Patterns, timing, and environmental drivers of growth in two coexisting green-stemmed Mediterranean alpine shrubs species.

The Mediterranean alpine is one of the most vulnerable ecosystems under future environmental change. Yet, patterns, timing and environmental controls of plant growth are poorly investigated. We aimed at an improved understanding of growth processes, as well as stem swelling and shrinking patterns, by examining two common coexisting green-stemmed shrub species. Using dendrometers to measure daily stem diameter changes, we separated these changes into water-related shrinking and swelling and irreversible growth. Implementing correlation analysis, linear mixed effects models, and partial least squares regression on time series of stem diameter changes, with corresponding soil temperature and moisture data as environmental predictors, we found species-specific growth patterns related to different drought-adaptive strategies. We show that the winter-cold-adapted species Cytisus galianoi uses a drought tolerance strategy combined with a high ecological plasticity, and is, thus, able to gain competitive advantages under future climate warming. In contrast, Genista versicolor is restricted to a narrower ecological niche using a winter-cold escape and drought avoidance strategy, which might be of disadvantage in a changing climate. Pregrowth environmental conditions were more relevant than conditions during growth, controlling the species' resource availability. Thus, studies focusing on current driver constellations of growth may fail to predict a species' ecological niche and its potential future performance.

Shifts in intra-annual growth dynamics drive a decline in productivity of temperate trees in Central European forest under warmer climate.

Climate change shifts tree growth phenology and dynamics in temperate forests. However, there is still little information on how warming climate changes intra-annual growth patterns and how these changes affect the productivity and carbon uptake of temperate trees. To address this knowledge gap, we used high-precision growth data from automatic dendrometers to quantify the impacts of unusually warm weather in 2022 (hot year) on growth phenology, dynamics and aboveground biomass (AGB) production in eight common temperate species (both conifers and broadleaved) in the Czech Republic. Mixed-effect models were used to investigate inter-annual changes in the start, end, and length of the growing season and intra-annual growth dynamics. We also modelled how changes in growth phenology, growth rates, and tree size affected yearly AGB production of individual trees. In the hot year, the growth started 5 days earlier, peaked 22 days earlier and ended 20 days earlier than in the climatically normal year, resulting in a shorter growing season with fewer growing days. AGB production decreased 36 % in the hot year, mainly due to fewer growing days and lower maximum growth rates, but with significant variation among tested species. The decline in AGB production in the hot year was most significant in the most productive species, which were also the species with the greatest reduction in the number of growing days. Tree size strongly enhanced AGB production, but its effect did not change with climate variation. Our findings suggest that climate change is likely to advance but also shorten the growing season of temperate trees, resulting in lower biomass production and carbon uptake. The results also indicate that the fast-growing and highly productive temperate tree species will have their growth reduced most by climate change, which will increasingly limit their high carbon sequestration potential.

Radial stem growth dynamics and phenology of a multi-stemmed species (Corylus avellana L.) across orchards in the Northern and Southern hemispheres.

Climate change and the global economy impose new challenges in the management of food-producing trees and require studying how to model plant physiological responses, namely growth dynamics and phenology. Hazelnut (Corylus avellana L.) is a multi-stemmed forest species domesticated for nut production and now widely spread across different continents. However, information on stem growth and its synchronization with leaf and reproductive phenology is extremely limited. This study aimed at (i) defining the sequencing of radial growth phases in hazelnut (onset, maximum growth and cessation) and the specific temperature triggering stem growth; and (ii) combining the stem growth phases with leaf and fruit phenology. Point dendrometers were installed on 20 hazelnut trees across eight orchards distributed in the Northern and Southern hemisphere during a period of three growing seasons between 2015 and 2018. The radial growth variations and climatic parameters were averaged and recorded every 15 min. Leaf and reproductive phenology were collected weekly at each site. Results showed that stem radial growth started from day of year 84 to 134 in relation to site and year but within a relatively narrow range of temperature (from 13 to 16.5 °C). However, we observed a temperature-related acclimation in the cultivar Tonda di Giffoni. Maximum growth always occurred well before the summer solstice (on average 35 days) and before the maximum annual air temperatures. Xylogenesis developed rapidly since the time interval between onset and maximum growth rate was about 3 weeks. Importantly, the species showed an evident delay of stem growth onset with respect to leaf emergence (on average 4-6 weeks) rarely observed in tree species. These findings represent the first global analysis of radial growth dynamics in hazelnut, which is an essential step for developing models on orchard functioning and management on different continents.

