al. 2020 ). In this study, we explored the adaptive potential and plasticity of sugar maple on seedlings from 30 Canadian origins planted in two common gardens near and at the northern boundary of the species range. We aimed to quantify the relative contribution of adaptation and phenotypic plasticity to the variance in bud phenology. We tested the hypothesis that phenotypic plasticity is more important than local adaptation in regulating bud phenology due to the high variability in short-term weather events experienced in this area of the North American continent. Materials And Methods Seed origin A total of 30 stands of sugar maple located across the natural range in Canada were used in this study (Fig. 1 ). Seeds collected from different stands represent different seed origins. This area is included in the bioclimatic domains of deciduous and mixed forests of the northern temperate zone, which is dominated by both broadleaves and conifers, mainly sugar maple, red maple ( Acer rubrum L.), yellow birch ( Betula alleghaniensis Britt) and fir ( Abies balsamea (L.) Mill.). The climate is continental, with cold winters and warm summers. Plant material and common garden The seeds were collected either from single mother trees (National Tree Seed Centre, Natural Resources Canada), or at stand level (Ministère des Forêts, de la Faune et des Parcs) (Table S1). Some seed origins were collected from the same area (Mavis Mills). In 2018, the seeds were used to grow seedlings that were planted in two common gardens located in Chicoutimi and Ripon (QC, Canada) in spring 2019 at a distance of 3 m × 3.5 m. Between 6 and 10 seedlings per origin were planted and submitted to phenological observations, for a total of 217 and 242 seedlings in Chicoutimi and Ripon, respectively. Phenological observations We monitored leaf phenology of seedlings twice per week in 2020. Eight phases of bud and leaf development were recorded according to Skinner and Parker (1994): (1) bud swell, with reddish scales and enlarging bud; (2) bud elongation, with a yellowish color between the scales; (3) green tip stage, with the tip and area between the scales light green but closed bud; (4) bud break, with loosened scales but barely visible expanding leaf tips; (5) extended bud break, with leaf bundle expanded beyond the scales but no separated leaves; (6) initial leaf emergence, with the leaves starting to expand perpendicularly to the base of the bud; (7) initial leaf expansion, with light green, small, and wrinkled leaves; and (8) full leaf expansion, with flattened and fully expanded leaves. Climate date extraction and bioclimatic index computation Minimum, maximum, mean temperature and precipitation during 1979–2020 were extracted according to the coordinates of all seed origins and common gardens, from ERA5 dataset on Google Earth Engine (Copernicus Climate Change Service 2017; Gorelick et al. 2017). The 19 bioclimatic variables proposed by O’Donnell and Ignizio (2012) were extracted with the biovars function of the dismo package (Hijmans et al. 2020). Statistical analyses A Principal Component Analysis (PCA) was performed based on the 19 bioclimatic variables to describe the annual trend, seasonality and extreme conditions across seed origins and between common gardens. The contribution of each bioclimatic variable to the total variance explained by the principal components was determined. We applied a generalized linear model to test the effects of phase, site (common garden), and their interaction on bud phenology of sugar maple. The effects of provenance and individuals were set as random factors. We tested the effect of plasticity (i.e. the factor site) and adaptation (i.e. the factor seed origin) on variance partitioning for the different phenological phases using the non-parametric procedure PERmutational Multivariate Analysis Of Variance (PERMANOVA) (Anderson et al. 2011) based on the adonis function, with metric Euclidean distance matrix (McArdle and Anderson 2001) and 10000 permutations. The relative contribution of site, seed origin, and their interaction to the variance of bud phenology were further calculated using PERMANOVA. All the statistics were performed in R (R Core Team 2020) using the MuMin and vegan packages (Oksanen et al. 2020). Results Climate and bioclimate of the study sites The mean annual temperature of the seed origins ranged between 2.6 and 8.4°C, recorded at seed origin 30 and 1, respectively (Table S1). On average of all seed origins, minimum and maximum mean annual temperature was − 7.9 and 18.3°C, respectively. With a minimum mean annual temperature of -11.1°C, and a maximum mean annual temperature of 16.6°C, seed origin 30 was the coldest site. Seed origin 1 was the warmest site, showing a minimum mean annual temperature of -4.8°C, and a maximum of 21.9°C (Table S1). The mean annual temperature in Ripon was 4.5°C, warmer than that in Chicoutimi (3.2°C). Annual precipitation ranged between 947 and 4924 mm, increasing towards east. PCA extracted three main principal components (PC), explaining 44.4, 26.3, and 21.2% of the variability in the bioclimatic variables for seed origins and common gardens (Table S2). PC1 separated continental and oceanic climates, situated on the left and right quadrants, respectively (Fig. 2 ). Thus, the seed origins located inland had a more continental climate (higher temperature variability at seasonal and yearly scale represented by bio 4 and bio 7) than those close to the Gulf of Saint Lawrence River (Fig. 1 ). PC2 separated locations characterized by different precipitation regimes. The seed origins located in the westernmost part were generally drier (lower precipitation during the wettest period, represented by bio 16 and bio 13) than those close to the Saint Lawrence River. PC3 separated seed origins by the isothermality, i.e. the ratio between mean diurnal range and temperature annual range. Seed origins located inland showed a higher isothermality (bio 3) than those close to the Gulf of Saint Lawrence River (Fig. 2 ). Ripon and Chicoutimi, the two common gardens, were located in the upper quadrant on the right side of PCA (Fig. 2 ). Compared with most seed origins, they were characterized by colder winter temperatures (bio 6 and bio 9) and higher seasonality and annual range in temperature (bio 4 and bio 7). Chicoutimi had a higher temperature seasonality (bio 4) than Ripon. Bud and leaf phenology Bud and leaf phenology occurred earlier in Ripon than Chicoutimi (Fig. 3 ). On average, phase 1 started on DOY 121 in Ripon, 12 days earlier than Chicoutimi (DOY 133). The difference in bud phenology between the two sites decreased with the successive phases. Seedlings in Ripon exhibited phase 8 on DOY 150, only 4 days earlier than Chicoutimi (DOY 154). Overall, the period of leafing lasted 29 and 21 days in Ripon and Chicoutimi, respectively. The generalized linear model demonstrated the differences observed, calculating significant effects of phase (F = 5691.4, p < 0.0001), site (F = 523.2, p < 0.0001), and their interaction (F = 253.2, p < 0.0001) in determining the timing of bud burst. The marginal R 2 and conditional R 2 of the generalized linear model were 0.93 and 0.95, respectively, demonstrating the high fitness of the model. Variance components PERMANOVA confirmed the effect of site for all phases of leaf development ( p < 0.001), with Ripon showing the earliest growth reactivation. The effect of seed origin was significant at the beginning (phases 1–3) and ending (phases 7–8) of leaf development ( p < 0.05, Table 2). The effect of seed origin indicated that populations had different timings of bud and leaf development. The interaction site×seed origin was significant for phases 2, 7 and 8 (p < 0.05), suggesting that the difference of bud phenology between the two common gardens changed according to the seed origin. The factor site explained most variance in leaf development, between 7.4 and 88.3% (Fig. 4 ). The importance of the site was highest at the beginning, decreasing with the successive phenological phases. Overall, seed origin and site×seed origin contributed with a lower proportion of the variance, between 11.7 and 26.7%. For all phases, the variance component of seed origin (9.2–25.5%) was higher than the interaction site×seed origin (1.1–9.3%). Both