Influence of climatic factors upon tree rings of Larix decidua and L. decidua × L. kaempferi from Pulawy,Poland

Summary Dendroclimatological techniques are used to assess the impact of climatic factors on tree-ring width of Larix decidua and L. decidua × L. kaempferi (= L. x eurolepis) growing in two experimental plots established in 1914 in south-west Poland. One plot included F₁ progeny grown from seeds of an artificial crossing between European and Japanese larch. The other plot included progeny from maternal trees of European larch. Total ring width, earlywood width and latewood widths were dated, standardized and related to monthly climatic data using response function and stepwise multiple regression analyses. Wide rings in larch are associated with high precipitation in May–July, cool conditions in July–September of the preceding year, and cool dry conditions in August. Ring widths in L. x eurolepis are more dependent upon precipitation than ring widths in L. decidua. Latewood widths in L. x eurolepis are more dependent on high temperatures in June and July than latewood in L. decidua as well as total width and earlywood measurements. Variations in latewood were relatively independent of variations in earlywood and total wood. The variability of ring widths in these larches was greater than the variability reported for larches in many alpine sites and for other conifer species in some regions of North America.     

黑龙江漠河兴安落叶松与樟子松树轮生长特性及其对气候的响应

树木年轮 (简称树轮 ) 气候学是监测与重建全球气候变化的重要方法之一。针叶树树轮的生长能反馈出气温的变化, 在高纬度地带尤为明显。该文分析了生长在我国最北部的兴安落叶松 (Larixgmelinii) 与樟子松 (Pinussylvestrisvar.mongolica) 的树轮密度和宽度的特性。落叶松最大密度、晚材平均密度、早晚材宽度和轮宽都远高于樟子松。樟子松的所有密度变量的样本方差都明显高于兴安落叶松, 宽度变量的样本方差却明显低于兴安落叶松。两树种密度变量的差值年表显著相关, 宽度变量之间没有显著相关关系。落叶松与樟子松的晚材密度的形成受 7、8月的最高温控制。另外, 樟子松的晚材还与生长季节的长短相关。落叶松的年轮宽度对生长季节开始前的温度敏感, 而樟子松的轮宽对气候变量没有很好的响应。结果表明, 落叶松与樟子松的树轮最大密度都与生长季后期的温度显著相关, 两树种的树轮信息对气候变化的重建有很大的潜力。     

气候变暖背景下杉木年轮密度对气候因子的响应

为探讨杉木年轮密度与气候因子的响应关系,采用树木年轮学方法,以60年生杉木种源林为研究对象,测定杉木整轮密度、早材密度、晚材密度、晚材最大密度和早材最小密度,分析在气候变暖条件下主要气候因子（温度、降水、相对湿度）对杉木年轮密度及其生长的影响。结果表明：杉木不同年轮密度指标均受到温度、降水和相对湿度的显著影响。早材密度与当年夏季最高温度、当年5月降水量,最大密度与当年10月、当年秋季降水量,最小密度与前一年秋季降水量、最小相对湿度呈显著负相关。滑动相关分析表明气候因子在短时间尺度上对杉木生长影响的稳定性有显著影响,其中杉木年轮最大密度与当年10月、秋季的平均相对湿度和最小相对湿度,最小密度与当年2、3月的平均相对湿度和前一年秋季的平均相对湿度、最小相对湿度、降水量的负相关关系最为稳定。杉木年轮最小密度对前一年气候要素的响应存在滞后效应,且晚材密度对当年春季的气候要素响应也存在滞后效应。研究结果对开展亚热带针叶树种年轮生态学和年轮气候学研究具有重要参考价值,建议选择武夷山的天然林获取更长年表用于重建古气候。     

X-ray microdensitometry applied to subfossil tree-rings: growth characteristics of ancient pines from the southern boreal forest zone in Finland at intra-annual to centennial time-scales

