Stem Radius Changes Research Articles

Using thermal imagery and changes in stem radius to assess water stress in two coniferous tree species

With a warming climate and greater evaporative demand, many forest ecosystems are increasingly affected by water limitation as prolonged water deficits reduce tree-level growth and survival. Water deficit can be monitored in several ways, including from daily measurements of sap flow or changes in stem radius from automated sensors mounted on individual trees. As an alternative approach, we evaluated the use of airborne thermal imagery from unmanned aerial vehicles as a rapid, scalable tool for assessing tree-level water stress. Plant water stress leads to higher leaf temperatures when soil moisture is low and evaporative demand is high. To detect this response, we modeled the difference between leaf and air temperature (∆T) as a function of local soil moisture, vapor pressure deficit, and wind speed for two tree species, lodgepole pine (Pinus contorta var. latifolia) and white spruce (Picea glauca). We used those same weather and soil conditions to model dendrometer-based measurements of daily changes in internal tree water deficit (∆TWD). While canopy leaf temperature and daily change in tree water deficit showed little direct correlation with one another, these variables both responded to soil moisture, vapor pressure deficit, and wind speed in a manner that reflects responses to water stress as soil became progressively drier over the summer months. The two species showed some differences related to species-specific strategies for drought avoidance. The application of thermal imagery to detect water stress in natural forest ecosystems can improve understanding of how trees experience water stress across species and environmental conditions.

Climatic control of high-resolution stem radius changes in a drought-limited southern boreal forest

Key messageStem radius changes measured at high temporal resolution in a drought-limited southern boreal forest were primarily related to soil temperature and partly soil moisture variation, but only weakly to VPD.Forest productivity at the southern fringe of the boreal forest biome in Inner Asia is strongly drought-limited, as is evident from dendrochronological analyses. Using electronic point dendrometers, we studied the climate response of stem radius changes at high temporal resolution (10-min intervals) in a mixed larch–birch forest in northern Mongolia in a drought year and two subsequent moist years. Larch trees showed stronger stem radius fluctuations than birches, and this difference was more pronounced in dry than in moist years. Stem radius changes were most tightly related to soil temperature variation, while soil moisture was the dominant controlling factor only in birch in the dry year. Correlations with the atmospheric vapor pressure deficit (VPD), and even more so with air temperature, were much weaker. While the linkage between radius change and VPD might primarily reflect diurnal transpiration-driven stem shrinkage and expansion, soil temperature is thought to directly affect cambial cell division and elongation during wood formation. We conclude that the phenology of stemwood increment is strongly controlled by soil temperature even in drought-limited southern boreal forests under continental climate due to the combination of cold and dry climate.

Reconstructing radial stem size changes of trees with machine learning.

Like many scientists, ecologists depend heavily on continuous uninterrupted data in order to understand better the object of their study. Although this might be straightforward to achieve under controlled laboratory conditions, the situation is easily complicated under field conditions where sensors and data transmission are affected by harsh weather, living organisms, changes in atmospheric conditions etc. This often results in parts of the data being corrupted or missing altogether. We propose the use of the most recent machine-learning techniques to reverse such data losses in multi-channel time series. In particular, we focus on tree stem growth data obtained from the TreeNet project, which monitors the changes in stem radius and environmental conditions of a few hundred trees across Switzerland. In the first part of the study, we test the performance of five architectures based on encoders and recurrent and convolutional neural networks, and we show that a deep neural network combining long short-term memory with one-dimensional convolutional layers performs the best. In the second part, we adopt this model to reconstruct the original TreeNet dataset, which we then use in a separate classification problem to show the effect of the proposed gap-filling procedure.

Snowmelt Water Use at Transpiration Onset: Phenology, Isotope Tracing, and Tree Water Transit Time