variance components of seed origin and site×seed increased from phase 1 to phase 7. The residuals accounted for an increasing variance in bud phenology, from 11.1% for phase 1 to 58% for phase 8. Discussion 4.1 Plasticity of bud phenology The significant effect of site on bud phenology indicated a significant phenotypic plasticity of leaf development for sugar maple. As sessile organisms, plants benefit from a high phenological plasticity to respond quickly to environmental changes, matching the optimal moment of the year for flushing, and avoiding the negative consequences of early or late growth reactivations (Allevato et al. 2019). Individuals with high phenological plasticity in bud phenology are favored during the evolutionary process (Vitasse et al. 2010). This phenological plasticity of sugar maple may be an important factor that enables a broad tolerance of the species to various environmental conditions, and probably contributes to the wide latitudinal range of the species across eastern North America. The interaction between site and seed origin at the beginning and ending of leaf development indicated differences in phenological plasticity among sugar maple populations. These different responses may be related to contrasting adaptation strategies among populations within the same species, and involving the trade-off between lengthening the growing season and avoiding frost damages. In six temperate deciduous species, the phenological plasticity was lower in populations originating from higher elevations (Vitasse et al. 2013). A lower plasticity limits the sensitivity to warmer spring temperatures, thus delaying bud burst and ensuring the avoidance of frost. A high plasticity is expensive for plants, and requires resources for the continuous monitoring of the environment through the synthesis or use of chemical substances (Van Buskirk and Steiner 2009). For Pinus contorta and Pinus monticola growing under stressing conditions, more resources were invested to resist drought and frost, thus, a reduced plasticity was observed for growth (Chuine et al. 2006). This different magnitude of plasticity among populations plays an important role in matching environmental changes and ensuring sustained growth in the long term. 4.2 Comparison of bud phenology in two common gardens Buds of sugar maple reactivated earlier in the southern plantation (Ripon). In our study, the mean spring temperature (April-May) in 2020 in Ripon was 4.6 ℃, which was higher than that in Chicoutimi (2.4 ℃). Warmer spring conditions in Ripon speed up the achievement of forcing temperatures, thus advancing bud burst. Our results are consistent with previous studies conducted in temperate and boreal ecosystems. The timings of leaf unfolding of Fagus sylvatica L. and Quercus petraea (Matt.) Liebl . advanced at a rate of 5.7 days per additional degree Celsius along five common gardens (Vitasse et al. 2010). Similarly, buds of Populus fremontii in Arizona flush earlier in trees growing in the common gardens of warmer regions (Cooper et al. 2019). According to the 19 bioclimatic variables, the climatic distance between the two common gardens could be considered small when compared to the broad geographical gradient of the seed origins used in this study. However, we detected a difference of 12 days for the beginning of leaf development between Ripon and Chicoutimi. The two common gardens are located near or at the border of the northern distribution of sugar maple. Individuals growing at the boundary of species distribution are more sensitive to the environment (Jump et al. 2006; Normand et al. 2009). For example, the effects of winter temperature on the radial growth of European beech ( Fagus sylvatica L.) were significant only at the colder part of the distribution (Weigel et al. 2018). Similarly, European beech was more sensitive to drought at the drier boundary of the range (Jump et al. 2006; Roibu et al. 2017). In our study, the difference in spring temperature between the two common gardens reached 2.2 ℃, which explained the observed delay of bud phenology in Chicoutimi. The time gap between Ripon and Chicoutimi decreased for the later leafing stages (7 and 8). The difference in photoperiod is an important factor for spring phenology. Day length during bud burst (phase 1) was 15.0 h in Chicoutimi, which was longer than that recorded in Ripon (14.1 h) due to the different latitudes and timings of growth reactivation between common gardens. A longer photoperiod could have speeded up bud development, thus resulting in similar timings of full leaf expansion. A previous study demonstrated that sugar maple leafing benefits from a longer day length, and photoperiod can outweigh the delaying effects of colder springs (Ren et al. 2020). However, the similar ending of leaf expansion in the two common gardens remains partially unexplained, and could result from the weather events occurring during the studied year. Thus, a better understanding of the impact of current weather on leaf development is needed and requires a long-term monitoring of bud phenology in the two common gardens. 4.3 Plasticity vs local adaptation Our study revealed a higher contribution of plasticity to variance compared with local adaptation, mainly at the beginning of leaf development. Similar results were also observed in other species of boreal and temperate ecosystems (Baliuckas and Pliura 2003; Vitasse et al. 2010). The contribution of plasticity to bud burst phenology ranged from 55 to 86% in seven deciduous species in Europe, one order of magnitude higher than that of local adaptation (0.3-9%) (Vitasse et al. 2013). In another study combining experiments in situ and common garden, the genetic differentiation explained <28% of variance in the morphological and physiological traits of leaves of sessile oak and European beech, suggesting a minor effect of local adaptation on leaf functional traits (Bresson et al. 2011). Local adaptation and phenotypic plasticity act as concurring processes in the response of plants to changes in the environment, playing a different role at spatial and temporal scales. Under stable local environments, well adapted populations could maintain competitive fitness to survive. However, in a context of rapidly changing conditions, which requires fast response mechanisms (Van Kleunen and Fischer 2005), phenotypic plasticity can be favoured. Species with wide distributions, especially under continental climates, can experience a wide inter-annual variability in environmental conditions, principally temperature, one of the main limiting factors for bud phenology in spring. Individuals with high plasticity can respond to weather events quickly and gain a comparably longer period for carbon fixation and high competitive abilities (Kramer 1995). In addition, because of the long lifespan and slow and intermittent regeneration periods (masting years) of trees, individuals could rely predominantly on phenotypic plasticity for survival and growth rather than adaptation (Fox et al. 2019). It has been predicted that the mid-latitudes of North America will have experienced warming up to 7 °C at the end of the 21st century (Feng et al. 2014). On the one hand, plastic species may benefit from these changes by lengthening the growing season and increasing fitness. A previous study demonstrated that the recent advancement of 13 flowering days under global warming has helped plastic trees to improve their fitness by 40% (Anderson et al. 2012). On the other hand, an earlier bud burst seriously increases the risk of frost damage to the young developing tissues and leaves (Howe et al. 2003). Under the ongoing global changes, the climatic variability increases in magnitude, resulting in more frequent extreme weather events such heat waves (Hegerl et al. 2011; Min et al. 2011) or cooling (Wang et al. 2011). These extreme events have proved to be detrimental for tree growth and survival in the short term. Phenotypic plasticity is an important functional trait to deal with the increased uncertainty of climate in the future (Harmon et al. 2009; Donohue et al. 2013). In this study, we observed a high contribution of residuals to the variance in bud phenology, which is in agreement with previous studies (Sole-Medina et al. 2020; Varsamis et al. 2018). This large