A collection of subfossil wood of Pinus sylvestris (Scots pine) was exposed to X-ray densitometry. The collection of 64 samples from the southern boreal forest zone was dendrochronologically cross-dated to a.d. 673-1788. Growth characteristics were determined by performing density profiles including the following parameters: minimum density, earlywood and latewood boundary density, maximum density, earlywood width, earlywood density, latewood width, latewood density, annual ring width and annual ring density. Seven out of the nine parameters were found to contain non-climatic growth trends and six were found to be heteroscedastic in their variance. Tree-specific records were indexed, to remove the non-climatic growth trends and stabilize the variance, and combined into nine parameter-specific tree-ring chronologies. Growth characteristics of the pines changed in parallel with the generally agreed climatic cooling from the Medieval Warm Period to the Little Ice Age: pine tree-rings showed decreasing maximum densities from the period a.d. 975-1150 to a.d. 1450–1625. A concomitant change in the intra-annual growth characteristics was detected between these periods. The findings indicate that not only the trees growing near the species’ distributional limits are sensitive to large-scale climatic variations but also the trees growing in habitats remote from the timberline have noticeably responded to past climate changes.     

A tree-ring densitometric transect from Alaska to Labrador

We describe a recently completed network of densitometric tree-ring time series representing various aspects of tree-growth for up to 200 years at 69 sites spread across the northern North American conifer zone from Yukon to Labrador. Duplicate cores, from 12 to 15 trees per site, provide time series for a suite of growth parameters including earlywood (spring), latewood (summer) and total (annual) ring widths and mean earlywood, mean latewood, minimum and maximum ring density. These data form the basis for extensive analyses of intra- and inter-site parameter comparisons and regional climate/tree-growth comparisons. Five large-scale regional chronologies do not suggest that any anomalous growth increases have occurred in recent decades, at least on these regional scales, despite the observed changes in atmospheric composition and climate.     

Climate-tree-growth relationships of European beech (Fagus sylvatica L.) in the French Permanent Plot Network (RENECOFOR)

The influence of climate on the radial growth of Fagus sylvatica was investigated using 15 chronologies developed from mature stands of the French Permanent Plot Network (RENECOFOR) growing under different climatic and soil conditions. The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis. Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. The three most frequent negative pointer years (1959, 1989, 1976) result from a particularly intense and durable drought, whereas positive years (1977, 1958) coincide with wet conditions. The total ring chronology variance attributable to climate averages 34.1% (15.8% –57%). Current early-summer soil water deficit enters in 10 models and the deficit in June explains alone a large part of the radial growth variability (mean value: 26.6%). Temperature or soil water deficit for the other months and weather conditions during the previous season were of little consistency across stands. The response pattern of earlywood is very similar and the percentage of variance explained is higher (16.2% –57.8%). Latewood widths present a different response pattern. High minimum temperature in August and/or September often favour wide latewood widths and monthly water deficits play a secondary role. The percentage of variance explained ranges from 8.8% to 67.4%. Soil water capacity strongly modulates ring characteristics and climate-growth relationships. Mean sensitivity, expressed population signal, signal-to-noise ratio and the strength of growth-climate correlations increase with decreasing soil water capacity.     

Crossdating and analysis of eucalypt tree rings exhibiting terminal and reverse latewood

We investigated crossdating and climate sensitivity in tree-ring series from Eucalyptus delegatensis Baker, R.T. and E. obliqua L'Herit. We first visually crossdated the measured ring width series and then independently verified this crossdating using Xmatch and cross-correlation significance tests. Crossdating was verified in 28 of the 32 study trees. Crossdating success differed between E. delegatensis and E. obliqua. In E. delegatensis crossdating success appears to be related to tree dominance and elevation. In E. obliqua radial azimuth appears to affect crossdating success. We developed two chronologies for each of the species studied. The first of these chronologies was based on all visually crossdated radii and the other on radii for which crossdating had been independently verified. Signal strength was higher in the verified chronologies. Correlation analysis between the verified chronologies and climate data revealed no significant correlation between precipitation and ring width for either species. E. obliqua ring width was significantly correlated with mean minimum and maximum air temperature and vapour pressure deficit during summer of the growing season. The E. delegatensis chronology was significantly correlated with air temperature and frequency of frost during the preceding winter. Ring width in both species was significantly correlated with air temperature during the preceding summer. Potential physiological explanations for these results are discussed. Further study is required to verify the results of climatological analysis and to explore the causes of variation in signal strength within and between trees.     

