Separating continuously measured stem radius (SR) fluctuations into growth-induced irreversible stem expansion (GRO) and tree water deficit-induced reversible stem shrinkage (TWD) requires a ...conceptualization of potential growth processes that may occur during periods of shrinking and expanding SR below a precedent maximum. Here, we investigated two physiological concepts: the linear growth (LG) concept, assuming linear growth, versus the zero growth (ZG) concept, assuming no growth during periods of stem shrinkage.
We evaluated the physiological mechanisms underlying these two concepts and assessed their respective plausibilities using SR data obtained from 15 deciduous and evergreen trees.
The application of the LG concept produced steady growth rates, whereas growth rates varied strongly under the ZG concept, more in accordance with mechanistic expectations. Further, growth increased for a maximum of 120 min after periods of stem shrinkage, indicating limited growth activity during those periods. However, this extra growth was found to be a small fraction of total growth only. Furthermore, TWD under the ZG concept was better explained by a hydraulic plant model than TWD under the LG concept.
We conclude that periods of stem shrinkage allow for very little growth in the four tree species investigated. However, further studies should focus on obtaining independent growth data to ultimately validate these findings.
Abstract
Direct evidence for the link between stem diameter variations (SDV) and the daily canopy water status, i.e., daily water potentials (Ψ), is rare, particularly for tall trees. It thus remains ...unclear up to what degree SDV readings are useful to estimate daily canopy Ψ. We measured SDV with point dendrometers at the stem base of tall, mature individuals of six European forest tree species in a near-natural temperate forest and compared them with daily canopy Ψ during the growing seasons of 2014 (wet) and 2015 (dry). Stem diameter variations were de-trended for growth with two different approaches leading to the so-called tree water deficit (TWD). We found that midday Ψ can be predicted from TWD, independent of the growth-de-trending procedure to obtain TWD from SDV. Further, daily TWD was a better indicator for daily midday Ψ, particularly under dry conditions, than maximum daily shrinkage, another common quantity derived from SDV. Based on data from six temperate tree species, we conclude that TWD measured at the stem base is a consistent proxy for daily canopy midday Ψ of tall trees over the entire range of measured conditions.
Why trees grow at night Zweifel, Roman; Sterck, Frank; Braun, Sabine ...
New phytologist,
September 2021, Letnik:
231, Številka:
6
Journal Article
Recenzirano
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Summary
The timing of diel stem growth of mature forest trees is still largely unknown, as empirical data with high temporal resolution have not been available so far. Consequently, the effects of ...day–night conditions on tree growth remained uncertain.
Here we present the first comprehensive field study of hourly‐resolved radial stem growth of seven temperate tree species, based on 57 million underlying data points over a period of up to 8 yr.
We show that trees grow mainly at night, with a peak after midnight, when the vapour pressure deficit (VPD) is among the lowest. A high VPD strictly limits radial stem growth and allows little growth during daylight hours, except in the early morning. Surprisingly, trees also grow in moderately dry soil when the VPD is low. Species‐specific differences in diel growth dynamics show that species able to grow earlier during the night are associated with the highest number of hours with growth per year and the largest annual growth increment.
We conclude that species with the ability to overcome daily water deficits faster have greater growth potential. Furthermore, we conclude that growth is more sensitive than carbon uptake to dry air, as growth stops before stomata are known to close.
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•Trees at mid-latitudes of their distribution show greater sensitivity to drought.•Water deficit mainly controls tree resistance and spatial sensitivity to drought.•At wetter sites, ...higher drought frequency trains tree memory response to drought.•At drier sites, higher drought frequency causes substantial damage to junipers.
The resistance of forests to extreme climatic events such as drought shapes their sensitivity to future extreme events in space and time. To a large extent, the ability of trees to learn from prior droughts explains how trees adjust their sensitivity to water deficit. We use tree-ring width data collected from 1565 juniper trees (Juniperus prezwalskii and Juniperus tibetica) across 57 sites on the Tibetan Plateau to model tree resistance to water deficit and to map drought sensitivity across the species’ distribution. We test the effect of both the frequency and severity of drought on the drought memory of trees. We find that trees at mid-latitudes and in the northwestern part of the juniper distribution range exhibit higher drought sensitivity. Water deficit is the main factor controlling tree resistance and thus affects spatial sensitivity to drought. At wetter sites, higher drought frequency enhances tree adaptability through ecological memory, thereby promoting tree resistance and decreasing tree sensitivity to extreme drought events. At drier sites, higher drought frequency causes junipers’ growth decline that is not beneficial for tolerance to extreme drought events. Regional drought conditions and the frequency of pre-droughts affect tree resistance and sensitivity to extreme drought events. This explains the spatial pattern of drought risk for juniper forests on the Tibetan Plateau, and helps us to better understand the vulnerability of this high-elevation forest ecosystem. Such information is important for maintaining forest health and informing the sustainable development of the Tibetan Plateau under a changing climate.