heterogeneity in phenology unexplained by our factors may suggest a high variability among individuals (Perrin et al. 2017), in addition to the potential effect of microsite conditions and sampling errors during field observations. An important effect of microsite on plant phenology seems unlikely, because the two common gardens are located in cropland areas, which were submitted to crop production in the past, and are therefore expected to be more homogeneous than natural sites. The wide variation in bud phenology within the same population may represent a diverse gene reservoir for the long-term survival of the species (Rousi and Heinonen 2007). These various genotypes ensure a potential matching between some individuals of the populations and the environmental conditions, thus allowing local persistence of the species. Conclusion Local adaptation and phenotypic plasticity act as concurring processes in the response of plants to environment, whose changes are critical for the survival of local populations. In this study, we explored the adaptive potential and phenological plasticity of sugar maple seedlings from 30 Canadian origins planted in two common gardens near and at the northern boundary of the species range. Plasticity explained up to 88.3% of the total variance for bud burst, while the variance component of adaptation ranged between 9.2 and 25.5%. Sugar maple is distributed over a wide geographical area, at different latitudes and distances from the sea. Under such diverging conditions, this species deals with complex climatic dynamics, involving different patterns of growing season length and frost risk (Guo et al. 2020). Adaptive mechanisms allow phenology to better match with local climates, thus ensuring a tolerance to wide environmental conditions. On the other hand, weather events are partially unstable, and populations face such a changing environment throughout their lifespan. In this context, the plasticity allows individuals to respond quickly to environmental changes, despite the high costs in resources required to maintain plasticity. Our study assessed the importance of plasticity and adaptation and interpreted their role under different temporal and spatial scales, respectively. Phenology could benefit from the high plasticity observed in sugar maple to cope with the challenge that climate warming will impose for the survival of local populations. Declarations ACKNOWLEDGEMENTS This work was funded by Ministère des Forêts, de la Faune et des Parcs du Québec, Natural Sciences and Engineering Research Council of Canada (Alliance Grants and Engage Grants), Fonds de Recherche Nature et Technologies Québec (Établissement de nouveaux chercheurs), National Natural Science Foundation of China (41861124001), Natural Science Foundation of Guangdong Province (2019B121202007), and the International Collaborative Key Project of the CAS (GJHZ1752). X. Guo received a State Scholarship Fund (201904910365) provided by the China Scholarship Council to conduct this research. The authors thank P. Benoît, F. Gagnon and P. Ren for technical support, Y. Gobeil for permitting the study on his property, and A. Garside for checking the English text. Conflict of Interest The authors declare that they have no conflict of interest Ethics approval (include appropriate approvals or waivers) 'Not applicable' Consent to participate (include appropriate statements) All patients included in this study gave written informed consent to participate in this research. Consent for publication (include appropriate statements) All patients included in this research gave written informed consent to publish the data and images contained within this case report. Availability of data and material (data transparency) The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request. Code availability (software application or custom code) The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request. 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Oecologia 171:663–678. doi: 10.1007/s00442-012-2580-9 Wang X et al (2011) Spring temperature change and its implication in the change of vegetation growth in North America from 1982 to 2006. Proc Natl Acad Sci U S A 108:1240–1245. doi: 10.1073/pnas.1014425108 Weigel R et al (2018) Winter matters: Sensitivity to winter climate and cold events increases towards the cold distribution margin of European beech (Fagus sylvaticaL.). J Biogeogr 45:2779–2790. doi: 10.1111/jbi.13444 Table Table 1 Effect of site, seed origin and their interaction on bud and leaf phenology in sugar maple seedlings. One, two, and three asterisks indicate p < 0.05, p<0.01, and p < 0.001, respectively Site Seed origin Site×seed origin Phase 1 1581.67*** 2.84*** 1.24 Phase 2 1255.77*** 3.01*** 0.01** Phase 3 1051.76*** 3.04*** 0.61 Phase 4 337.88*** 1.44 0.74 Phase 5 139.51*** 1.15 0.65 Phase 6 37.94*** 1.23 0.86 Phase 7 52.07*** 2.46*** 1.69* Phase 8 159.85*** 1.74* 2.53*** Supplementary Files supportinginformation.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-627097","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":36524022,"identity":"b6d45a2a-3030-4342-894c-31dc2793e74f","order_by":0,"name":"guo xiali","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAtElEQVRIiWNgGAWjYLCCBxUMDAakaUk4Q7KWxDZStMhHJB+TSJxXJ2/Ov4Dxww8GuzyCWgxvpCUbJG47bLhzxgNmyR6G5GLCWmbkGD5I3HYgweDGAQZpBoYDiQ1EaDE4kDinDqSF+TdRWuQlQLY0MCcYnG9gI84WA55nyQYJx0B+YWyz7DFIJsKWdmCIfagBhdjhwzd+VNgRYcsBGEsCpJiY2JGHG8p/ALeqUTAKRsEoGNkAAHCHPbJUZPNLAAAAAElFTkSuQmCC","orcid":"","institution":"Chinese Academy of Sciences","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"guo","middleName":"","lastName":"xiali","suffix":""},{"id":36524023,"identity":"94ad0623-0ced-48a4-a6f7-fe0628778b76","order_by":1,"name":"Valentina Buttò","email":"","orcid":"","institution":"University of Quebec: Universite du Quebec","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Valentina","middleName":"","lastName":"Buttò","suffix":""},{"id":36524024,"identity":"525a7e95-fd70-47a0-84dd-ddc70523768f","order_by":2,"name":"Yann Surget-Groba","email":"","orcid":"","institution":"University of Quebec: Universite du Quebec","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yann","middleName":"","lastName":"Surget-Groba","suffix":""},{"id":36524025,"identity":"c727f6d1-dad4-49d2-8bee-166a1a623604","order_by":3,"name":"Jian-Guo Huang","email":"","orcid":"","institution":"South China Botanical Garden","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Jian-Guo","middleName":"","lastName":"Huang","suffix":""},{"id":36524026,"identity":"3a21ba04-a752-4edc-8cae-aee5016bc776","order_by":4,"name":"Sylvain Delagrange","email":"","orcid":"","institution":"University of Quebec at Outaouais: Universite du Quebec en Outaouais","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Sylvain","middleName":"","lastName":"Delagrange","suffix":""},{"id":36524027,"identity":"05714228-62ba-4fbf-bab3-303245d9bc8f","order_by":5,"name":"Sergio Rossi","email":"","orcid":"","institution":"University of Quebec at Chicoutimi: Universite du Quebec a Chicoutimi","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Sergio","middleName":"","lastName":"Rossi","suffix":""}],"badges":[],"createdAt":"2021-06-16 06:47:53","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-627097/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-627097/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":11198631,"identity":"99651b86-7685-4870-9155-332e51d8acd1","added_by":"auto","created_at":"2021-07-07 11:28:32","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1220621,"visible":true,"origin":"","legend":"Location of the 30 seed origins of sugar maple. The stars mark the location of the two common gardens in Chicoutimi and Ripon. ","description":"","filename":"fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/13112682b3d633927aa2678d.png"},{"id":11198353,"identity":"890fa020-9f36-4f8d-a5ad-a61038351d3f","added_by":"auto","created_at":"2021-07-07 11:25:32","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":193251,"visible":true,"origin":"","legend":"Principal component analysis of the climatic variability between seed origins and common gardens. Bio1: annual mean temperature; bio 2: mean diurnal range; bio3: isothermality; bio4: temperature seasonality; bio5: max temperature of warmest