树木叶片稳定碳同位素分馏对环境梯度的响应

除了自身的遗传因素外,植物对稳定碳同位素的分馏还受温度、水分状况、光照等多种环境因素的影响。通过对生长在中国北热带、亚热带和温带的13个气候区树木的稳定碳同位素分馏与环境要素的关系的研究得出,不同气候条件下树木叶片稳定碳同位素比率(δ13C)和稳定碳同位素分辨率(Δ)差异明显,分别在-23.759‰--33.914‰和14.581‰-24.354‰的范围内。且由南向北,δ13C值呈现逐渐加重的趋势、而Δ值逐渐减小。年均温度、相对湿度、降雨量及无霜期等与δ13C值呈显著负相关、与Δ值显著正相关;年均日照时数、气温年较差、蒸发量、纬度及海拔等与δ13C值呈显著正相关、而与Δ值显著负相关;而经度、年极端最高气温等与δ13C和Δ值的相关性较弱。δ13C和Δ值的变化幅度和趋势在年均温度、年较差、无霜期、活动积温分别达到约19℃、23℃、309d、6000℃出现转折,这些指标在此基础上继续增加反而使叶片δ13C值增大、Δ值减小。而当年均降雨量和相对湿度分别超过1105mm、75%以及蒸发量小于约1700mm后,δ13C和Δ值随之变化的幅度也变小。结果表明在亚热带和温带的分界线以南,水热条件均能满足植物正常生理活动的需要,环境条件的相对改变对植物生理活动及叶片稳定碳同位素分馏的影响较小;而在北方,干旱、低温等极端的外界环境条件加剧了对植物生长和生理指标的影响、造成了不同环境梯度下叶片稳定碳同位素分馏的差异。     

天山南北坡树轮稳定碳同位素对气候的响应差异

利用采自天山南坡阿克苏河流域、天山北坡伊犁河流域和吉尔吉斯斯坦伊塞克湖流域的树木年轮样本,按照树轮稳定碳同位素研究步骤,分别建立了天山南坡和天山北坡各2条树轮稳定碳同位素去趋势序列（δ¹³C_corr）。采用相关函数和共线性分析揭示了天山南北坡树轮稳定碳同位素与各气候要素之间的关系。结果表明,天山南坡树轮稳定碳同位素与生长季降水、相对湿度显著负相关,而与温度没有明显的相关关系,表明影响天山南坡树轮稳定碳同位素分馏的主要气候因子是生长季水分状况,尤其是降水;天山北坡树轮稳定碳同位素与生长季尤其是夏季的平均气温、平均最高气温和饱和水汽压亏缺显著正相关,而与降水量、相对湿度显著负相关,表明影响天山北坡树轮稳定碳同位素分馏的气候因子可能较为复杂。进一步分析表明,夏季温度、降水和相对湿度共同调控着天山北坡雪岭云杉树轮稳定碳同位素分馏。     

树木年轮中不同组分稳定碳同位素含量对气候的响应

通过研究贺兰山油松树轮中全木、综纤维素、α纤维素中占δ^13C值的变化,来探讨树木生长过程中引起其中同位素差别的原因,以及它们对环境变化的响应。研究发现不同组分的占δ^13C值在过去30a中变化并不一致,其中,树轮中α-纤维素中占δ^13C序列含有最强周围环境变化的信息。δ^13C序列和周围大气平均气温和降雨量关系密切,其中和6～8月份的平均气温相关系数为0．427(α=0．05);和2～7月份总降水量的相关系数为(r=-0．514,a=0．01)。分析发现树轮α-纤维素序列和树轮宽度序列呈显著负相关(r=-0．545,α=0．01)。对比树轮各种组分的占δ^13C值和气候之间的相关性及和树木的生长量的关系可以发现,树轮α-纤维素中碳在经历了光合作用的固碳过程中的同位素分馏后,其占δ^13C值就保持了相对稳定,是研究过去环境变化的良好载体。     