AbstractStudies of tree water source partitioning have primarily focused on the growing season. However, little is yet known about the source of transpiration before, during, and after snowmelt when trees rehydrate and recommence transpiration in the spring. This study investigates tree water use during spring snowmelt following tree's winter stem shrinkage. We document the source of transpiration of three boreal forest tree species—Pinus banksiana, Picea mariana, and Larix laricina—by combining observations of weekly isotopic signatures (δ18O and δ2H) of xylem, soil water, rainfall and snowmelt with measurements of soil moisture dynamics, snow depth and high‐resolution temporal measurements of stem radius changes and sap flow. Our data shows that the onset of stem rehydration and transpiration overlaps with snowmelt for evergreens. During rehydration and transpiration onset, xylem water at the canopy reflected a constant pre‐melt isotopic signature likely showing late fall conditions. As snowmelt infiltrates the soil and recharges the soil matrix, soil water shows a rapid isotopic shift to depleted‐snowmelt water values. While there was an overlap between snowmelt and transpiration timing, xylem and soil water isotopic values did not overlap during transpiration onset. Our data showed 1–2‐week delay in the shift in xylem water from pre‐melt to clear snowmelt‐depleted water signatures in evergreen species. This delay appears to be controlled by tree water transit time that was in the order of 9–18 days. Our study shows that snowmelt is a key source for stem rehydration and transpiration in the boreal forest during spring onset.

High growth recovery ability of Eucalyptus grandis trees following a 3-year period of 80% throughfall reduction

Eucalyptus plantations are already planted in - or expanding to - water-limited regions with a high risk of severe drought. In future drier and more variable climate including extreme events, the ability of trees to recover after severe droughts emerges as a crucial for the sustainability of forest plantations.An original experiment involving 80% reduction in throughfall was set up in Brazil to gain insight into the responses of Eucalyptus grandis trees to prolonged (3-year) extreme water deficit and the ability of this species to recover following water stress. Our study focused on the changes in basal area, stem radius, and total height measured by high-temporal resolution dendrometers and periodical surveys of trees affected by 80% throughfall reduction (treatment group) and in a control group. The differences in basal area, stem radius, and total height growth rate were compared between groups over (i) 37 months of 80% throughfall reduction and (ii) 31 months following the end of 80% throughfall reduction. The correlations among growth rates, stem radius fluctuations, and meteorological variables in each group were determined to gain insights into the trees’ responses to environmental conditions and stem water status.The 80% reduction in throughfall over 3 years significantly reduced tree growth rates by 73% in basal area and 95% in total height. However, under normal water availability following throughfall reduction, the basal area growth rate of water-stressed trees was 97% higher than that of the control trees while total height growth rate was only 8% higher. Despite the severe water stress, no tree mortality was observed. Trees recovered 51% of their basal area over the 31-month recovery period. In contrast, only 5% of total height was recovered.In the treatment group, rapid responses to variation in rainfall events were observed during the 80% throughfall reduction period. Also, correlations between stem radius fluctuations and vapor pressure deficit indicate increased transpiration following the end of throughfall reduction. These relationships indicate high conservation of the integrity of the xylem vascular system over the 3 years of 80% throughfall reduction, a key factor in the increased resilience of the trees. In the absence of tree mortality, our results suggest the 80% throughfall reduction had a severe impact on tree growth but demonstrate great recovery ability of Eucalyptus grandis trees in basal area growth following such a severe water deficit.

Linking variability of tree water use and growth with species resilience to environmental changes

Tree growth is an indicator of tree vitality and its temporal variability is linked to species resilience to environmental changes. Second‐order statistics that quantify the cross‐scale temporal variability of ecophysiological time series (statistical memory) could provide novel insights into species resilience. Species with high statistical memory in their tree growth may be more affected by disturbances, resulting in lower overall resilience and higher vulnerability to environmental changes. Here, we assessed the statistical memory, as quantified with the decay in standard deviation with increasing time scale, in tree water use and growth of co‐occurring European larch Larix decidua and Norway spruce Picea abies along an elevational gradient in the Swiss Alps using measurements of stem radius changes, sap flow and tree‐ring widths. Local‐scale interspecific differences between the two conifers were further explored at the European scale using data from the International Tree‐Ring Data Bank. Across the analysed elevational gradient, tree water use showed steeper variability decay with increasing time scale than tree growth, with no significant interspecific differences, highlighting stronger statistical memory in tree growth processes. Moreover, Norway spruce displayed slower decay in growth variability with increasing time scale (higher statistical memory) than European larch; a pattern that was also consistent at the European scale. The higher statistical memory in tree growth of Norway spruce in comparison to European larch is indicative of lower resilience of the former in comparison to the latter, and could potentially explain the occurrence of European larch at higher elevations at the Alpine treeline. Single metrics of resilience cannot often summarize the multifaceted aspects of ecosystem functioning, thus, second‐order statistics that quantify the strength of statistical memory in ecophysiological time series could complement existing resilience indicators, facilitating the assessment of how environmental changes impact forest growth trajectories and ecosystem services.