month; bio6: min temperature of coldest month; bio7: temperature annual range; bio8: mean temperature of wettest quarter; bio9: mean temperature of driest quarter; bio10: mean temperature of warmest quarter; bio11: mean temperature of coldest quarter; bio12: annual precipitation; bio13: precipitation of wettest month; bio14: precipitation of driest month; bio15: precipitation seasonality; bio16: precipitation of wettest quarter; bio17: precipitation of driest quarter; bio18: precipitation of warmest quarter; bio19: precipitation of coldest quarter.","description":"","filename":"fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/0450d62dac874c0950d19c8e.png"},{"id":11198633,"identity":"b5fceefd-8c1e-4abc-8c98-639ec7fe38a0","added_by":"auto","created_at":"2021-07-07 11:28:32","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":44298,"visible":true,"origin":"","legend":"Occurrence of the different leafing phases of sugar maple in the two common gardens. Values are reported as average and standard deviation.","description":"","filename":"fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/fcb4671a5b6b168f5c70ad1b.png"},{"id":11199205,"identity":"4b39e2c3-d35f-4397-afd6-f8d2264ddc9d","added_by":"auto","created_at":"2021-07-07 11:31:32","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":61765,"visible":true,"origin":"","legend":"Relative contribution of the effects of site, seed origin and their interaction to the variance of bud phenology in sugar maple. ","description":"","filename":"fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/b9a2f9200c84c3e142cfb031.png"},{"id":18402260,"identity":"0ec27829-7ab4-49d2-a9b3-e35bbb36d35b","added_by":"auto","created_at":"2022-02-19 19:40:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1776961,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/b397009d-ea6e-4301-89e6-73550f5dd430.pdf"},{"id":11198634,"identity":"6783ba6d-6eba-4f7b-bce8-2374b72e3f37","added_by":"auto","created_at":"2021-07-07 11:28:32","extension":"docx","order_by":9,"title":"","display":"","copyAsset":false,"role":"supplement","size":50706,"visible":true,"origin":"","legend":"","description":"","filename":"supportinginformation.docx","url":"https://assets-eu.researchsquare.com/files/rs-627097/v1/89bf05b3ec408dac247185e9.docx"}],"financialInterests":"","formattedTitle":"\u003cp\u003ePhenological Plasticity and Adaptive Potential of Sugar Maple Populations\u003c/p\u003e","fulltext":[{"header":"Introduction","content":" \u003cp\u003eClimate change is altering the growing conditions of plants in terrestrial ecosystems, and mainly tree growth at mid and high latitudes. In order to survive under such a rapid change, plants can either track their usual conditions through migration (Angert et al. 2011; Chen et al., 2011), or change their growth strategies to match the novel climate \u003cem\u003ein situ\u003c/em\u003e (Aitken et al. 2008; Hoffmann and Sgr\u0026ograve; 2011). However, given their long lifespan, slow regeneration cycles, the presence of geographical barriers (Scheller and Mladenoff \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), and the rapid rates of climate change (Gray and Hamann \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), trees could be unable to match the new environmental conditions. Under such a context, the ability to deal locally with a changing climate could be a critical factor for the survival of local populations in the medium and long term.\u003c/p\u003e \u003cp\u003ePhenology describes the recurring sequence of biological events during the plant cycle, such as bud burst, flowering, and leaf senescence (Richardson et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Among this sequence, the timings of bud burst in spring and the beginning of the growing season have received a lot attention given the close relationship with carbon sequestration and growth. An earlier bud burst is related to a longer growing season (Leinonen and Kramer \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), allowing plants to produce more carbohydrates to invest in growth and reserves. The accumulation of cold temperatures in autumn and winter (chilling), heat in spring (forcing), and hours of light (photoperiod) are the important factors involved in growth reactivation (Arora et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and can affect the annual time window when trees can perform photosynthesis. Previous studies have demonstrated that climate warming has strongly advanced bud phenology due to the earlier fulfilment of forcing, resulting in an increased carbon sequestration in trees (Bronson et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). However, the question of the adaptive potential of phenology to climate variations remains unanswered.\u003c/p\u003e \u003cp\u003eLocal adaptation is a long-term evolutionary process driven by natural selection. In a given site, the genotypes better matching local growing conditions survive and are maintained within the population. Thus, local genotypes show a higher fitness in their home environment (Kawecki and Ebert 2004). Numerous studies have shown the local adaptation regarding a number of functional traits, such as plant diameter (G\u0026aacute;rate-Escamilla et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), height (Beaulieu et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), leaf area (Roybal and Butterfield \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), and bud phenology (Salmela et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Silvestro et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). For example, in a cold environment, where the growing season is limited by low temperatures, trees tend to have a lower forcing requirement, allowing bud burst to start earlier to take advantage of the favorable growing conditions (Guo et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePhenotypic plasticity is the ability of a genotype to express different phenotypes within a range of environmental conditions (Nicotra et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Plants invest resources in the synthesis of special compounds for maintenance costs, which allows information to be obtained on the surrounding conditions (Auld et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Once the signals of environmental change are detected, alternative reaction norms, a series of phenotypes, could be induced to cope with the new conditions. These multiple phenotypes have different fitness, some of them inevitably mismatching the environmental conditions, with consequent maladaptation (Ghalambor et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). As an important component of fitness, phenotypic plasticity enables species to respond quickly, and more quicker/faster than genotypic adaptation, to the environment changes (Chevin et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Franks et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). However, high phenotypic plasticity is expensive in terms of maintenance costs, developmental instability (Van Buskirk and Steiner \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), and the potential occurrence of maladaptation.\u003c/p\u003e \u003cp\u003eLocal adaptation and plasticity are two important strategies for plants to deal with climate change. However, no consensus has been reached on the relative importance of them. For example, some studies demonstrated a dominant role of local adaptation in mediating the relationship between functional traits and environment (Leimu \u0026amp; Fischer 2008; Laughlin \u0026amp; Messier 2015). Others have shown that plasticity exceeds adaptation in explaining the variation of tree height (G\u0026aacute;rate-Escamilla et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), leaf area (Bresson et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and bud phenology (Vitasse et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). A precise estimation of the relative contribution of local adaptation and plasticity can quantify the ability of species to perform across wide distribution ranges, and predict the potential changes of this range under global warming.