Long-term changes in wood density and radial growth of Quercus petraea Liebl. in northern France since the middle of the nineteenth century

Long-term changes in sessile oak (Quercus petraea Liebl.) growth and wood density were studied using cores collected from 99 even-aged high forest stands between 56 and 187 years old, located in northeastern and north-central France. Growth and density trends were tested by analysis of variance and covariance. Two models were applied to two samples, sample A and sample B (sample B being a sub-sample with limited cambial age and calendar date ranges). Model 1 showed a significant increase in radial growth: +35%, +87% and +66% in earlywood width, latewood width and ring width, respectively, from 1811 to 1993 for sample A. Consequently, there was a positive trend in latewood ratio (+14%). A slight decrease in wood density was found: -3.3% and -5.4% for earlywood and latewood density, respectively. Despite an increase in latewood percentage, mean ring density showed a -2.0% decrease. Model 1 applied to a biomass indicator (density2ring width) showed a 62% increase from 10.4 to 16.8 kg m^-3 between 1811 and 1993 for sample A. Results for sample B were slightly different: the increase in latewood ratio was not detected. Model 2 showed a change with time in the positive hyperbolic relationship between mean density and ring width. The results are discussed. The decrease in wood density cannot be explained by N atmospheric deposition or by long-term changes in average temperature. Increasing atmospheric CO₂ levels cannot be invoked owing to the present lack of studies. Finally, hypotheses concerning long-term changes in wood anatomical characteristics are proposed.     

Chronologies of earlywood vessels and latewood width disentangle climate drivers of oak growth in a mild oceanic region

We obtained tree-ring chronologies for pedunculate oak along an altitudinal gradient of four sites in NW Iberia, which covers most of the species range within the region. Trees grow under Atlantic climate, with a rainfall maximum in autumn-winter, and a minimum during summer, but lacking a remarkable drought. Chronologies included several earlywood anatomical features and ring widths, which were compared to each other. Latewood width and a subset of six earlywood variables were selected for further analyses, considering the period 1954–2003.The lowest site considerably differed from the rest of the gradient due to its milder conditions. Earlywood and latewood were unrelated at the three upper sites, and both compartments responded to different climatic factors. Hydraulic conductivity determined by vessel size was used to describe tree performance, which was modulated by temperature along the gradient. We hypothesize that the main processes involved are the timing of earlywood formation and carbohydrate dynamics. Water availability during late spring and summer affected latewood width, but only at low elevation.This paper illustrates the complexity of analyzing climate-growth relationships in oceanic areas under the absence of a prevailing limiting factor, while providing a feasible explanation of potential mechanisms involved.     

Climate–tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA

Knowledge of tree growth/climate response relationships is important to dendroecological studies and dendroclimatic reconstructions, particularly in the Southeastern Coastal Plain where few such studies have been attempted. To this end, we developed tree-ring chronologies of total ring width, earlywood width, and latewood width from longleaf pine (Pinus palustris Mill.) at three sites in the Southeastern Coastal Plain to examine the climate–growth relationships for this tree species. The length of these chronologies is unprecedented for southern pine chronologies in the Southeast. We compared the tree-ring chronologies to monthly temperature, precipitation, Palmer drought severity index (PDSI), and Palmer hydrological drought index (PHDI) data from the pertinent climate divisions. We found that PDSI and PHDI have the highest correlation with longleaf pine growth, and the strongest relationships between longleaf pine growth and these variables occur between July and November. Precipitation in the spring and summer was also positively related to growth at all sites. The relationship between temperature and growth was the weakest among all climate variables, but warm summer temperatures had a consistent, negative relationship with longleaf pine growth. The climate signal in the latewood was generally more robust than for total ring width and earlywood width.     

