Stable Isotopes In Tree Rings Research Articles

Inter- and Intra-Annual Variations in Oak Tree Ring δ13C Values across Different Elevations and Their Climatic Responses in Qinling Mountains

The Qinling Mountains, serving as a natural geographical and climatic boundary in China, require comprehensive climatic records to elucidate the trends in climate changes across the country. While stable isotopes in tree rings are widely employed to indicate historical environmental changes, investigations into tree ring isotopes in the Qinling Mountains, particularly within the widespread broad-leaf oaks, remain limited. In this study, we investigated both intra- and inter-annual variations in the δ13C values of tree rings and their correlations with climatic signals over the past two decades for Quercus aliena var. acuteserrata, a dominant species among oak trees on the main peak of the Qinling Mountains. Our results reveal that responses to climate differ among altitudes and individual trees, with trees at higher altitudes exhibiting higher sensitivity to extreme climate, which is low temperatures and rainfall fluctuations during the growth period in intra-annual δ13C variations. Furthermore, the positive correlations are observed between temperature during growing season and both tree growth and the inter-annual δ13C variations. However, the climate signal appears to be hampered by oak-specific factors, such as intense competition among individuals and the age of trees. Therefore, we suggest a more rigorous selection of sampling and propose further investigations into isotopic fractionation processes in oaks for future studies.

Stable Water Isotope Signals and Their Relation to Stratiform and Convective Precipitation in the Tropical Andes

AbstractStratiform and convective precipitation are known to be associated with distinct isotopic fingerprints in the tropics. Such rain type specific isotope signals are of key importance for climate reconstructions derived from climate proxies (e.g., stable isotopes in tree rings). Recently, the relation between rain type and isotope signal in present‐day climate has been intensively discussed. While some studies point out the importance of deep convection, other studies emphasize the role of stratiform precipitation for strongly depleted isotope signals in precipitation. Uncertainties arise from observational studies due to data scarcity while modeling approaches with global climate models cannot explicitly resolve convective processes and rely on parameterizations. High‐resolution climate models are particularly important for studies over complex topography and for the simulation of convective cloud formation and organization. Therefore, we applied the isotope‐enabled version of the high‐resolution climate model from the Consortium for Small‐Scale Modeling (COSMOiso) over the Andes of tropical south Ecuador, South America, to investigate the influence of stratiform and convective rain on the stable oxygen isotope signal of precipitation (δ18OP). Our results highlight the importance of deep convection for depleting the isotopic signal of precipitation and increasing its deuterium excess. Due to the opposing effect of shallow and deep convection on the δ18OP signal, the use of a stratiform fraction might be misleading. We therefore propose to use a shallow and deep convective fraction to analyze the effect of rain types on δ18OP.

Rising utilization of stable isotopes in tree rings for climate change and forest ecology

Analyses of stable isotopes (C, O, H) in tree rings are increasingly important cross-disciplinary programs. The rapid development in this field documented in an increasing number of publications requires a comprehensive review. This study includes a bibliometric analysis-based review to better understand research trends in tree ring stable isotope research. Overall, 1475 publications were selected from the Web of Science Core Collection for 1974–2023. The findings are that: (1) numbers of annual publications and citations increased since 1974. From 1974 to 1980, there were around two relevant publications per year. However, from 2020 to 2022, this rose sharply to 109 publications per year. Likewise, average article citations were less than four per year before 1990, but were around four per article per year after 2000; (2) the major subjects using tree ring stable isotopes include forestry, geosciences, and environmental sciences, contributing to 42.5% of the total during 1974–2023; (3) the top three most productive institutions are the Chinese Academy of Sciences (423), the Swiss Federal Institute for Forest, Snow and Landscape Research (227), and the University of Arizona (204). These achievements result from strong collaborations; (4) review papers, for example, (Dawson et al., Annu Rev Ecol Syst 33:507–559, 2002) and (McCarroll and Loader, Quat Sci Rev 23:771–801, 2004), are among the most cited, with more than 1000 citations; (5) tree ring stable isotope studies mainly focus on climatology and ecology, with atmospheric CO2 one of the most popular topics. Since 2010, precipitation and drought have received increasing attention. Based on this analysis, the research stages, key findings, debated issues, limitations and directions for future research are summarized. This study serves as an important attempt to understand the progress on the use of stable isotopes in tree rings, providing scientific guidance for young researchers in this field.