\u003c/p\u003e \u003cp\u003eDue to its wide distribution range, strong sensitivity to climate, and the economic importance for eastern North America, sugar maple (\u003cem\u003eAcer saccharum\u003c/em\u003e Marsh.) has been regarded as a model to study the response of species to climate change (Li et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Putnam and Reich \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Previous studies showed evidence of ecotypic differentiation of bud phenology, mainly regarding the minimum temperatures occurring in spring at the seed origin (Guo et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ren et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In this study, we explored the adaptive potential and plasticity of sugar maple on seedlings from 30 Canadian origins planted in two common gardens near and at the northern boundary of the species range. We aimed to quantify the relative contribution of adaptation and phenotypic plasticity to the variance in bud phenology. We tested the hypothesis that phenotypic plasticity is more important than local adaptation in regulating bud phenology due to the high variability in short-term weather events experienced in this area of the North American continent.\u003c/p\u003e "},{"header":"Materials And Methods","content":"\u003cdiv class=\"Section2\" id=\"Sec3\"\u003e\n \u003ch2\u003eSeed origin\u003c/h2\u003e\n \u003cp\u003eA total of 30 stands of sugar maple located across the natural range in Canada were used in this study (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). Seeds collected from different stands represent different seed origins. This area is included in the bioclimatic domains of deciduous and mixed forests of the northern temperate zone, which is dominated by both broadleaves and conifers, mainly sugar maple, red maple (\u003cem\u003eAcer rubrum\u003c/em\u003e L.), yellow birch (\u003cem\u003eBetula alleghaniensis\u003c/em\u003e Britt) and fir (\u003cem\u003eAbies balsamea\u003c/em\u003e (L.) Mill.). The climate is continental, with cold winters and warm summers.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec4\"\u003e\n \u003ch2\u003ePlant material and common garden\u003c/h2\u003e\n \u003cp\u003eThe seeds were collected either from single mother trees (National Tree Seed Centre, Natural Resources Canada), or at stand level (Minist\u0026egrave;re des For\u0026ecirc;ts, de la Faune et des Parcs) (Table S1). Some seed origins were collected from the same area (Mavis Mills). In 2018, the seeds were used to grow seedlings that were planted in two common gardens located in Chicoutimi and Ripon (QC, Canada) in spring 2019 at a distance of 3 m \u0026times; 3.5 m. Between 6 and 10 seedlings per origin were planted and submitted to phenological observations, for a total of 217 and 242 seedlings in Chicoutimi and Ripon, respectively.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec5\"\u003e\n \u003ch2\u003ePhenological observations\u003c/h2\u003e\n \u003cp\u003eWe monitored leaf phenology of seedlings twice per week in 2020. Eight phases of bud and leaf development were recorded according to Skinner and Parker (1994): (1) bud swell, with reddish scales and enlarging bud; (2) bud elongation, with a yellowish color between the scales; (3) green tip stage, with the tip and area between the scales light green but closed bud; (4) bud break, with loosened scales but barely visible expanding leaf tips; (5) extended bud break, with leaf bundle expanded beyond the scales but no separated leaves; (6) initial leaf emergence, with the leaves starting to expand perpendicularly to the base of the bud; (7) initial leaf expansion, with light green, small, and wrinkled leaves; and (8) full leaf expansion, with flattened and fully expanded leaves.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec6\"\u003e\n \u003ch2\u003eClimate date extraction and bioclimatic index computation\u003c/h2\u003e\n \u003cp\u003eMinimum, maximum, mean temperature and precipitation during 1979\u0026ndash;2020 were extracted according to the coordinates of all seed origins and common gardens, from ERA5 dataset on Google Earth Engine (Copernicus Climate Change Service 2017; Gorelick et al. 2017). The 19 bioclimatic variables proposed by O\u0026rsquo;Donnell and Ignizio (2012) were extracted with the biovars function of the dismo package (Hijmans et al. 2020).\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec7\"\u003e\n \u003ch2\u003eStatistical analyses\u003c/h2\u003e\n \u003cp\u003eA Principal Component Analysis (PCA) was performed based on the 19 bioclimatic variables to describe the annual trend, seasonality and extreme conditions across seed origins and between common gardens. The contribution of each bioclimatic variable to the total variance explained by the principal components was determined. We applied a generalized linear model to test the effects of phase, site (common garden), and their interaction on bud phenology of sugar maple. The effects of provenance and individuals were set as random factors. We tested the effect of plasticity (i.e. the factor site) and adaptation (i.e. the factor seed origin) on variance partitioning for the different phenological phases using the non-parametric procedure PERmutational Multivariate Analysis Of Variance (PERMANOVA) (Anderson et al. 2011) based on the adonis function, with metric Euclidean distance matrix (McArdle and Anderson 2001) and 10000 permutations. The relative contribution of site, seed origin, and their interaction to the variance of bud phenology were further calculated using PERMANOVA. All the statistics were performed in R (R Core Team 2020) using the \u003cem\u003eMuMin\u003c/em\u003e and \u003cem\u003evegan\u003c/em\u003e packages (Oksanen et al. 2020).\u003c/p\u003e\n\u003c/div\u003e"},{"header":"Results","content":"\u003cdiv class=\"Section2\" id=\"Sec9\"\u003e\n \u003ch2\u003eClimate and bioclimate of the study sites\u003c/h2\u003e\n \u003cp\u003eThe mean annual temperature of the seed origins ranged between 2.6 and 8.4\u0026deg;C, recorded at seed origin 30 and 1, respectively (Table S1). On average of all seed origins, minimum and maximum mean annual temperature was \u0026minus;\u0026thinsp;7.9 and 18.3\u0026deg;C, respectively. With a minimum mean annual temperature of -11.1\u0026deg;C, and a maximum mean annual temperature of 16.6\u0026deg;C, seed origin 30 was the coldest site. Seed origin 1 was the warmest site, showing a minimum mean annual temperature of -4.8\u0026deg;C, and a maximum of 21.9\u0026deg;C (Table S1). The mean annual temperature in Ripon was 4.5\u0026deg;C, warmer than that in Chicoutimi (3.2\u0026deg;C). Annual precipitation ranged between 947 and 4924 mm, increasing towards east.\u003c/p\u003e\n \u003cp\u003ePCA extracted three main principal components (PC), explaining 44.4, 26.3, and 21.2% of the variability in the bioclimatic variables for seed origins and common gardens (Table S2). PC1 separated continental and oceanic climates, situated on the left and right quadrants, respectively (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). Thus, the seed origins located inland had a more continental climate (higher temperature variability at seasonal and yearly scale represented by bio 4 and bio 7) than those close to the Gulf of Saint Lawrence River (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). PC2 separated locations characterized by different precipitation regimes. The seed origins located in the westernmost part were generally drier (lower precipitation during the wettest period, represented by bio 16 and bio 13) than those close to the Saint Lawrence River. PC3 separated seed origins by the isothermality, i.e. the ratio between mean diurnal range and temperature annual range. Seed origins located inland showed a higher isothermality (bio 3) than those close to the Gulf of Saint Lawrence River (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eRipon and Chicoutimi, the two common gardens, were located in the upper quadrant on the right side of PCA (Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). Compared with most seed origins, they were characterized by colder winter temperatures (bio 6 and bio 9) and higher seasonality and annual range in temperature (bio 4 and bio 7). Chicoutimi had a higher temperature seasonality (bio 4) than Ripon.