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 15 min over 7 years) 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.

Does phloem osmolality affect diurnal diameter changes of twigs but not of stems in Scots pine?

Diel stem diameter changes measured at the stem base of temperate tree species can be mostly explained by a hydraulic system of flow and storage compartments passively driven by transpiration. Active, osmotic processes are considered to play a minor role only. Here we explore whether such osmotic processes have a stronger impact on diel changes in twig diameter than in stem diameter because twigs are closer to the leaves, the main source of newly acquired carbon. We investigated stem and twig diameter changes of wood and bark of pine trees in parallel to fluctuations of the osmolality in needles and in the bark at the stem base. We found consistent twig bark size increments concurrent with twig wood size decreases during daylight hours whereas needle osmolality was not consistently increasing even on sunny days. The size changes of bark and wood either reversed or ran in parallel from late afternoon onwards until the next morning. No such patterns were measurable at the stem base. Stem wood was hardly changing in size, whereas stem bark followed the regular pattern of a decrease during the daylight hours and an increase during the night. Osmolality at the stem base showed no particular course over 24 h. We conclude that assimilates from the needles were rapidly transported to the twigs where they increased the osmolality of the bark tissue by sugar loading, explaining the bark size increase (over-) compensating the xylem size decrease. The stem base largely followed the expectation of a passive, hydraulic system without a measurable role of osmoregulation. Diameter changes thus follow different diurnal dynamics in twigs and at the stem base.

Rhizophoraceae Mangrove Saplings Use Hypocotyl and Leaf Water Storage Capacity to Cope with Soil Water Salinity Changes.

Some of the most striking features of Rhizophoraceae mangrove saplings are their voluminous cylinder-shaped hypocotyls and thickened leaves. The hypocotyls are known to serve as floats during seed dispersal (hydrochory) and store nutrients that allow the seedling to root and settle. In this study we investigate to what degree the hypocotyls and leaves can serve as water reservoirs once seedlings have settled, helping the plant to buffer the rapid water potential changes that are typical for the mangrove environment. We exposed saplings of two Rhizophoraceae species to three levels of salinity (15, 30, and 0-5‰, in that sequence) while non-invasively monitoring changes in hypocotyl and leaf water content by means of mobile NMR sensors. As a proxy for water content, changes in hypocotyl diameter and leaf thickness were monitored by means of dendrometers. Hypocotyl diameter variations were also monitored in the field on a Rhizophora species. The saplings were able to buffer rapid rhizosphere salinity changes using water stored in hypocotyls and leaves, but the largest water storage capacity was found in the leaves. We conclude that in Rhizophora and Bruguiera the hypocotyl offers the bulk of water buffering capacity during the dispersal phase and directly after settlement when only few leaves are present. As saplings develop more leaves, the significance of the leaves as a water storage organ becomes larger than that of the hypocotyl.

High light decreases xylem contribution to fruit growth in tomato.

Recently, contradicting evidence has been reported on the contribution of xylem and phloem influx into tomato fruits, urging the need for a better understanding of the mechanisms involved in fruit growth. So far, little research has been performed on quantifying the effect of light intensity on the different contributors to the fruit water balance. However, as light intensity affects both transpiration and photosynthesis, it might be expected to induce important changes in the fruit water balance. In this study, tomato plants (Solanum lycopersicum L.) were grown in light and shade conditions and the fruit water balance was studied by measuring fruit growth of girdled and intact fruits with linear variable displacement transducers combined with a model-based approach. Results indicated that the relative xylem contribution significantly increased when shading lowered light intensity. This resulted from both a higher xylem influx and a lower phloem influx during the daytime. Plants from the shade treatment were able to maintain a stronger gradient in total water potential between stem and fruits during daytime, thereby promoting xylem influx. It appeared that the xylem pathway was still functional at 35 days after anthesis and that relative xylem contribution was strongly affected by environmental conditions.