Fire affects wood formation dynamics and ecophysiology of Pinus pinaster Aiton growing in a dry Mediterranean area

The increase in frequency and intensity of wildfires is seriously affecting forest ecosystems, especially in drought-prone areas. Trees’ recovery after fire is related to direct tree damage and is influenced by climate conditions, such as warm temperature and water shortage. In this study, we evaluate the post-fire effects on a Pinus pinaster Aiton forest growing in a hot and dry area of the Mediterranean region by comparing burned trees with severe crown reduction against unburned and not-defoliated trees. Inter-annual analyses of dendrochronology and stable isotopes in tree rings were combined with xylogenesis monitoring to investigate the effects of fire on tree growth, ecophysiological processes and wood formation. Tree-ring and isotope data showed a growth reduction and a decrease in photosynthetic activity in the burned trees, compared to control individuals, in the three years after fire. Further, the monitoring of cambial activity demonstrated a negative influence of warm and dry periods on wood formation, low xylem production, a delay in phenology and a reduction in xylem plasticity in burned trees. Our findings suggest that substantial photosynthetic limitations caused by crown defoliation and recurrent drought events could lead to severe growth decrease and reduction of trees ability to regain the pre-disturbance productivity rates.

Variable Response in Alpine Tree-Ring Stable Isotopes Following Volcanic Eruptions in the Tropics and Iceland

The importance of the stable isotopes in tree rings for the study of the climate variations caused by volcanic eruptions is still unclear. We studied δ18O, δD, δ13C stable isotopes of larch and cembran pine cellulose around four major eruptions with annual resolution, along with a superposed epoch analysis of 34 eruptions with 5-year resolution. Initial analysis of the tropical Tambora (1815 CE) and Samalas (1257 CE) eruptions showed a post-eruption decrease in δ18O values attributed to post-volcanic cooling and increased summer precipitation in Southern Europe, as documented by observations and climate simulations. The post-volcanic cooling was captured by the δD of speleothem fluid inclusion. The δ18O decrease was also observed in the analysis of 34 major tropical eruptions over the last 2000 years. In contrast, the eruptions of c. 750, 756, and 764 CE attributed to Icelandic volcanoes left no significant responses in the cellulose isotopes. Further analysis of all major Icelandic eruptions in the last 2000 years showed no consistent isotopic fingerprints, with the exception of lower post-volcanic δ13C values in larch. In summary, the δ18O values of cellulose can provide relevant information on climatic and hydroclimatic variations following major tropical volcanic eruptions, even when using the 5-year resolution wood samples of the Alpine Tree-Ring Isotope Record database.

CO2, nitrogen deposition and a discontinuous climate response drive water use efficiency in global forests

Reduced stomatal conductance is a common plant response to rising atmospheric CO2 and increases water use efficiency (W). At the leaf-scale, W depends on water and nitrogen availability in addition to atmospheric CO2. In hydroclimate models W is a key driver of rainfall, droughts, and streamflow extremes. We used global climate data to derive Aridity Indices (AI) for forests over the period 1965–2015 and synthesised those with data for nitrogen deposition and W derived from stable isotopes in tree rings. AI and atmospheric CO2 account for most of the variance in W of trees across the globe, while cumulative nitrogen deposition has a significant effect only in regions without strong legacies of atmospheric pollution. The relation of aridity and W displays a clear discontinuity. W and AI are strongly related below a threshold value of AI ≈ 1 but are not related where AI > 1. Tree ring data emphasise that effective demarcation of water-limited from non-water-limited behaviour of stomata is critical to improving hydrological models that operate at regional to global scales.

Paludification reduces black spruce growth rate but does not alter tree water use efficiency in Canadian boreal forested peatlands