川西米亚罗林区不同海拔岷江冷杉生长对气候变化的响应

为分析青藏高原东缘半湿润区树木生长与气候关系随海拔变化的规律,于川西米亚罗林区分别在高、中、低海拔选取3个采样点,共采集132棵岷江冷杉年轮样芯,建立了最长达170a(1842-2011年)3个海拔高度的差值年表.岷江冷杉年轮指数与气候因子的相关分析表明:随海拔高度降低,温度与生长的负相关呈增加趋势.高海拔岷江冷杉径向生长与前—年冬季最低温呈显著正相关,中低海拔与当年春季均温、最高温和年均最高温呈显著负相关.低海拔岷江冷杉与当年4月降水呈显著正相关,随海拔升高降水与岷江冷杉生长的相关性降低.中低海拔岷江冷杉年表与4、5月帕尔默干旱指数(P DSI)呈显著正相关,表明在中低海拔存在春季干旱胁迫,抑制了岷江冷杉的生长.另外,大龄树木比小龄树木对气候变化的响应更敏感.     

玉龙雪山东坡不同海拔长苞冷杉径向生长与气候因子的关系

基于树木年代学方法,利用玉龙雪山东坡长苞冷杉树轮宽度资料,分别建立了3个海拔(低、中、高)长苞冷杉差值年表(RES),并利用响应分析与冗余分析(RDA)研究其径向生长与温度和降水的相关关系,分析影响玉龙雪山长苞冷杉径向生长的主要气候因子.结果表明: 玉龙雪山不同海拔长苞冷杉径向生长受气候变化的影响具有一致性,温度和降水共同作用于径向生长,这种一致性与玉龙雪山地形陡峭以及长苞冷杉的高海拔分布与浅根特性有关.不同海拔长苞冷杉径向生长均与当年7月和上年11月平均温以及当年1月降水量呈显著正相关,而与当年6月降水量呈显著负相关.响应分析与冗余分析的综合应用能够准确揭示环境梯度上树木生长与气候因子的关系.本研究可为该地区历史气候重建以及森林经营管理提供科学依据.     

川西米亚罗林区主要树木生长对气候响应的差异

为分析青藏高原东缘半湿润区不同树种树木生长对气候变化的响应规律,于川西米亚罗林区海拔3000 m左右(低海拔)采集铁杉、岷江冷杉、紫果云杉,海拔4000 m左右林线位置(高海拔)采集岷江冷杉、四川红杉,共计182棵树木年轮样芯,建立了不同树种的树轮宽度年表,对不同树种的年轮指数与各月气候因子进行相关分析.结果表明： 在低海拔处,树木生长与4、5月气温呈负相关,与4、5月降雨呈正相关,受到春季干旱胁迫的影响;但树种之间存在显著差异： 铁杉的生长受春季干旱胁迫影响最严重,岷江冷杉次之,紫果云杉所受影响很小.在高海拔处,树木生长主要受生长季温度的影响,岷江冷杉年轮指数与当年2、7月最低气温呈显著正相关,与上一年10月最高气温亦呈正相关;四川红杉年轮指数与5月最高气温呈显著正相关,但与2月均温、3月最低气温呈显著负相关.近几十年青藏高原东北缘气候有干暖化趋势,如果这种趋势持续发生,低海拔紫果云杉长势将超过铁杉和岷江冷杉;高海拔处的升温更有利于岷江冷杉的生长.     