An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner‐Alpine regions, the species ...composition in low elevation forests is changing: The sub‐boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub‐Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger‐scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed‐effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small‐diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.
Radial stem size changes, measured with automated dendrometers at intra-daily resolution, offer great potential to link environmental conditions with tree physiology at the seasonal scale. Such ...measurements need to be time-aligned, cleaned of outliers and shifts, gap-filled and analysed for reversible (water-related) and irreversible (growth-related) fractions to obtain physiologically meaningful data. Therefore, comprehensive tools are needed for reproducible data processing and analytics of dendrometer data. Here we present a transparent method, compiled in the R package treenetproc, to turn raw dendrometer data into clean, physiologically interpretable information, i.e., stem growth, tree water deficit, growth phenological phases, mean daily shrinkage and their respective timings. The removal of errors is facilitated by additional functions and supported with graphical visualizations. To ensure reproducible data handling, the processing parameters and induced changes to the raw data are documented in the output and, thus, are a step towards a standardized processing of automatically measured stem radius time series. We discuss examples, such as the seasonality of growth or the dependence of growth on atmospheric and soil drought. The presented growth and water-related physiological variables at high temporal resolution offer novel physiological insights into the seasonally varying responses of trees to changing environmental conditions.
Wood is a sustainable natural resource and an important global commodity. According to the 'moon wood theory', the properties of wood, including its growth and water content, are believed to ...oscillate with the lunar cycle. Despite contradicting our current understanding of plant functioning, this theory is commonly exploited for marketing wooden products. To examine the moon wood theory, we applied a wavelet power transformation to series of 2,000,000 hourly stem radius records from dendrometers. We separated the influence of 74 consecutive lunar cycles and meteorological conditions on the stem variation of 62 trees and six species. We show that the dynamics of stem radius consist of overlapping oscillations with periods of 1 day, 6 months, and 1 year. These oscillations in stem dimensions were tightly coupled to oscillations in the series of air temperature and vapour pressure deficit. By contrast, we revealed no imprint of the lunar cycle on the stem radius variation of any species. We call for scepticism towards the moon wood theory, at least as far as the stem water content and radial growth are concerned. We foresee that similar studies employing robust scientific approaches will be increasingly needed in the future to cope with misleading concepts.
•How long-term soil moisture change affects the sensitivity of transpiration to environmental variability is unknown.•Exposure to long-term soil moisture reduction decreased sap flux sensitivity to ...VPD and REW.•Long-term irrigation increased sap flux sensitivity to VPD and REW but at highly water-limited sites only.•Relative sensitivity is generalizable across forest types suggesting common adjustment mechanisms to soil moisture status.
Tree transpiration depends on biotic and abiotic factors that might change in the future, including precipitation and soil moisture status. Although short-term sap flux responses to soil moisture and evaporative demand have been the subject of attention before, the relative sensitivity of sap flux to these two factors under long-term changes in soil moisture conditions has rarely been determined experimentally. We tested how long-term artificial change in soil moisture affects the sensitivity of tree-level sap flux to daily atmospheric vapor pressure deficit (VPD) and soil moisture variations, and the generality of these effects across forest types and environments using four manipulative sites in mature forests. Exposure to relatively long-term (two to six years) soil moisture reduction decreases tree sap flux sensitivity to daily VPD and relative extractable water (REW) variations, leading to lower sap flux even under high soil moisture and optimal VPD. Inversely, trees subjected to long-term irrigation showed a significant increase in their sensitivity to daily VPD and REW, but only at the most water-limited site. The ratio between the relative change in soil moisture manipulation and the relative change in sap flux sensitivity to VPD and REW variations was similar across sites suggesting common adjustment mechanisms to long-term soil moisture status across environments for evergreen tree species. Overall, our results show that long-term changes in soil water availability, and subsequent adjustments to these novel conditions, could play a critical and increasingly important role in controlling forest water use in the future.
Abstract
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.