BackgroundBlack spruce (Picea mariana (Mill.) BSP)-forested peatlands are widespread ecosystems in boreal North America in which peat accumulation, known as the paludification process, has been shown to induce forest growth decline. The continuously evolving environmental conditions (e.g., water table rise, increasing peat thickness) in paludified forests may require tree growth mechanism adjustments over time. In this study, we investigate tree ecophysiological mechanisms along a paludification gradient in a boreal forested peatland of eastern Canada by combining peat-based and tree-ring analyses. Carbon and oxygen stable isotopes in tree rings are used to document changes in carbon assimilation rates, stomatal conductance, and water use efficiency. In addition, paleohydrological analyses are performed to evaluate the dynamical ecophysiological adjustments of black spruce trees to site-specific water table variations.ResultsIncreasing peat accumulation considerably impacts forest growth, but no significant differences in tree water use efficiency (iWUE) are found between the study sites. Tree-ring isotopic analysis indicates no iWUE decrease over the last 100 years, but rather an important increase at each site up to the 1980s, before iWUE stabilized. Surprisingly, inferred basal area increments do not reflect such trends. Therefore, iWUE variations do not reflect tree ecophysiological adjustments required by changes in growing conditions. Local water table variations induce no changes in ecophysiological mechanisms, but a synchronous shift in iWUE is observed at all sites in the mid-1980s.ConclusionsOur study shows that paludification induces black spruce growth decline without altering tree water use efficiency in boreal forested peatlands. These findings highlight that failing to account for paludification-related carbon use and allocation could result in the overestimation of aboveground biomass production in paludified sites. Further research on carbon allocation strategies is of utmost importance to understand the carbon sink capacity of these widespread ecosystems in the context of climate change, and to make appropriate forest management decisions in the boreal biome.

Stable Isotopes in Tree Rings of Pinus heldreichii Can Indicate Climate Variability over the Eastern Mediterranean Region

A long-term context is important for understanding past climatic variability. Although tree-ring widths (TRWs) are widely used as a proxy for reconstructing past climate, the use of annually-resolved values of δ13C and δ18O tree-ring stable isotopes (TRSIs) is increasing and may provide further valuable information. Here, we present a 487-year-long TRW chronology and 240-year-long TRSI chronology for Bosnian pine (Pinus heldreichii H. Christ) and compare them to each other. We demonstrate that both δ13C and δ18O values are better proxies for temperature, precipitation, and drought than TRW. The correlations between these climate parameters and TRSIs are strongest for the combined summer (JJA) period. The results of temporal and spatial field correlation indicate that TRSI chronologies are stable, reliable proxies for JJA precipitation reconstruction over the whole Balkan Peninsula and surrounding eastern Mediterranean region. However, the stability of the temperature signal of the both δ13C and δ18O chronologies declines after the 1950s. Our work supports the emerging evidence that TRSI data track climate variability more accurately than a conventional TRW approach and can be subsequently used for the reconstruction of past climate.

Drought and cold spells trigger dieback of temperate oak and beech forests in northern Spain

Dieback in temperate forests is understudied, despite this biome is predicted to be increasingly affected by more extreme climate events in a warmer world. To evaluate the potential drivers of dieback we reconstructed changes in radial growth and intrinsic water-use efficiency (iWUE) from stable isotopes in tree rings. Particularly, we compared tree size, radial-growth trends, growth responses to climate (temperature, precipitation, cloudiness, number of foggy days) and drought, and changes in iWUE of declining and non-declining trees showing contrasting canopy dieback and defoliation. This comparison was done in six temperate forests located in northern Spain and based on three broadleaved tree species (Quercus robur, Quercus humilis, Fagus sylvatica). Declining trees presented lower radial-growth rates than their non-declining counterparts and tended to show lower growth variability, but not in all sites. The growth divergence between declining and non-declining trees was significant and lasted more in Q. robur (15–30 years) than in F. sylvatica (5–10 years) sites. Dieback was linked to summer drought and associated atmospheric patterns, but in the wettest Q. robur sites cold spells contributed to the growth decline. In contrast, F. sylvatica was the species most responsive to summer drought in terms of growth reduction followed by Q. humilis which showed coupled changes in growth and iWUE as a function of tree vigour. Low growth rates and higher iWUE characterized declining Q. robur and F. sylvatica trees. However, declining F. sylvatica trees became less water-use efficient close to the dieback onset, which could indicate impending tree death. In temperate forests, dieback and growth decline can be triggered by climate extremes such as dry and cold spells, and amplified by climate warming and rising drought stress.

Climate signals in tree-ring δ18 O and δ13 C from southeastern Tibet: insights from observations and forward modelling of intra- to interdecadal variability.