升温突变对川西马尔康树木生长的影响

以采自川西马尔康林区的岷江冷杉(Abies faxoniana)和岷江柏(Cupressus chengiana)树轮为研究对象,对树轮指数与气候要素进行相关分析,研究1995年升温突变前后该区主要针叶树种的树轮响应变化。结果显示:1955—1994年时段,高、中海拔岷江冷杉径向生长对温度正响应,年轮指数呈缓慢的上升趋势,低海拔岷江柏受温度影响不大,年轮指数上升较快;1995—2012年时段,随着温度升高,3样点树木年轮指数并未上升,甚至呈下降趋势,表现出"响应分异现象"。对月气候要素的响应,树种间存在差异:高海拔岷江冷杉在前一时段主要表现为与冬季温度的正相关,后一阶段则转变为与4月温度的显著负相关及与4月份降水的显著正相关,受到显著的4月干旱胁迫影响;中海拔岷江冷杉后一时段温度敏感性消失,5月干旱胁迫加剧;低海拔岷江柏主要受降水影响,后一时段4月份干旱胁迫加剧。升温突变后,川西马尔康林区岷江冷杉和岷江柏均表现出"响应分异现象",在今后的气候重建及碳循环模拟中应加以考虑。     

升温突变对川西不同坡向和海拔岷江冷杉径向生长的影响

坡向和海拔作为重要的地形因子决定着林木立地条件的水热分配，为揭示川西地区升温突变前后制约岷江冷杉(Abies fargesii var.faxoniana)径向生长的气候因子变化及其在坡向和海拔上的响应格局，通过树木年轮学方法构建了4个坡向(NE、N、NW和W)的3个海拔梯度(≥3650 m)共12个岷江冷杉的标准年表，采用皮尔逊相关分析和回归分析的方法分析了升温突变前后(1980年)限制岷江冷杉径向生长主要气候因子的变化及径向生长的变化趋势。结果表明：升温前，生长季(7月)低温和降水、前一年冬季(10—11月)最高温和平均温制约岷江冷杉的生长，而当年春季(3月)温度的升高和生长季前(5月)较多的降水不利于其生长。升温后，4个坡向的林线、东北坡中海拔，西坡中、低海拔岷江冷杉的径向生长与大部分月温度表现为正相关关系，且上述样点树木径向生长明显加速。1980年升温前，制约不同样点岷江冷杉径向生长的气候因子具有一致性——坡向和海拔对其径向生长与温度相关关系的干扰和影响较小。增温促进了各坡向林线岷江冷杉的径向生长，且偏阳坡在更大的海拔范围内对增温表现出正反馈。研究对未来气候变化背景下川西不同坡...     

高山林线急尖长苞冷杉不同器官的稳定碳同位素组成分布特征

通过分析青藏高原东南部色季拉山林线物种急尖长苞冷杉不同年龄叶片、嫩枝、枝条、树干及根系的稳定碳同位素比值（6BC）及其空间分布特征,研究了植物光合作用后稳定碳同位素分馏及其影响因素．结果表明：植物不同器官δ^13C值差异显著（P〈0．001）,为树干（-24．19‰）〉枝条（-24．56‰）〉根部（-25．05‰）〉嫩枝（-25．12‰）〉叶片（-27．25‰）,说明从光合作用器官到非光合作用器官有明显的碳同位素分馏,且非光合作用器官之间也存在差异．随着急尖长苞冷杉叶片或嫩枝年龄的增加,叶片δ^13C值降低,而嫩枝δ^13C值升高（P〈0．01）．冠层上部叶片δ^13C值明显高于冠层下部（P〈0．01）,嫩枝δ^13C值则无显著性差异（P〉0．05）．远离树干2．5m的枝条δ^13C值有明显的高度变化（P〈0．01）,而离树干较近（1．5或0．5m）的枝条及树干在不同高度之间无差异（P〉0．05）．在同一冠层高度,随着与树干距离加大,枝条δ^13C值降低,且在中部和下部枝条尤为明显．说明林线地区冷杉光合作用后存在明显的碳同位素分馏;特定冠层高度树干与枝条生长所需的碳并不是全部来源于同高度的叶片光合作用合成的碳．     