A simple program to measure and analyse tree rings using Excel,R and SigmaScan

I present a new software that links a program for image analysis (SigmaScan), one for spreadsheets (Excel) and one for statistical analysis (R) for applications of tree-ring analysis. The first macro measures ring width marked by the user on scanned images, stores raw and detrended data in Excel and calculates the distance to the pith and inter-series correlations. A second macro measures darkness along a defined path to identify latewood–earlywood transition in conifers, and a third shows the potential for automatic detection of boundaries. Written in Visual Basic for Applications, the code makes use of the advantages of existing programs and is consequently very economic and relatively simple to adjust to the requirements of specific projects or to expand making use of already available code.     

Couplings in cell differentiation kinetics mitigate air temperature influence on conifer wood anatomy

Conifer trees possess a typical anatomical tree‐ring structure characterized by a transition from large and thin‐walled earlywood tracheids to narrow and thick‐walled latewood tracheids. However, little is known on how this characteristic structure is maintained across contrasting environmental conditions, due to its crucial role to ensure sap ascent and mechanical support. In this study, we monitored weekly wood cell formation for up to 7 years in two temperate conifer species (i.e., Picea abies (L.) Karst and Larix decidua Mill.) across an 8°C thermal gradient from 800 to 2,200 m a.s.l. in central Europe to investigate the impact of air temperature on rate and duration of wood cell formation. Results indicated that towards colder sites, forming tracheids compensate a decreased rate of differentiation (cell enlarging and wall thickening) by an extended duration, except for the last cells of the latewood in the wall‐thickening phase. This compensation allows conifer trees to mitigate the influence of air temperature on the final tree‐ring structure, with important implications for the functioning and resilience of the xylem to varying environmental conditions. The disappearing compensation in the thickening latewood cells might also explain the higher climatic sensitivity usually found in maximum latewood density.     

Variations in Environmental Signals in Tree-Ring Indices in Trees with Different Growth Potential

We analysed two groups of Quercus robur trees, growing at nearby plots with different micro-location condition (W-wet and D-dry) in the floodplain Krakovo forest, Slovenia. In the study we compared the growth response of two different tree groups to environmental variables, the potential signal stored in earlywood (EW) structure and the potential difference of the information stored in carbon isotope discrimination of EW and latewood (LW). For that purpose EW and LW widths and carbon isotope discrimination for the period 1970–2008 AD were measured. EW and LW widths were measured on stained microscopic slides and chronologies were standardised using the ARSTAN program. α-cellulose was extracted from pooled EW and LW samples and homogenized samples were further analysed using an elemental analyser and IRMS. We discovered that W oaks grew significantly better over the whole analysed period. The difference between D and W oaks was significant in all analysed variables with the exception of stable carbon isotope discrimination in latewood. In W oaks, latewood widths correlated with summer (June to August) climatic variables, while carbon isotope discrimination was more connected to River Krka flow during the summer. EW discrimination correlated with summer and autumn River Krka flow of the previous year, while latewood discrimination correlated with flow during the current year. In the case of D oaks, the environmental signal appears to be vague, probably due to less favourable growth conditions resulting in markedly reduced increments. Our study revealed important differences in responses to environmental factors between the two oak groups of different physiological conditions that are preconditioned by environmental stress. Environmental information stored in tree-ring features may vary, even within the same forest stand, and largely depends on the micro-environment. Our analysis confirmed our assumptions that separate EW and LW analysis of widths and carbon isotope discrimination provides complementary information in Q. robur dendroecology.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

Comparison of methods for the demarcation between earlywood and latewood in tree rings of Norway spruce