Stable isotopes in tree rings are increasingly used as proxies for climatic and ecophysiological changes. However, uncertainties remain about the strength and consistency of their response to environmental variation at different temporal (i.e. seasonal to inter-decadal) scales. We developed 5yr of intra-seasonal and 62yr of early- and late-wood δ13 C and δ18 O series of Smith fir (Abies georgei var. smithii) on the southeastern Tibetan Plateau, and used a process-based forward model to examine the relative importance of environmental and physiological controls on the isotopic data. In this temperate high-altitude region, the response, both δ18 O and δ13 C, is primarily to variations in relative humidity, but by different processes. In δ18 O, the response is via source water δ18 O but also arises from leaf water 18 O enrichment. In δ13 C, the response is via changes in stomatal conductance but is modified by carry-over effects from prior periods. We conclude that tree-ring δ18 O may be a more robust climate proxy than δ13 C, and δ13 C may be more suited to studies of site-related physiological responses to the local environment.

Using stable isotopes of oxygen from tree-rings to study the origin of past flood events: first results from the iberian peninsula

Tree-ring studies have been used for over fifty years to date and quantify past flood events. Stable C and O isotopes in tree-rings have also been extensively applied to the reconstruction of past environmental conditions and their changes over time. However, the two approaches have not previously been combined. In this study we explore whether the meteorological origin of precipitation causing past flood events might be assessed by investigating oxygen stable isotopes in tree rings. It is well known that floods may have different origins, e.g. heavy convective rainstorms, frontal precipitation, snow melting, etc.; each of these floodwater sources bears a particular isotopic fingerprint. This paper presents the first results of this methodology applied to a recent flash flood event occurring in Central Spain. For this purpose, a well-known heavy-rain convective event was chosen from the recent flood record. In the forested area affected by this event, six cores from each of four selected species (Pinus sylvestris L., Pinus pinaster Ait., Quercus pyrenaica L. and Alnus glutinosa (L) Gaertn), were sampled using an incremental borer. After α-cellulose extraction, the oxygen-isotope composition (δ18O) in tree-rings was analyzed and compared with climate data and δ18O values in precipitation. The isotope signal in tree-ring cellulose was dominated by spring climatic conditions but, after removing this “spring signal”, a clear and site-specific signal in latewood δ18O emerged, which was associated with the heavy-rain event. These preliminary results encourage more in-depth analyses aimed at recognizing specific precipitation sources and separating different populations of past floods according to their cause.

Stable isotopes in tree rings: towards a mechanistic understanding of isotope fractionation and mixing processes from the leaves to the wood.

The mechanistic understanding of isotope fractionation processes is increasing but we still lack detailed knowledge of the processes that determine the isotopic composition of the tree-ring archive over the long term. Especially with regard to the path from leaf photosynthate production to wood formation, post-assimilation fractionations/processes might cause at least a partial decoupling between the leaf isotope signals that record processes such as stomatal conductance, transpiration and photosynthesis, and the wood or cellulose signals that are stored in the paleophysiological record. In this review, we start from the rather well understood processes at the leaf level such as photosynthetic carbon isotope fractionation, leaf water evaporative isotope enrichment and the issue of the isotopic composition of inorganic sources (CO2 and H2O), though we focus on the less explored 'downstream' processes related to metabolism and transport. We further summarize the roles of cellulose and lignin as important chemical constituents of wood, and the processes that determine the transfer of photosynthate (sucrose) and associated isotopic signals to wood production. We cover the broad topics of post-carboxylation carbon isotope fractionation and of the exchange of organic oxygen with water within the tree. In two case studies, we assess the transfer of carbon and oxygen isotopic signals from leaves to tree rings. Finally we address the issue of different temporal scales and link isotope fractionation at the shorter time scale for processes in the leaf to the isotopic ratio as recorded across longer time scales of the tree-ring archive.