Similar diurnal, seasonal and annual rhythms in radial root expansion across two coexisting Mediterranean oak species.

Dendrometers are being increasingly used to measure stem radius changes in trees and to unravel the mechanisms underlying stem daily rhythms of radial expansion and contraction. Nevertheless, automated dendrometers have not been often used to measure root radius dynamics, their relationship with environmental variables and the influence of endogenous processes, especially in drought-prone Mediterranean areas. Here, we measured root radius dynamics of two coexisting oak species (the evergreen Quercus ilex L. and the deciduous Quercus faginea Lam). Our goals were to describe annual, seasonal and diurnal scale root radius patterns and to disentangle the role of different environmental parameters as drivers. Long-term high-resolution measurements (every 15min over 7years) were collected with automated point dendrometers on the main tree roots of five individuals per species. Root radius annual change patterns were bimodal and similar for both oak species. Quercus faginea Lam showed three times larger root increment in the spring than Q. ilex, but the bimodal pattern was stronger in Q. ilex, which showed a larger root increment in autumn. Quercus faginea Lam showed an earlier root phenological activation in the spring and in late summer compared with Q. ilex. The effects of environmental drivers across species were similar at daily scales: root radius increased with air temperature and soil moisture, and it decreased with rising vapor pressure deficit. Furthermore, daily root radius variations for both oak species were maintained after extracting statistically the environmental effects, which points toward a significant role of endogenous drivers. These differences in root radius change patterns at seasonal to daily scales likely result from the differences in leaf phenology and growth strategy. Quercus faginea Lam is deciduous and has a faster growing rate in spring than the evergreen Q. ilex, which can grow more in summer.

Daytime stem swelling and seasonal reversal in the peristaltic depletion of stored water along the stem of Avicennia marina (Forssk.) Vierh.

Diurnal courses of stem radial water dynamics represent the sum of all internal and external conditions affecting tree water relations. Changes in stem radius due to early morning water depletion and night time refilling of storage tissues is generally well documented. This study seeks to understand the unusual daytime refilling of stem elastic storage tissues present in the mangrove species Avicennia marina (Forssk.) Vierh, which deviates from our traditional understanding of hydraulics in terrestrial trees. We explored the relationship of this pattern to other water-related physiological processes and environmental variables, and investigated the seasonal changes in the timing and time lags of peak swelling at different stem heights, in order to understand the 'peristaltic' depletion of internally stored water. Our findings show that daytime stem swelling occurs year-round, even on days when leaf water potentials dropped to values lower than -4 MPa. The amplitude of stem swelling was strongly positively correlated to daily light sums more often than to measures of water availability in air and soil, especially in winter. There was also a clear seasonal reversal in the timing and direction of the 'peristaltic' depletion of water along the stem, with an earlier onset of shrinking in the upper (median = 10:00 h) than in the lower stem (median = 12:00 h) in winter, but an earlier onset of shrinking in the lower (median = 08:00 h) than in the upper stem (median = 11:00 h) in summer. This time lag was closely correlated to daily temperature, with a clear switch in the direction of peristaltic stem shrinking at the start of the growing season. We propose that sugar loading/unloading and changes in source-sink activity play a role in the endogenous osmotic adjustment responsible for daytime stem swelling and the seasonal switch in the direction of peristaltic water storage depletion in A. marina.

Leaf water maintains daytime transpiration in young Cryptomeria japonica trees.

Compared with stem water storage, leaf water storage is understudied although it may be important for alleviating water stress by contributing quickly and directly to transpiration demand. To quantify the relative contribution of stem versus leaf water storage to daily water deficit, we measured diurnal changes in transpiration rate, sap-flow velocity and stem radius of 10-year-old Cryptomeria japonica D. Don trees. We assumed that the duration of time lags between transpiration rate and sap-flow velocity reflected stored water in the stem and leaf, and that stem volume change represented water content of elastic tissue. The relationship between fresh mass and water potential of the whole tree indicated that the study trees had capacity to store, on average, 91.4 ml of water per kg fresh mass at turgor loss. Leaves, sapwood and elastic tissue contributed around 51%, 29% and 20% of stored water, respectively. During morning, transpiration rates were higher than sap-flow velocity suggesting depletion of stored water. During the first 2 h after onset of transpiration, stored water contributed more than 100% of whole-tree transpiration. Depletion of leaf water (PLeaf) and sapwood water (PSap) coincided with the onset of transpiration and became maximum around 15:00 h. Depletion of elastic tissue water (PElastic) lagged behind that of PLeaf and PSap by 1-2 h, indicating that replenishment of stored water occurs late in the day when low leaf water potentials resulting from daytime transpiration drive water uptake. Maximum depletion of PLeaf was about 1-3 times and 5-10 times that of PSap and PElastic, respectively. The contribution of PLeaf to total daily transpiration was 5-8%, while those of PSap and PElastic were 3-4% and 0.7-1%, respectively. Our results suggest the importance of leaf water storage in maintaining daily transpiration in young C. japonica trees.