普达措国家公园四种针叶树径向生长对气候因子的响应

运用树木年代学的原理和方法,对普达措国家公园大果红杉、长苞冷杉、高山松和麦吊云杉4个优势针叶树种的年轮宽度进行测量,建立年轮宽度差值年表。分析年表与香格里拉气象站的日、月气候数据的相关性,研究4个优势针叶树种的径向生长对气候因子的响应。结果表明: 大果红杉的年生长速率最高,长苞冷杉的年生长速率最低;4种针叶树径向生长对气候因子的响应存在物种特异性,大果红杉与气候因子的相关性最强,麦吊云杉的径向生长对气候因子的响应不敏感;长苞冷杉树轮宽度年表与上年冬季(11、12月)和当年夏季(7月)的平均温度呈显著正相关;大果红杉树轮宽度年表与生长季早期(6月)温度呈显著正相关,与同期降水量和相对湿度呈显著负相关;而高山松树轮宽度年表与生长季早期(5月)的降水量和相对湿度呈显著正相关,与同期最高温度呈显著负相关,表明高山松的径向生长主要受生长季早期水分可利用性的影响。     

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.     

Consequences of larch budmoth outbreaks on the climatic significance of ring width and stable isotopes of larch

Tree-ring widths and stable carbon and oxygen isotopes of five European larch trees from Lötschental, Switzerland were investigated for the period 1900–2004. The objective was to test the suitability of each of these parameters for high-frequency climate reconstructions. This is of special interest with regard to the problem of cyclic larch budmoth (LBM) infestations of alpine larch trees. The results clearly demonstrate that tree-ring width chronologies are not suitable for high-frequency reconstructions because infestations lead to variably reduced tree-ring increments, largely suppressing climate signals. On the other hand, the stable isotope chronologies proved less affected by larch budmoth outbreaks, independent of the strength of the infestations. The correlation of the carbon isotopes with summer temperatures was especially high (r = 0.73) and with precipitation lower but nevertheless significant (r = ?0.43). Oxygen isotopes were also correlated with summer temperature (r = 0.46); however, a certain perturbation of normal oxygen isotope signatures due to LBM outbreaks was evident. Contrary to tree-ring widths, none of the LBM outbreaks caused a significant disturbance of the current year’s isotopic climate signal and, most importantly, there were no delayed effects in the following years. Thus, stable carbon isotopes in tree-ring chronologies of the European larch provide an excellent opportunity for high-frequency temperature reconstructions.     

Significant Mean and Extreme Climate Sensitivity of Norway Spruce and Silver Fir at Mid-Elevation Mesic Sites in the Alps

Climate forcing is the major abiotic driver for forest ecosystem functioning and thus significantly affects the role of forests within the global carbon cycle and related ecosystem services. Annual radial increments of trees are probably the most valuable source of information to link tree growth and climate at long-term time scales, and have been used in a wide variety of investigations worldwide. However, especially in mountainous areas, tree-ring studies have focused on extreme environments where the climate sensitivity is perhaps greatest but are necessarily a biased representation of the forests within a region. We used tree-ring analyses to study two of the most important tree species growing in the Alps: Norway spruce (Picea abies) and silver fir (Abies alba). We developed tree-ring chronologies from 13 mesic mid-elevation sites (203 trees) and then compared them to monthly temperature and precipitation data for the period 1846–1995. Correlation functions, principal component analysis and fuzzy C-means clustering were applied to 1) assess the climate/growth relationships and their stationarity and consistency over time, and 2) extract common modes of variability in the species responses to mean and extreme climate variability. Our results highlight a clear, time-stable, and species-specific response to mean climate conditions. However, during the previous-year''s growing season, which shows the strongest correlations, the primary difference between species is in their response to extreme events, not mean conditions. Mesic sites at mid-altitude are commonly underrepresented in tree-ring research; we showed that strong climatic controls of growth may exist even in those areas. Extreme climatic events may play a key role in defining the species-specific responses on climatic sensitivity and, with a global change perspective, specific divergent responses are likely to occur even where current conditions are less limited.     