On the reconstruction of precipitation in temperature-limited environments by the use of stable isotopes in tree-rings

On the reconstruction of precipitation in temperature-limited environments by the use of stable isotopes in tree-rings

The influence of climate on 13C/ 12C and 18O/ 16O ratios in tree ring cellulose of Pinus sylvestris L. growing in the central Scandinavian Mountains

The stable isotope composition of tree rings is known to contain valuable information of past climatic and environmental changes, which may be used as a complement to tree ring width and maximum latewood density in climate reconstructions. In this study we examine the character and strength of the climate signal captured in the δ 18O and δ 13C values of Scots pine ( Pinus sylvestris L.) trees growing at the tree-line in the central Scandinavian Mountains. Between 4 and 14 trees were pooled to produce annual records of carbon and oxygen isotope ratios from tree ring cellulose, spanning the period AD 1736–2006. Weather conditions of the current growing season most strongly influenced the carbon and oxygen isotope ratios in a given tree ring. Both records showed positive correlations with temperature, sunshine and air pressure, and negative associations with precipitation. The overall climate signal in δ 13C appeared to be slightly stronger than that of δ 18O. Spatial correlation analysis with gridded instrumental data demonstrated that the carbon series captured the summer temperature signal in a broad region of mid-west Sweden and the eastern part of Norway, greatly exceeding the spatial coverage of the signal derived from pine ring widths growing in the central Scandinavian mountains. A weak relationship between δ 13C and precipitation and a much stronger temperature and sunshine dependence may imply that photosynthetic rate rather than stomatal conductance is more important in controlling the inter-annual tree ring δ 13C variability in the area. Moreover, it was shown that, the climate– δ 18O and δ 13C relationship was temporally unstable throughout the twentieth century, which was linked to large-scale shifts in climate that may have altered the isotope-climate dependence. Our results thus demonstrate that, stable isotopes in tree rings from maritime high-altitude Scandinavia can provide high-resolution regional climate information, especially regarding parameters associated with temperature. The non-stationary nature of the isotope–precipitation relationship may provide important information on past changes in the large-scale atmospheric circulation during summer. However, these issues needs to be further investigated before pine tree ring isotopes from this region can be confidently used in palaeoclimate reconstructions.

Comparison of whole wood and cellulose carbon and oxygen isotope series from Pinus nigra ssp. laricio (Corsica/France)

Stable isotopes in tree rings have widely been used for palaeoclimate reconstructions since tree rings record climatic information at annual resolution. However, various wood components or different parts of an annual tree-ring may differ in their isotopic compositions. Thus, sample preparation and subsequent laboratory analysis are crucial for the isotopic signal retained in the final tree-ring isotope series used for climate reconstruction and must therefore be considered for the interpretation of isotope–climate relationships. This study focuses on wood of Corsican Pine trees (Pinus nigra ssp. laricio) as this tree species allows to reconstruct the long-term climate evolution in the western Mediterranean. In a pilot study, we concentrated on methodological issues of sample preparation techniques in order to evaluate isotope records measured on pooled whole tree-ring cellulose and whole tree-ring bulk wood samples. We analysed 80-year long carbon and oxygen chronologies of Corsican Pine trees growing near the upper tree line on Corsica. Carbon and oxygen isotope records of whole tree-ring bulk wood and whole tree-ring cellulose from a pooled sample of 5 trees were correlated with the climate parameters monthly precipitation, temperature and the self-calibrating Palmer Drought Severity Index (sc-PDSI). Results show that the offsets in carbon and oxygen isotopes of bulk wood and cellulose are not constant over time. Both isotopes correlate with climate parameters from late winter and summer. The carbon and oxygen isotope ratios of cellulose are more sensitive to climatic variables than those of bulk wood. The results of this study imply that extraction of cellulose is a pre-requisite for the reconstruction of high-resolution climate records from stable isotope series of P. nigra ssp. laricio.

A 350 year drought reconstruction from Alpine tree ring stable isotopes

Climate reconstructions based on stable isotopes in tree rings rely on the assumption that fractionation‐controlling processes are strongly linked to meteorological variables. In this context, we investigated the climate sensitivity of 350 years of carbon and oxygen isotope ratios of tree ring cellulose from European larch obtained at a high‐elevation site in the Swiss Alps (∼2100 m above sea level). Unlike tree ring width and maximum latewood density, which contain only summer temperature information at this site, we found that our stable isotope series reveal additionally to temperature a striking sensitivity to precipitation (mainly for carbon) and sunshine duration (mainly for oxygen) during July and August. A drought index reflecting the combined temperature and precipitation influence provided the most stable correlations over time for the carbon isotope series. All of these climate‐isotope relationships are preserved in the isotope series obtained from younger trees at the same site, while strong intertree correlations further emphasize the high climate sensitivity. We thus present the first carbon‐isotope‐based summer drought reconstruction for the Swiss Alps, which provides new evidence for interannual to long‐term changes in summer regional moisture variability from 1650 to 2004 in Europe, revealing extreme drought summers in the second half of the 18th century and throughout the 20th century.