Temperate tree species show identical response in tree water deficit but different sensitivities in sap flow to summer soil drying.

Temperate forests are expected to be particularly vulnerable to drought and soil drying because they are not adapted to such conditions and perform best in mesic environments. Here we ask (i) how sensitively four common temperate tree species (Fagus sylvatica, Picea abies, Acer pseudoplatanus and Fraxinus excelsior) respond in their water relations to summer soil drying and seek to determine (ii) if species-specific responses to summer soil drying are related to the onset of declining water status across the four species. Throughout 2012 and 2013 we determined tree water deficit (TWD) as a proxy for tree water status from recorded stem radius changes and monitored sap flow rates with sensors on 16 mature trees studied in the field at Lägeren, Switzerland. All tree species responded equally in their relative maximum TWD to the onset of declining soil moisture. This implies that the water supply of all tree species was affected by declining soil moisture and that none of the four species was able to fully maintain its water status, e.g., by access to alternative water sources in the soil. In contrast we found strong and highly species-specific responses of sap flow to declining soil moisture with the strongest decline in P. abies (92%), followed by F. sylvatica (53%) and A. pseudoplatanus (48%). F. excelsior did not significantly reduce sap flow. We hypothesize the species-specific responses in sap flow to declining soil moisture that occur despite a simultaneous increase in relative TWD in all species reflect how fast these species approach critical levels of their water status, which is most likely influenced by species-specific traits determining the hydraulic properties of the species tree.

Radial stem variations - a source of tree physiological information not fully exploited yet.

Radial stem variations - a source of tree physiological information not fully exploited yet.

Xylem as the main origin of stem radius changes in Eucalyptus

The state-of-the-art interpretation of stem radius changes (DRTotal) for tree water relations is based on knowledge from mostly slow growing tree species. The ratio between diurnal size fluctuations of the rigid xylem (DRXylem) and the respective fluctuations of the elastic bark (DRBark) is known to be small (<0.4) and is of importance for the localisation of water storage dynamics in stems. In this study, fast growing Eucalyptus globulus Labill. in Tasmania were investigated by point dendrometers in order to investigate tree water relations. Unexpectedly, DRXylem was found to be the main driver of DRTotal with the bark acting as a passive layer on top of the fluctuating xylem under most conditions. Accordingly, the ratio between the diurnal fluctuations of the two tissues was found to be much higher (0.6-1.6) than everything reported before. Based on simulations using a hydraulic plant model, the high tissue-specific elasticity of the Eucalyptus xylem was found to explain this atypical response and not osmotically-driven processes or species-specific flow resistances. The wide zone of secondary thickening xylem in various stages of lignification is proposed to be an important component of the high wood elasticity. The tissue acts as additional water storage like the bark and may positively affect the water transport efficiency.

Stem water storage in five coexisting temperate broad-leaved tree species: significance, temporal dynamics and dependence on tree functional traits

The functional role of internal water storage is increasingly well understood in tropical trees and conifers, while temperate broad-leaved trees have only rarely been studied. We examined the magnitude and dynamics of the use of stem water reserves for transpiration in five coexisting temperate broad-leaved trees with largely different morphology and physiology (genera Fagus, Fraxinus, Tilia, Carpinus and Acer). We expected that differences in water storage patterns would mostly reflect species differences in wood anatomy (ring vs. diffuse-porous) and wood density. Sap flux density was recorded synchronously at five positions along the root-to-branch flow path of mature trees (roots, three stem positions and branches) with high temporal resolution (2 min) and related to stem radius changes recorded with electronic point dendrometers. The daily amount of stored stem water withdrawn for transpiration was estimated by comparing the integrated flow at stem base and stem top. The temporal coincidence of flows at different positions and apparent time lags were examined by cross-correlation analysis. Our results confirm that internal water stores play an important role in the four diffuse-porous species with estimated 5-12 kg day(-1) being withdrawn on average in 25-28 m tall trees representing 10-22% of daily transpiration; in contrast, only 0.5-2.0 kg day(-1) was withdrawn in ring-porous Fraxinus. Wood density had a large influence on storage; sapwood area (diffuse- vs. ring-porous) may be another influential factor but its effect was not significant. Across the five species, the length of the time lag in flow at stem top and stem base was positively related to the size of stem storage. The stem stores were mostly exhausted when the soil matrix potential dropped below -0.1 MPa and daily mean vapor pressure deficit exceeded 3-5 hPa. We conclude that stem storage is an important factor improving the water balance of diffuse-porous temperate broad-leaved trees in moist periods, while it may be of low relevance in dry periods and in ring-porous species.