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec10\"\u003e\n \u003ch2\u003eBud and leaf phenology\u003c/h2\u003e\n \u003cp\u003eBud and leaf phenology occurred earlier in Ripon than Chicoutimi (Fig. \u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e). On average, phase 1 started on DOY 121 in Ripon, 12 days earlier than Chicoutimi (DOY 133). The difference in bud phenology between the two sites decreased with the successive phases. Seedlings in Ripon exhibited phase 8 on DOY 150, only 4 days earlier than Chicoutimi (DOY 154). Overall, the period of leafing lasted 29 and 21 days in Ripon and Chicoutimi, respectively. The generalized linear model demonstrated the differences observed, calculating significant effects of phase (F\u0026thinsp;=\u0026thinsp;5691.4, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001), site (F\u0026thinsp;=\u0026thinsp;523.2, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001), and their interaction (F\u0026thinsp;=\u0026thinsp;253.2, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) in determining the timing of bud burst. The marginal R\u003csup\u003e2\u003c/sup\u003e and conditional R\u003csup\u003e2\u003c/sup\u003e of the generalized linear model were 0.93 and 0.95, respectively, demonstrating the high fitness of the model.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv class=\"Section2\" id=\"Sec11\"\u003e\n \u003ch2\u003eVariance components\u003c/h2\u003e\n \u003cp\u003ePERMANOVA confirmed the effect of site for all phases of leaf development (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with Ripon showing the earliest growth reactivation. The effect of seed origin was significant at the beginning (phases 1\u0026ndash;3) and ending (phases 7\u0026ndash;8) of leaf development (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05, Table\u0026nbsp;2). The effect of seed origin indicated that populations had different timings of bud and leaf development. The interaction site\u0026times;seed origin was significant for phases 2, 7 and 8 (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), suggesting that the difference of bud phenology between the two common gardens changed according to the seed origin.\u003c/p\u003e\n \u003cp\u003eThe factor site explained most variance in leaf development, between 7.4 and 88.3% (Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003e). The importance of the site was highest at the beginning, decreasing with the successive phenological phases. Overall, seed origin and site\u0026times;seed origin contributed with a lower proportion of the variance, between 11.7 and 26.7%. For all phases, the variance component of seed origin (9.2\u0026ndash;25.5%) was higher than the interaction site\u0026times;seed origin (1.1\u0026ndash;9.3%). Both variance components of seed origin and site\u0026times;seed increased from phase 1 to phase 7. The residuals accounted for an increasing variance in bud phenology, from 11.1% for phase 1 to 58% for phase 8.\u003c/p\u003e\n\u003c/div\u003e"},{"header":"Discussion","content":"\u003ch2\u003e4.1 Plasticity of bud phenology\u003c/h2\u003e\n\u003cp\u003eThe significant effect of site on bud phenology indicated a significant phenotypic plasticity of leaf development for sugar maple.\u0026nbsp;As sessile organisms, plants benefit from\u0026nbsp;a\u0026nbsp;high phenological plasticity to respond quickly to environmental changes, matching the optimal moment of the year for flushing, and avoiding the negative consequences of early or late growth reactivations\u0026nbsp;(Allevato et al. 2019). Individuals with high phenological plasticity in bud phenology are favored during the evolutionary process (Vitasse et al. 2010). This phenological plasticity of sugar maple may be an important factor that enables a broad tolerance of the species to various environmental conditions, and probably contributes to the wide latitudinal range of the species across eastern North America.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe interaction between site and seed origin at the beginning and ending of leaf development indicated differences in phenological plasticity among sugar maple populations. These different responses may be related to contrasting adaptation strategies among populations within the same species, and involving the trade-off between lengthening the growing season and avoiding frost damages. In six temperate deciduous species, the phenological plasticity was lower in populations originating from higher elevations\u0026nbsp;(Vitasse et al. 2013). A lower plasticity limits the sensitivity to warmer spring temperatures, thus delaying bud burst and ensuring the avoidance of frost. A high plasticity is expensive for plants, and requires resources for the continuous monitoring of the environment through the synthesis or use of chemical substances \u0026nbsp;(Van Buskirk and Steiner 2009). For \u003cem\u003ePinus contorta\u0026nbsp;\u003c/em\u003eand \u003cem\u003ePinus monticola\u0026nbsp;\u003c/em\u003egrowing under stressing conditions, more resources were invested to resist drought and frost, thus, a reduced plasticity was observed for growth\u0026nbsp;(Chuine et al. 2006). This different magnitude of plasticity among populations plays an important role in matching environmental changes and ensuring sustained growth in the long term. \u0026nbsp;\u003c/p\u003e\n\u003ch2\u003e4.2 Comparison of bud phenology in two common gardens\u003c/h2\u003e\n\u003cp\u003eBuds of sugar maple reactivated earlier in the southern plantation (Ripon). In our study, the mean spring temperature (April-May) in 2020 in Ripon was 4.6\u0026nbsp;℃, which was higher than that in Chicoutimi (2.4\u0026nbsp;℃).\u0026nbsp;Warmer spring conditions in Ripon speed up the achievement of forcing temperatures, thus advancing bud burst.\u0026nbsp;Our results are consistent with previous studies conducted in temperate and boreal ecosystems. The timings of leaf unfolding of\u0026nbsp;\u003cem\u003eFagus sylvatica\u003c/em\u003e L. and \u003cem\u003eQuercus petraea\u003c/em\u003e (Matt.) Liebl\u003cem\u003e.\u0026nbsp;\u003c/em\u003eadvanced at a rate of 5.7 days per additional degree Celsius along five common gardens\u0026nbsp;(Vitasse et al. 2010). Similarly, buds of \u003cem\u003ePopulus fremontii\u003c/em\u003e in Arizona flush earlier in trees growing in the common gardens of warmer regions\u0026nbsp;(Cooper et al. 2019).\u003c/p\u003e\n\u003cp\u003eAccording to the 19 bioclimatic variables, the climatic distance between the two common gardens could be considered small when compared to the broad geographical gradient of the seed origins used in this study. However, we detected a difference of 12 days for the beginning of leaf development between Ripon and Chicoutimi. The two common gardens are located near or at the border of the northern distribution of sugar maple. Individuals growing at the boundary of species distribution are more sensitive to the environment\u0026nbsp;(Jump et al. 2006; Normand et al. 2009). For example, the effects of winter temperature on the radial growth of European beech (\u003cem\u003eFagus sylvatica\u003c/em\u003e L.) were significant only at the colder part of the distribution\u0026nbsp;(Weigel et al. 2018). Similarly, European beech was more sensitive to drought at the drier boundary of the range (Jump et al. 2006; Roibu et al. 2017). \u0026nbsp;In our study, the difference in spring temperature between the two common gardens reached 2.2\u0026nbsp;℃, which explained the observed delay of bud phenology in Chicoutimi. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe time gap between Ripon and Chicoutimi decreased for the later leafing stages (7 and 8). The difference in photoperiod is an important factor for spring phenology. Day length during bud burst (phase 1) was 15.0 h in Chicoutimi, which was longer than that recorded in Ripon (14.1 h) due to the different latitudes and timings of growth reactivation between common gardens. A longer photoperiod could have speeded up bud development, thus resulting in similar timings of full leaf expansion. A previous study demonstrated that sugar maple leafing benefits from a longer day length, and photoperiod can outweigh the delaying effects of colder springs\u0026nbsp;(Ren et al. 2020). However, the similar ending of leaf expansion in the two common gardens remains partially unexplained, and could result from the weather events occurring during the studied year. Thus, a better understanding of the impact of current weather on leaf development is needed and requires a long-term monitoring of bud phenology in the two common gardens.