Influence of climate and disturbance on the growth of Tsuga canadensis at its southern limit in eastern North America

It has long been hypothesized that trees growing at range limits likely also occur near the limit of their ecological amplitude and thus, should be more sensitive to climate variability than individuals growing nearer the range core. We developed a tree-ring chronology using Tsuga canadensis individuals from three disjunct stands at the species’ southern limit to quantify the influence of climate and disturbance on radial growth patterns. The tree-ring record extended 158 years from 1850 to 2007. Significant negative relationships were found between the STANDARD chronology and monthly mean temperature, monthly maximum temperature, and monthly minimum temperature during the previous and current summer, while significant positive relationships were documented between the STANDARD chronology and monthly minimum temperature for September and October of the current year. Also, significant positive relationships were documented between the STANDARD chronology and monthly total precipitation for September of the previous year and May of the current year. Response function analysis showed that monthly climate variables (r ² = 0.22) and prior growth (r ² = 0.40) explained 62% of the variance in the T. canadensis tree-ring chronology. A time series plot for the T. canadensis chronology showed that actual tree growth agreed relatively well with the predicted growth based on significant climate variables. However, positive departures from the predicted growth were noted. Dendroecological analysis revealed these departures were likely related to disturbance events. Our results indicated that T. canadensis individuals at its southernmost extent are sensitive to regional climate, but not more so than trees nearer the range core. We hypothesize that microenvironmental conditions of T. canadensis stands at its southern limit are similar to conditions within the contiguous distribution of the species, which may explain this pattern.     

Summer temperature dependency of larch budmoth outbreaks revealed by Alpine tree-ring isotope chronologies

Larch budmoth (LBM, Zeiraphera diniana Gn.) outbreaks cause discernable physical alteration of cell growth in tree rings of host subalpine larch (Larix decidua Mill.) in the European Alps. However, it is not clear if these outbreaks also impact isotopic signatures in tree-ring cellulose, thereby masking climatic signals. We compared LBM outbreak events in stable carbon and oxygen isotope chronologies of larch and their corresponding tree-ring widths from two high-elevation sites (1800–2200 m a.s.l.) in the Swiss Alps for the period AD 1900–2004 against isotope data obtained from non-host spruce (Picea abies). At each site, two age classes of tree individuals (150–250 and 450–550 years old) were sampled. Inclusion of the latter age class enabled one chronology to be extended back to AD 1650, and a comparison with long-term monthly resolved temperature data. Within the constraints of this local study, we found that: (1) isotopic ratios in tree rings of larch provide a strong and consistent climatic signal of temperature; (2) at all sites the isotope signatures were not disturbed by LBM outbreaks, as shown, for example, by exceptionally high significant correlations between non-host spruce and host larch chronologies; (3) below-average July to August temperatures and LBM defoliation events have been coupled for more than three centuries. Dampening of Alps-wide LBM cyclicity since the 1980s and the coincidence of recently absent cool summers in the European Alps reinforce the assumption of a strong coherence between summer temperatures and LBM defoliation events. Our results demonstrate that stable isotopes in tree-ring cellulose of larch are an excellent climate proxy enabling the analysis of climate-driven changes of LBM cycles in the long term.     

Freezing and chemical preservatives alter the stable isotope values of carbon and nitrogen of the Asiatic clam (Corbicula fluminea)

We tested the impacts of most common sample preservation methods used for aquatic sample materials on the stable isotope ratios of carbon and nitrogen in clams, a typical baseline indicator organism for many aquatic food web studies utilising stable isotope analysis (SIA). In addition to common chemical preservatives ethanol and formalin, we also assessed the potential impacts of freezing on δ¹³C and δ¹⁵N values and compared the preserved samples against freshly dried and analysed samples. All preservation methods, including freezing, had significant impacts on δ¹³C and δ¹⁵N values and the effects in general were greater on the carbon isotope values (1.3–2.2‰ difference) than on the nitrogen isotope values (0.9–1.0‰ difference). However, the impacts produced by the preservation were rather consistent within each method during the whole 1 year experiment allowing these to be accounted for, if clams are intended for use in retrospective stable isotope studies.     