Variable hydraulic resistances and their impact on plant drought response modelling.

Plant drought responses are still not fully understood. Improved knowledge on drought responses is, however, crucial to better predict their impact on individual plant and ecosystem functioning. Mechanistic models in combination with plant measurements are promising for obtaining information on plant water status and can assist us in understanding the effect of limiting soil water availability and drought stress. While existing models are reliable under sufficient soil water availability, they generally fail under dry conditions as not all appropriate mechanisms seem yet to have been implemented. We therefore aimed at identifying mechanisms underlying plant drought responses, and in particular investigated the behaviour of hydraulic resistances encountered in the soil and xylem for grapevine (Vitis vinifera L.) and oak (Quercus robur L.). A variable hydraulic soil-to-stem resistance was necessary to describe plant drought responses. In addition, implementation of a variable soil-to-stem hydraulic resistance enabled us to generate an in situ soil-to-stem vulnerability curve, which might be an alternative to the conventionally used vulnerability curves. Furthermore, a daily recalibration of the model revealed a drought-induced increase in radial hydraulic resistance between xylem and elastic living tissues. Accurate information on plant hydraulic resistances and simulation of plant drought responses can foster important discussions regarding the functioning of plants and ecosystems during droughts.

How to catch the patch? A dendrometer study of the radial increment through successive cambia in the mangrove Avicennia.

BACKGROUND AND AIMS: Successive vascular cambia are involved in the secondary growth of at least 200 woody species from >30 plant families. In the mangrove Avicennia these successive cambia are organized in patches, creating stems with non-concentric xylem tissue surrounded by internal phloem tissue. Little is known about radial growth and tree stem dynamics in trees with this type of anatomy. This study aims to (1) clarify the process of secondary growth of Avicennia trees by studying its patchiness; and (2) study the radial increment of Avicennia stems, both temporary and permanent, in relation to local climatic and environmental conditions. A test is made of the hypothesis that patchy radial growth and stem dynamics enable Avicennia trees to better survive conditions of extreme physiological drought. Methods Stem variations were monitored by automatic point dendrometers at four different positions around and along the stem of two Avicennia marina trees in the mangrove forest of Gazi Bay (Kenya) during 1 year. KEY RESULTS: Patchiness was found in the radial growth and shrinkage and swelling patterns of Avicennia stems. It was, however, potentially rather than systematically present, i.e. stems reacted either concentrically or patchily to environment triggers, and it was fresh water availability and not tidal inundation that affected radial increment. CONCLUSIONS: It is concluded that the ability to develop successive cambia in a patchy way enables Avicennia trees to adapt to changes in the prevailing environmental conditions, enhancing its survival in the highly dynamic mangrove environment. Limited water could be used in a more directive way, investing all the attainable resources in only some locations of the tree stem so that at least at these locations there is enough water to, for example, overcome vessel embolisms or create new cells. As these locations change with time, the overall functioning of the tree can be maintained.

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.

Using automated point dendrometers to analyze tropical treeline stem growth at Nevado de Colima, Mexico.

The relationship between wood growth and environmental variability at the tropical treeline of North America was investigated using automated, solar-powered sensors (a meteorological station and two dendrometer clusters) installed on Nevado de Colima, Mexico (19° 35' N, 103° 37' W, 3,760 m a.s.l.). Pure stands of Pinus hartwegii Lindl. (Mexican mountain pine) were targeted because of their suitability for tree-ring analysis in low-latitude, high-elevation, North American Monsoon environments. Stem size and hydroclimatic variables recorded at half-hour intervals were summarized on a daily timescale. Power outages, insect outbreaks, and sensor failures limited the analysis to non-consecutive months during 2001-2003 at one dendrometer site, and during 2002-2005 at the other. Combined data from the two sites showed that maximum radial growth rates occur in late spring (May), as soil temperature increases, and incoming short-wave radiation reaches its highest values. Early season (April-May) radial increment correlated directly with temperature, especially of the soil, and with solar radiation. Stem expansion at the start of the summer monsoon (June-July) was mostly influenced by moisture, and revealed a drought signal, while late season relationships were more varied.