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003e4.3 Plasticity vs local adaptation\u0026nbsp;\u003c/h2\u003e\n\u003cp\u003eOur study revealed a higher contribution of plasticity to variance compared with local adaptation, mainly at the beginning of leaf development. Similar results were also observed in other species of boreal and temperate ecosystems\u0026nbsp;(Baliuckas and Pliura 2003; Vitasse et al. 2010). The contribution of plasticity to bud burst phenology ranged from 55 to 86% in seven deciduous species in Europe, one order of magnitude higher than that of local adaptation (0.3-9%)\u0026nbsp;(Vitasse et al. 2013). In another study combining experiments in situ and common garden, the genetic differentiation explained \u0026lt;28% of variance in the morphological and physiological traits of leaves of sessile oak and European beech, suggesting a minor effect of local adaptation on leaf functional traits\u0026nbsp;(Bresson et al. 2011).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eLocal adaptation and phenotypic plasticity act as concurring processes in the response of plants to changes in the environment, playing a different role at spatial and temporal scales. Under stable local environments, well adapted populations could maintain competitive fitness to survive. However, in a context of rapidly changing conditions, which requires fast response mechanisms (Van Kleunen and Fischer 2005), phenotypic plasticity can be favoured. Species with wide distributions, especially under continental climates, can experience a wide inter-annual variability in environmental conditions, principally temperature, one of the main limiting factors for bud phenology in spring. Individuals with high plasticity can respond to weather events quickly and gain a comparably longer period for carbon fixation and high competitive abilities (Kramer 1995). In addition, because of the long lifespan and slow and intermittent regeneration periods (masting years) of trees, individuals could rely predominantly on phenotypic plasticity for survival and growth rather than adaptation (Fox et al. 2019).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIt has been predicted that the mid-latitudes of North America will have experienced warming up to 7 \u0026deg;C at the end of the 21st century (Feng et al. 2014). On the one hand, plastic species may benefit from these changes by lengthening the growing season and increasing fitness. A previous study demonstrated that the recent advancement of 13 flowering days under global warming has helped plastic trees to improve their fitness by 40% (Anderson et al. 2012). On the other hand, an earlier bud burst seriously increases the risk of frost damage to the young developing tissues and leaves (Howe et al. 2003). Under the ongoing global changes, the climatic variability increases in magnitude, resulting in more frequent extreme weather events such heat waves (Hegerl et al. 2011; Min et al. 2011) or cooling (Wang et al. 2011). These extreme events have proved to be detrimental for tree growth and survival in the short term. Phenotypic plasticity is an important functional trait to deal with the increased uncertainty of climate in the future (Harmon et al. 2009; Donohue et al. 2013).\u003c/p\u003e\n\u003cp\u003eIn this study, we observed a high contribution of residuals to the variance in bud phenology, which is in agreement with previous studies (Sole-Medina et al. 2020; Varsamis et al. 2018). This large heterogeneity in phenology unexplained by our factors may suggest a high variability among individuals (Perrin et al. 2017), in addition to the potential effect of microsite conditions and sampling errors during field observations. An important effect of microsite on plant phenology seems unlikely, because the two common gardens are located in cropland areas, which were submitted to crop production in the past, and are therefore expected to be more homogeneous than natural sites. The wide variation in bud phenology within the same population may represent a diverse gene reservoir for the long-term survival of the species (Rousi and Heinonen 2007). These various genotypes ensure a potential matching between some individuals of the populations and the environmental conditions, thus allowing local persistence of the species.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eLocal adaptation and phenotypic plasticity act as concurring processes in the response of plants to environment, whose changes are critical for the survival of local populations. In this study, we explored the adaptive potential and phenological plasticity of sugar maple seedlings from 30 Canadian origins planted in two common gardens near and at the northern boundary of the species range. Plasticity explained up to 88.3% of the total variance for bud burst, while the variance component of adaptation ranged between 9.2 and 25.5%. Sugar maple is distributed over a wide geographical area, at different latitudes and distances from the sea. Under such diverging conditions, this species deals with complex climatic dynamics, involving different patterns of growing season length and frost risk (Guo et al. 2020). Adaptive mechanisms allow phenology to better match with local climates, thus ensuring a tolerance to wide environmental conditions. On the other hand, weather events are partially unstable, and populations face such a changing environment throughout their lifespan. In this context, the plasticity allows individuals to respond quickly to environmental changes, despite the high costs in resources required to maintain plasticity. Our study assessed the importance of plasticity and adaptation and interpreted their role under different temporal and spatial scales, respectively. Phenology could benefit from the high plasticity observed in sugar maple to cope with the challenge that climate warming will impose for the survival of local populations.\u003c/p\u003e\n"},{"header":"Declarations","content":"\u003ch2\u003eACKNOWLEDGEMENTS\u003c/h2\u003e\n\u003cp\u003eThis work was funded by Minist\u0026egrave;re des For\u0026ecirc;ts, de la Faune et des Parcs du Qu\u0026eacute;bec, Natural Sciences and Engineering Research Council of Canada (Alliance Grants and Engage Grants), Fonds de Recherche Nature et Technologies Qu\u0026eacute;bec (\u0026Eacute;tablissement de nouveaux chercheurs),\u0026nbsp;National Natural Science Foundation of China (41861124001), Natural Science Foundation of Guangdong Province (2019B121202007), and the International Collaborative Key Project of the CAS (GJHZ1752). X. Guo received a State Scholarship Fund (201904910365) provided by the China Scholarship Council to conduct this research.\u0026nbsp;The authors thank P. Beno\u0026icirc;t, F. Gagnon and P. Ren for technical support, Y. Gobeil for permitting the study on his property, and A. Garside for checking the English text.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eConflict of Interest\u003c/h2\u003e\n\u003cp\u003eThe authors declare that they have no conflict of interest\u003c/p\u003e\n\u003ch2\u003eEthics approval (include appropriate approvals or waivers)\u003c/h2\u003e\n\u003cp\u003e\u0026apos;Not applicable\u0026apos;\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eConsent to participate (include appropriate statements)\u003c/h2\u003e\n\u003cp\u003eAll patients included in this study gave written informed consent to participate in this research.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eConsent for publication (include appropriate statements)\u003c/h2\u003e\n\u003cp\u003eAll patients included in this research gave written informed consent to publish the data and images contained within this case report.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eAvailability of data and material (data transparency)\u003c/h2\u003e\n\u003cp\u003eThe datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eCode availability (software application or custom code)\u003c/h2\u003e\n\u003cp\u003eThe datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.