Long‐term stem CO2 concentration measurements in Norway spruce in relation to biotic and abiotic factors

Stem CO(2) concentrations (stem [CO(2)]) undergo large temporal variations that need to be understood to better link tree physiological processes to biosphere-atmosphere CO(2) exchange. During 19 months, stem [CO(2)] was continuously measured in mature subalpine Norway spruce trees (Picea abies) and jointly analysed with stem, soil and air temperatures, sap flow rates, stem radius changes and CO(2) efflux rates from stem and soil on different time scales. Stem [CO(2)] exhibited a strong seasonality, of which over 80% could be explained with stem and soil temperatures. Both physical equilibrium processes of CO(2) between water and air according to Henry's law as well as physiological effects, including sap flow and local respiration, concurrently contributed to these temporal variations. Moreover, the explanatory power of potential biological drivers (stem radius changes, sap flow and soil respiration) varied strongly with season and temporal resolution. We conclude that seasonal and daily courses of stem [CO(2)] in spruce trees are a combined effect of physical equilibrium and tree physiological processes. Furthermore, we emphasize the relevance of axial diffusion of CO(2) along air-filled spaces in the wood, and potential wound response processes owing to sensor installation.

Link between continuous stem radius changes and net ecosystem productivity of a subalpine Norway spruce forest in the Swiss Alps

*Continuous stem radius changes (DR) include growth and water-related processes on the individual tree level. DR is assumed to provide carbon turnover information complementary to net ecosystem productivity (NEP) which integrates fluxes over the entire forest ecosystem. Here, we investigated the unexpectedly close relationship between NEP and DR and asked for causalities. *NEP (positive values indicate carbon sink) measured by eddy covariance over 11 yr was analysed at three time scales alongside automated point dendrometer DR data from a Swiss subalpine Norway spruce forest. *On annual and monthly scales, the remarkably close relationship between NEP and DR was positive, whereas on a half-hourly scale the relationship was negative. Gross primary production (GPP) had a similar explanatory power at shorter time scales, but was significantly less correlated with DR on an annual scale. *The causal explanation for the NEP-DR relationship is still fragmentary; however, it is partially attributable to the following: radial stem growth with a strong effect on monthly and annual increases in NEP and DR; frost-induced bark tissue dehydration with a parallel decrease in both measures on a monthly scale; and transpiration-induced DR shrinkage which is negatively correlated with assimilation and thus with NEP on a half-hourly scale.

Ultrasonic acoustic emissions in drought‐stressed trees – more than signals from cavitation?

Ultrasonic acoustic emission (UAE) in trees is often related to collapsing water columns in the flow path as a result of tensions that are too strong (cavitation). However, in a decibel (dB) range below that associated with cavitation, a close relationship was found between UAE intensities and stem radius changes. UAE was continuously recorded on the stems of mature field-grown trees of Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens) at a dry inner-Alpine site in Switzerland over two seasons. The averaged 20-Hz records were related to microclimatic conditions in air and soil, sap-flow rates and stem-radius fluctuations detrended for growth (Delta W). Within a low-dB range (27 +/- 1 dB), UAE regularly increased and decreased in a diurnal rhythm in parallel with DeltaW on cloudy days and at night. These low-dB emissions were interrupted by UAE abruptly switching between the low-dB range and a high-dB range (36 +/- 1 dB) on clear, sunny days, corresponding to the widely supported interpretation of UAE as sound from cavitations. It is hypothesized that the low-dB signals in drought-stressed trees are caused by respiration and/or cambial growth as these physiological activities are tissue water-content dependent and have been shown to produce courses of CO(2) efflux similar to our courses of Delta W and low-dB UAE.