The precise demarcation between earlywood and latewood is important for the detailed analysis of intra-annual tree ring features. Different techniques based on visual assessment, wood anatomy analysis and X-ray densitometry have been developed and are currently used for this purpose. Depending on the chosen method, tree species and environmental conditions, the results can significantly vary. Thus, it is important to determine the technique optimal for a particular research. Here, we investigated Norway spruce (Picea abies) tree rings to examine the agreement among the following demarcation methods: (1) direct visual assessment, (2) Mork’s index (anatomical definition of the transition from earlywood to latewood based on cell wall-lumen ratio) and (3) fixed and floating density thresholds applied to intra-ring density profiles. The aim was to modify both the Mork’s criterion and density thresholds on the basis of reference values given by visual identification of earlywood/latewood transition. A total of 231 tree rings were analysed by all methods. Our results showed that the usage of floating threshold (defined for each ring separately based on density profiles) is more reliable in comparison with fixed threshold (the same threshold value used for all tree rings and samples). Statistical analysis revealed the best correspondence between visual identification of earlywood/latewood transition and demarcation based on the standard Mork’s index and the floating density threshold derived as 80 % of maximum latewood density. In terms of Mork’s index calibration, the results showed that to determine latewood cells in Norway spruce trees growing in temperate conditions, it is sufficient to use an index value equal to 0.83. The results are applicable for the studied spruce population growing in a temperate climate. The methodology itself, however, is universal and can help to calibrate criteria for earlywood-latewood demarcation under specific conditions.     

The utility of bulk wood density for tree-ring research

Bulk wood density measurements are recognized for their utility in ecology, industry, and biomass estimations. In tree-ring research, microdensitometric techniques are widely used, but their ability to determine the correct central tendency has been questioned. Though rarely used, it may be possible to use bulk wood density as a tool to check the accuracy of and even correct microdensitometric measurements. Since measuring bulk wood density in parallel with X-ray densitometry is quickly and easily done, we suspect that its omission is largely due to a lack of awareness of the procedure and/or its importance. In this study, we describe a simple protocol for measuring bulk wood density tailored for tree-ring researchers and demonstrate a few possible applications. To implement real-world examples of the applications, we used a sample of existing X-ray and Blue Intensity (BI) measurements from 127 living and dead Pinus sylvestris trees from northern Sweden to produce new measurements of bulk wood density.We can confirm that the central tendency in this sample material is offset using X-ray densitometry and that the diagnosis and correction of X-ray density is easily done using bulk wood density in linear transfer functions. However, this approach was not suitable for our BI measurements due to heavy discoloration. Nevertheless, we were able to use bulk wood density to diagnose and improve the use of deltaBI (latewood BI – earlywood BI) with regard to its overall trends and multi-centennial variability in a dendroclimatological application. Moreover, we experimented with percent of latewood width, scaled with bulk wood density, as a time- and cost-effective proxy for annual ring density. Although our reconstruction only explains about half of the variation in ring density, it is most likely superior to using fixed literature values of density in allometric equations aimed at biomass estimations.With this study, we hope to raise new awareness regarding the versatility and importance of bulk wood density for dendrochronology by demonstrating its simplicity, relevance, and applicability.     

Effect of climate on the growth of annual rings in the main roots of perennial forbs in an Inner Mongolian semi‐arid grassland,China

Question: Is there a pattern in growth of annual rings in roots of perennial forbs in relation to climate and climate extremes in grassland ecosystems? Location: Semi‐arid grassland in Duolun (42°27′N, 116°41′E, 1380 m a.s.l.), central Inner Mongolia, China. Methods: Main roots of three perennial species, Potentilla anserina L., Cymbaria dahurica L. and Lespedeza daurica Schindl., were sampled. Cross‐sections (10–15‐μm thick) were produced from the proximal end of sampled roots using a sledge microtome. Annual growth rings in the main roots were identified and measured by differentiating between earlywood and latewood in the secondary xylem. Relationships between annual growth rings and monthly mean temperature and total monthly precipitation were identified using correlation analysis. Differences in an annual ring width to the previous and following years were examined by calculating a distinctness score. Results: The three perennial forbs showed clearly demarcated annual growth rings in all individuals and the same fluctuation patterns. Their ring widths were generally positively correlated with precipitation from April to October (except for August) and with temperature from February to June (except June for L. daurica), September to October, and the annual mean. Strong deviations of annual ring widths from their neighbour rings were observed in 1998 and 2000. The trend of absolute distinctness scores (D_m) increased significantly from 1988 to 2003, indicating an increase in the frequency of annual ring width variation. Conclusions: Annual growth rings in the main roots of three perennial forb species can be used as an indicator of the influence of climate on below‐ground grassland growth. The change in below‐ground conditions and effects on the functioning of grassland should receive more attention in future studies.     