\u0026nbsp;\u003cbr\u003e \u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllevato E et al (2019) Canopy damage by spring frost in European beech along the Apennines: effect of latitude, altitude and aspect. 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Plant Ecology Diversity 6:523\u0026ndash;536. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1080/17550874.2013.795627\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eScheller RM, Mladenoff DJ (2008) Simulated effects of climate change, fragmentation, and inter-specific competition on tree species migration in northern Wisconsin. USA Clim Res 36:191\u0026ndash;202\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSilvestro R, Rossi S, Zhang S, Froment I, Huang JG, Saracino A (2019) From phenology to forest management: ecotypes selection can avoid early or late frosts, but not both. For Ecol Manage 436:21\u0026ndash;26\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSole-Medina A et al (2020) Genetic variation in early fitness traits across European populations of silver birch (Betula pendula). AoB Plants 12:plaa019. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1093/aobpla/plaa019\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVan Buskirk J, Steiner UK (2009) The fitness costs of developmental canalization and plasticity. J Evol Biol 22:852\u0026ndash;860\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVan Kleunen M, Fischer M (2005) Constraints on the evolution of adaptive phenotypic plasticity in plants. New Phytol 166:49\u0026ndash;60\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVarsamis G et al (2018) Adaptive Diversity of Beech Seedlings Under Climate Change Scenarios. Front Plant Sci 9:1918. doi: \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3389/fpls.2018.01918\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVitasse Y, Bresson CC, Kremer A, Michalet R, Delzon S (2010) Quantifying phenological plasticity to temperature in two temperate tree species. Funct Ecol 24:1211\u0026ndash;1218. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/j.1365-2435.2010.01748.x\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVitasse Y et al (2013) Elevational adaptation and plasticity in seedling phenology of temperate deciduous tree species. Oecologia 171:663\u0026ndash;678. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s00442-012-2580-9\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang X et al (2011) Spring temperature change and its implication in the change of vegetation growth in North America from 1982 to 2006. Proc Natl Acad Sci U S A 108:1240\u0026ndash;1245. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1073/pnas.1014425108\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWeigel R et al (2018) Winter matters: Sensitivity to winter climate and cold events increases towards the cold distribution margin of European beech (Fagus sylvaticaL.). J Biogeogr 45:2779\u0026ndash;2790. doi:\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/jbi.13444\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Table","content":"\u003cp style=\"text-align: center;\"\u003eTable 1\u003c/p\u003e\n\u003cp style=\"text-align: center;\"\u003e\u0026nbsp;Effect of site, seed origin and their interaction on bud and leaf phenology in sugar maple seedlings. One, two, and three asterisks indicate p \u0026lt; 0.05, p\u0026lt;0.01, and p \u0026lt; 0.001, respectively\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellpadding=\"0\" cellspacing=\"0\" style=\"border-collapse: collapse; margin: 0px auto;\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003eSite\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003eSeed origin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003eSite\u0026times;seed origin\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e1581.67***\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e2.84***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e1.24\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e1255.77***\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e3.01***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e0.01**\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e1051.76***\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e3.04***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e0.61\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e337.88***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e1.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e0.74\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e139.51***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e1.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e0.65\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 6\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e37.94***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e1.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e0.86\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e52.07***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e2.46***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e1.69*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" width=\"17.884130982367758%\"\u003e\n \u003cp\u003ePhase 8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"24.937027707808564%\"\u003e\n \u003cp\u003e159.85***\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"26.19647355163728%\"\u003e\n \u003cp\u003e1.74*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" width=\"30.9823677581864%\"\u003e\n \u003cp\u003e2.53***\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Acer saccharum, bud burst, common garden, climate change, ecotype, temperature","lastPublishedDoi":"10.21203/rs.3.rs-627097/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-627097/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eGlobal changes affect the growing conditions of terrestrial ecosystems, mismatching the phenological adaptation of plants to local climates at mid and high latitudes. Their long lifespan and slow reproductive cycles prevent trees from tracking the quick shift in their usual climatic conditions, thus endangering the survival of local populations. In this study, we explored the phenological plasticity and adaptive potential of bud burst in sugar maple (\u003cem\u003eAcer saccharum\u003c/em\u003e Marsh.) seedlings from 30 Canadian origins with contrasting climates planted in two common gardens near and at the northern boundary of the species range. Bud development and leafing occurred in April-May, with complete bud burst lasting between 21 and 29 days. On average, bud swelling differed by 12 days between common gardens. However, this difference decreased to 4 days for complete leafing. Both factors site and seed origin affected bud burst, which represented the phenological plasticity and adaptation of sugar maple, respectively. Overall, the former (7.4\u0026ndash;88.3%) contributed more than the latter (9.2\u0026ndash;25.5%) to the variance in bud burst, despite the wide climatic range among the provenance origins compared with that at the two common gardens. Adaptation to local conditions provide the genetic tools for the survival of species across wide climatic ranges. Plasticity enables physiological responses of individuals to quick environmental changes. Our study demonstrated the major role of plasticity in bud phenology, and revealed the importance of investing resources in mechanisms dealing with the climatic challenges due to inter-annual variations in weather events.\u003c/p\u003e","manuscriptTitle":"Phenological Plasticity and Adaptive Potential of Sugar Maple Populations","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2021-07-07 11:25:30","doi":"10.21203/rs.3.rs-627097/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"dd77ca8b-d6d5-44f6-b205-d5986479c7d2","owner":[],"postedDate":"July 7th, 2021","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":5460450,"name":"Scientific Communication"},{"id":5460451,"name":"Environmental Engineering"}],"tags":[],"updatedAt":"2022-02-19T19:40:26+00:00","versionOfRecord":[],"versionCreatedAt":"2021-07-07 11:25:30","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-627097","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-627097","identity":"rs-627097","version":["v1"]},"buildId":"-HB7Z8yhvgn0wM9Nzuekk","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}