Daily patterns of stem size variation in irrigated and unirrigated Eucalyptus globulus

High resolution measurements of stem diameter variation provide a means to study short-term dynamics of tree growth and water status. In this 14-month study, daily changes in stem radius of Eucalyptus globulus Labill. seedlings were measured with electronic point dendrometers in a plantation in southern Tasmania, Australia. The daily patterns of stem expansion and shrinkage were classified into three phases: shrinkage; recovery; and increase in diameter from one maximum to the next, or increment. This study showed that rapid onset of even mild drought in irrigated trees caused distinct changes in daily patterns of stem diameter variation, particularly the duration of daily stem increment. The duration of the daily increment phase was directly related to increment magnitude. The dynamics of daily increment were significantly affected by mean minimum temperature, indicating a temperature limitation on metabolic processes underlying diameter growth in these trees. Most likely due to differences in conductance, the duration but not rate of the incremental daily expansion was greater in fast- than in slow-growing trees.

Dendrometer and intra-annual tree growth: What kind of information can be inferred?

Dendrometer measurements provide time series composed of the rhythm of water storage fluctuations over the year and seasonal tree growth. For slow-growing trees, however, difficulties have been found in the identification of crucial events such as growth onset, stem growth period and cessation, rendering it necessary to define what can be measured and at which time scale. Time scale means the time interval (from one day to one month) at which stem radius variation is extracted. In this study, two conifer species were monitored by an automatic band dendrometer to assess several time scales and analysis approaches. Data were collected from 8 trees of Picea abies (L.) Karst and Larix decidua L., growing at 1020 and 2080 m a.s.l. in the eastern Italian Alps, from 2000 to 2003. Time series of stem radius variation were extracted with different approaches, such as the stem cycle, daily mean and daily maximum. Several approaches can be used, as very similar time series of stem radius variations were produced with high coefficients of correlation among the series. At lower altitude, the approximate onset was identified at the beginning of May with a 10-day time scale, when the distribution of stem radius variation differed from zero. The main growth period, from May to June–July, corresponded mainly with earlywood cell formation. At higher altitude, a time scale of at least 15 days facilitated identification of the main period of stem growth only, corresponding with earlywood cell formation. Even if latewood cells were produced in August at both altitudes, the variability in stem radius changes was higher than the amount of growth in terms of cell-wood production. For a slow-growing species in a cold environment, an understanding of the growth period, assessed with several time scales, is necessary when using time series of stem radius variation to assess growth and climate relationships. The period used for growth and climate analysis should correspond only with the main period of stem growth.

Modeling tree water deficit from microclimate: an approach to quantifying drought stress

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaWE), with soil water potential (Psisoil) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth. We followed changes in DeltaW and DeltaWE in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaWE. This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psisoil or VPD alone provided poorer estimates of DeltaW than a model incorporating both factors. As a first approximation of DeltaWE, Psisoil can be weighted so that the negative mean Psisoil reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: approximately 65%, P. abies: approximately 70%, P. sylvestris: approximately 75%). The differences in DeltaW among species can be partially explained by a different weighting of Psisoil against VPD. The DeltaW of P. sylvestris was more dependent on Psisoil than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.

Seasonal changes in stem radius and production of new tracheids in Norway spruce.

The progress of xylem formation in Norway spruce (Picea abies (L.) Karst.) was measured during one growing season in southern Finland. Stem radius was monitored continuously with band dendrometers, and the formation of new tracheids was determined by examination of small increment cores taken twice weekly. Tracheid production started in June and ceased in August. Xylem formation was fastest in early July, when 0.75-1.25 new tracheids were formed per day. The rate of xylem formation was significantly correlated with mean daily temperature. Synchronous fluctuations in tracheid and lumen diameters were observed at the same relative positions within each annual ring, but no relationship existed between the diameters and weather variables. The timing of changes in stem radius differed from the timing of actual xylem formation. Stem radius increased in April and May, and the fastest daily increments were recorded in June. Increases in stem radius slowed in July, but small increases were measured more than a month after xylem formation had ceased. Daily changes in stem radius were correlated with daily precipitation, reflecting changes in stem water content. Therefore, dendrometers are of dubious value for measuring the timing of actual xylem formation. Small increment cores proved to be useful in assessing actual xylem formation, but the method is laborious.