Responses to climate by tree-ring widths and maximum latewood densities of two Abies species at upper and lower altitudinal distribution limits in central Japan

This study examined the effects of climate on tree-ring widths and maximum latewood densities of Abies veitchii and Abies mariesii at the upper and lower distribution limits in central Japan. A. veitchii and A. mariesii dominated at the lower and upper parts of the subalpine zone, respectively. Residual chronologies of tree-ring width and maximum latewood density were developed for the two Abies species at the upper and lower distribution limits, and were compared with monthly mean temperatures and monthly sums of precipitation. Tree-ring widths of the two Abies species at the upper and lower distribution limits positively correlated with temperatures during the beginning of the dormant season and during the growing season of the current year, except for A. veitchii at the lower distribution limit, which showed no positive correlation with temperature. Maximum latewood densities of the two Abies species at the upper and lower distribution limits positively and negatively correlated with temperatures and precipitation, respectively, during the growing season of the current year. Therefore, tree-ring widths and maximum latewood densities of the two Abies species were sensitive to low temperature, except for the tree-ring width of A. veitchii at the lower distribution limit with the warmest thermal conditions along the altitude. Global warming is suggested to affect maximum latewood densities and tree-ring widths of the two Abies species along the altitude.     

Influence of climate on tree-ring and earlywood vessel formation in Quercus robur in Latvia

We studied the effects of climatic factors on tree-ring width and vessel lumen area (VLA) in earlywood of English oak (Quercus robur L.) in Latvia. Cores were obtained from healthy canopy oaks in 40 stands located across Latvia. Tree-ring widths and VLA were measured. Principal component analysis was used to arrange the sites along gradients of response of tree-ring width and earlywood to environmental factors. Significant relationships of tree-ring width and mean VLA with climatic factors (mean monthly temperature and precipitation sum) were determined by correlation analysis. Relationships between tree-ring, early- and latewood widths were tested in three sampled stands. The patterns of response of VLA and tree-ring width to environmental factors differed in relation to a west–east gradient of increasing continentality. Three regions of Latvia (western, central and eastern) were distinguished along this gradient. Responses to climate differed between tree-ring width and mean VLA. Occurrence of significant correlations between climatic factors and the proxies differed between regions, likely due to regional differences in temperature and precipitation. Tree-ring width correlated with climatic factors (most commonly with March, May and June temperature and August precipitation of the current growing season and July–August temperatures of the previous growing season); VLA was more strongly related to climatic factors, particularly with temperature in winter and spring months. The proportion of significant correlation coefficients with climatic factors differed between the regions. Among sites, significant correlation of tree-ring width with temperature in spring and summer was more frequent in the western region, while correlation with winter temperature of the previous growing season and precipitation in August was more frequent in the eastern region. For VLA, the frequency of significant correlation coefficients with temperature in winter and spring was higher in the eastern region.     

Seasonal and perennial changes in the distribution of water in the sapwood of conifers in a sub-frigid zone

An analysis was made of progressive changes in patterns of cavitation in the sapwood of three species of conifer (Larix kaempferi, Abies sachalinensis, and Picea jezoensis) that were growing in a sub-frigid zone. In all three conifers, all tracheids of the newly forming outermost annual ring were filled with water or cytoplasm during the period from May to August. However, many tracheids in the transition zone from earlywood to latewood lost water in September, presumably through drought-induced cavitation. Cavitated tracheids tended to be continuously distributed in a tangential direction. Subsequently, some earlywood tracheids of the outermost annual ring lost water during the period from January to March. This was associated with freeze-thaw cycles. In the second and third annual rings from the cambium of all three conifers, the lumina of most tracheids in the transition zone from earlywood to latewood contained no water. In contrast, some latewood tracheids near the annual ring boundary and many earlywood tracheids retained water in their lumina. The third annual ring had more cavitated tracheids than the second annual ring. Our observations indicated that cavitation progressed gradually in the tracheids of the conifers and that they were never refilled once cavitation had occurred. The region involved in water transport in conifers did not include the entire sapwood and differed among annual rings.