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and ¹³C/¹²C in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (δ ¹³C) of wood. The sensitivity of ring width, wood density and δ ¹³C to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July–September). δ ¹³C variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by δ ¹³C of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season δ ¹³C of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and ¹³C composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season δ ¹³C values of wood between individual trees (−31 to −24‰) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930–990 g cm⁻³). The relationship between seasonal changes in tree-ring structure and ¹³C composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20–30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction.     

秦岭不同龄组巴山冷杉径向生长对气候因子的响应差异

秦岭地区树轮气候学研究已经引起众多学者的重视,但年龄因素对调节树木径向生长-气候响应关系的研究尚未在当地得到广泛关注。运用树轮气候学方法,建立秦岭牛背梁国家级自然保护区低、中、高3个龄组巴山冷杉(Abies fargesii)的树轮宽度差值年表,探究不同龄组巴山冷杉径向生长与覆盖研究区的0.5°×0.5°分辨率格点气候要素之间的响应关系,以期解译年龄差异对巴山冷杉树木径向生长-气候响应特征的潜在影响。结果表明：不同龄组巴山冷杉差值年表统计特征值存在明显差异,平均敏感度和样本总体代表性随年龄增大而递减,但标准差、样本间平均相关系数和第一特征根变异解释量均以中龄组最低,信噪比却以中龄组最高;春季(3-5月)降水的增加以及秋季(9-11月)气温的升高对低龄组巴山冷杉径向生长的促进作用最强,对中龄组树木生长的促进作用明显减弱,而对高龄组树木生长几乎没有影响;气温对巴山冷杉径向生长的促进作用随树龄增高而增强,差值年表与当年2月、8月、上年5月平均温、平均最高温的相关系数均呈现随龄级增大而逐渐递增的趋势。年龄因素对牛背梁国家级自然保护区巴山冷杉径向生长-气候响应关系存在明确影响,随着年龄的增加,巴山冷杉径向生长受降水的制约减弱,受气温的限制增强。因此,在进行树木年轮学研究以及制定森林经营管理策略时应该考虑不同树龄巴山冷杉对气候的响应差异。     

Hydrogen and oxygen isotope ratios of tree-ring cellulose for riparian trees grown long-term under hydroponically controlled environments

Saplings of three riparian tree species (alder, birch and cottonwood) were grown for over 5 months in a hydroponics system that maintained the isotopic composition of source water in six treatments, ranging from –120 to +180‰δD and –15 to +10‰δ¹⁸O. The trees were grown in two greenhouses maintained at 25°C and at either 40 or 75% relative humidity, creating differences in transpiration rates and leaf water isotopic evaporative enrichment. The cellulose produced in the annual growth ring was linearly related to source water with differences in both slope and offset associated with greenhouse humidity. The slope of the isotopic composition of source water versus tree-ring cellulose was less than 1 for both δD and δ¹⁸O indicating incomplete isotopic exchange of carbohydrate substrate with xylem water during cellulose synthesis. Tests using the outer portion of the tree-ring and new roots were similar and showed that the tree-ring values were representative of the cellulose laid down under the imposed environmental conditions. The fraction of H and O in carbohydrate substrate that isotopically exchange with medium water was calculated to be 0.36 and 0.42 respectively, and biochemical mechanisms for these observed fractions are discussed. A mechanistic model of the biochemical fractionation events for both δD and δ¹⁸O leading to cellulose synthesis was robust over the wide range of cellulose stable isotope ratios. The experimental results indicate that both water source and humidity information are indeed recorded in tree-ring cellulose. These results help to resolve some of the disparate observations regarding the interpretation of stable isotope ratios in tree-rings found in the literature. Received: 4 January 1999 / Accepted: 12 August 1999