Stable isotopes in tree rings from the Russian Arctic— a proxy for winter precipitation?

Dendroclimatology studies shed light on the climate conditions prior to the instrumental data period. The classical approach is to derive past summer temperatures from the tree ring width or more recently, from tree ring density. By studying stable carbon and oxygen isotopes in tree ring cellulose, additional climate parameters may be retrieved, though the underlying mechanisms and relations between climate and tree ring isotopes are complex and may vary for different locations (McCarroll and Loader, 2004).

Stable Isotopes in Tree Rings as Proxies for Winter Precipitation Changes in the Russian Arctic over the Past 150 Years

We present results from an analysis of tree ring width and stable carbon and oxygen isotopes in tree ring cellulose of Siberian Spruce collected from remote forest islands in the northwestern Russian tundra. Ring width is often considered a proxy for summer temperatures. The aim of this pilot study was to test whether stable isotopes can provide additional information about climate during the growth of trees in this extreme environment. Comparison of δ<sup>13</sup>C and δ<sup>18</sup>O with observed meteorological data shows that there is a link between stable isotopes and winter precipitation. This may be explained by the strong influence that snow exerts on the isotopic composition of soil moisture during spring and early summer, when the new cellulose is formed. Our results show that winter precipitation in the study area was increasing from 1865-1900, and thereafter decreasing until ~ 1930. The 1960-1980 period was again rather humid, followed by a drying trend until 1990. The study highlights the potential of stable carbon and oxygen isotopes in tree rings as proxies for winter precipitation.

Temporal stability of climate‐isotope relationships in tree rings of oak and pine (Ticino, Switzerland)

Climate reconstructions based on stable isotopes in tree rings commonly rely on the assumption that climate‐isotope relations are stable over time. However, studies of tree ring growth have revealed trends thought to result from either physiological changes or changes in the climate‐growth relationship. We investigated whether or not similar trends exist for tree ring stable isotopic ratios using a statistical approach. Correlations between climate (temperature and precipitation amount) and tree ring cellulose δ13C and δ18O of oak and pine from Ticino, Switzerland, were calculated for the period AD 1660–2000. Climate calibration of tree rings was enabled by long‐term monthly resolved temperature and precipitation data sets on the basis of instrumental and documentary proxy data. Overall, five findings have been identified: (1) Isotopic ratios in tree rings most strongly reflect conditions of the current growing season, (2) temporally stable climate signals are found in pine δ13C only, (3) all other correlations between tree ring isotopes and climate are temporally unstable and characterized by shifts in correlation sign and strength, (4) climate signals in oak are strongest in the 20th century, and (5) tree ring δ13C reflects local climatic conditions while δ18O is influenced by large‐scale synoptic circulation. The nonstationary relationships observed could reflect changes in the relationship between the climate variables or a physiological adaptation to warmer conditions. Our results provide a cautionary note for the calibration of long tree ring series with 20th century relationships, at least for trees located at ecologically nonextreme sites.

Quantification of Deuterium Isotopomers of Tree-Ring Cellulose Using Nuclear Magnetic Resonance

Stable isotopes in tree rings are important tools for reconstruction of past climate. Deuterium (D) is of particular interest since it may contain climate signals and report on tree physiology. Measurements of the D/H ratio of tree-ring cellulose have proven difficult to interpret, presumably because the D/H ratio of the whole molecule blends the abundances of the seven D isotopomers of cellulose. Here we present a method to measure the abundance of the D isotopomers of tree-ring cellulose by nuclear magnetic resonance spectroscopy (NMR). The method transforms tree-ring cellulose into a glucose derivative that gives highly resolved, quantifiable deuterium NMR spectra. General guidelines for measurement of D isotopomers by NMR are described. The transformation was optimized for yield and did not alter the original D isotopomer abundances, thus, conserving the original signals recorded in wood cellulose. In the tree-ring samples tested, the abundances of D isotopomers varied by approximately +/-10% (2% standard error). This large variability can only be caused by biochemistry processes and shows that more information is present in D isotopomer abundances, compared to the D/H ratio. Therefore, measurements of the D isotopomer distribution of tree rings may be used to obtain information on long-term adaptations to environmental changes and past climate change.