Roman Gebauer

Effect of thinning on anatomical adaptations of Norway spruce needles

Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.

Effects of different light conditions on the xylem structure of Norway spruce needles

Conifer needles are extraordinarily variable and much of this diversity is linked to the water transport capacity of the xylem and to xylem conduit properties. However, we still know little about how anatomical characteristics influence the hydraulic efficiency of needle xylem in different parts of the crown. In this study we evaluated needle function and anatomy in Norway spruce families exposed to different light conditions. We measured tracheid and needle characteristics of sun-exposed and shaded current-year needles in two experimental plots: a control plot and a thinned plot with 50% reduction in stand density. Sun-exposed needles had a larger tracheid lumen area than shaded needles, and this was caused by a larger maximum tracheid lumen diameter, while the minimum lumen diameter was less plastic. Sun-exposed needles had also higher theoretical hydraulic conductivity than shaded needles. Thinning leads to increased radiation to the lower branches, and presumably exposes the upper branches to stronger water stress than before thinning. Thinning affected several needle parameters both in sun-exposed and shaded needles; tracheid lumens were more circular and minimum tracheid lumen diameter was larger in the thinned plot, whereas maximum tracheid lumen diameter was less plastic on both plots. This study demonstrates that needle xylem structure in Norway spruce is clearly influenced by the light gradient within the tree crown.

Osmolality and Non-Structural Carbohydrate Composition in the Secondary Phloem of Trees across a Latitudinal Gradient in Europe.

Phloem osmolality and its components are involved in basic cell metabolism, cell growth, and in various physiological processes including the ability of living cells to withstand drought and frost. Osmolality and sugar composition responses to environmental stresses have been extensively studied for leaves, but less for the secondary phloem of plant stems and branches. Leaf osmotic concentration and the share of pinitol and raffinose among soluble sugars increase with increasing drought or cold stress, and osmotic concentration is adjusted with osmoregulation. We hypothesize that similar responses occur in the secondary phloem of branches. We collected living bark samples from branches of adult Pinus sylvestris, Picea abies, Betula pendula and Populus tremula trees across Europe, from boreal Northern Finland to Mediterranean Portugal. In all studied species, the observed variation in phloem osmolality was mainly driven by variation in phloem water content, while tissue solute content was rather constant across regions. Osmoregulation, in which osmolality is controlled by variable tissue solute content, was stronger for Betula and Populus in comparison to the evergreen conifers. Osmolality was lowest in mid-latitude region, and from there increased by 37% toward northern Europe and 38% toward southern Europe due to low phloem water content in these regions. The ratio of raffinose to all soluble sugars was negligible at mid-latitudes and increased toward north and south, reflecting its role in cold and drought tolerance. For pinitol, another sugar known for contributing to stress tolerance, no such latitudinal pattern was observed. The proportion of sucrose was remarkably low and that of hexoses (i.e., glucose and fructose) high at mid-latitudes. The ratio of starch to all non-structural carbohydrates increased toward the northern latitudes in agreement with the build-up of osmotically inactive C reservoir that can be converted into soluble sugars during winter acclimation in these cold regions. Present results for the secondary phloem of trees suggest that adjustment with tissue water content plays an important role in osmolality dynamics. Furthermore, trees acclimated to dry and cold climate showed high phloem osmolality and raffinose proportion.

Transpiration and stomatal conductance of mistletoe (Loranthus europaeus) and its host plant, downy oak (Quercus pubescens)

Sap flow rate was measured in the crown of a solitary specimen of downy oak (Quercus pubescens) infested by mistletoe (Loranthus europaeus). Five oak branches and two mistletoe plants were selected for analysis. The seasonal sum of transpired water expressed per leaf area unit was five times higher in the mistletoe than in the oak. In addition, the diurnal curves of sap flow were different between the plants. In the morning, the sap flow measured in the mistletoe lagged one hour behind the sap flow measured in an oak branch unencumbered by mistletoe. In contrast, no time lag was observed in the evening. The proportion of water transpired at night relative to the total transpiration was 7% in both species. The stomatal conductances derived from the inverted Penman-Monteith equation and their dependence on global radiation and the vapour pressure deficit (D) revealed that D exerts a different behaviour in stomatal control of transpiration in the mistletoe. We also determined that the concentration of calcium in the leaf mass could serve as a proxy for transpiration rate, however the relationship was not proportional.

Root Function: In Situ Studies Through Sap Flow Research

Sap flow measured by the Heat Field Deformation technique, HFD, is sensitive to flow responses to small changes in water potential gradients within the tree hydraulic systems. When these changes occur abruptly, under experimental treatments (severing, localized irrigation, heavy loading), sap flow movement can be used as a marker to study root functionality, for example root ability to redistribute water and withstand heavy machinery pressure. Experiments also showed that a compensation mechanism may operate in trees, with a temporary increase in the absorbed water due to a preferential use of one part of the root system when another part is damaged or when a water source is lost. Long-term measurements of root sap flow allow distinguishing between water uptake from shallow and deep rooted trees, at different exposures at a forest edge and from healthy and infected trees. Root sap flow can be used as an indicator of tree stress or of the prevailing mechanisms used by trees to survive drought, under irrigation or rain-fed conditions.

A comparative structural and functional study of leaf traits and sap flow in Dracaena cinnabari and Dracaena draco seedlings

Water relations for two remote populations of Dracaena tree species from the dragon tree group, Dracaena cinnabari Balfour f. and Dracaena draco (L.) L., were studied to test our hypothesis that morphological and anatomical differences in leaf structure may lead to varied functional responses to changing environmental conditions. Sap flow measurements were performed using the heat field deformation method for four Dracaena seedlings grown in one glasshouse and two greenhouses, and leaf traits related to plant-water relationships were characterised. All traits studied confirmed that D. cinnabari leaves are more xeric in their morpho-anatomical structure compared with D. draco leaves. No radial sap flow variability was detected in D. draco plant stems, whereas sap flow was found to be higher in the inner part of D. cinnabari stems. The regular occurrence of reverse sap flow at night in both Dracaena species was consistent with a staining experiment. Vapour pressure deficit (VPD) was found to be the main driver for transpiration for both Dracaena species. However, the relationship between VPD and sap flow appeared to be different for each species, with a clockwise or no hysteresis loop for D. draco and a counter-clockwise hysteresis loop for D. cinnabari. This resulted in a shorter transpiration cycle in D. cinnabari. The observed superior water-saving strategy of D. cinnabari corresponds to its more xeric morpho-anatomical leaf structure compared with D. draco.

Effects of prolonged drought on the anatomy of sun and shade needles in young Norway spruce trees

Abstract Predicted increases in the frequency and duration of drought are expected to negatively affect tree vitality, but we know little about how water shortage will influence needle anatomy and thereby the trees’ photosynthetic and hydraulic capacity. In this study, we evaluated anatomical changes in sun and shade needles of 20‐year‐old Norway spruce trees exposed to artificial drought stress. Canopy position was found to be important for needle structure, as sun needles had significantly higher values than shade needles for all anatomical traits (i.e., cross‐sectional needle area, number of tracheids in needle, needle hydraulic conductivity, and tracheid lumen area), except proportion of xylem area per cross‐sectional needle area. In sun needles, drought reduced all trait values by 10–40%, whereas in shade needles, only tracheid maximum diameter was reduced by drought. Due to the relatively weaker response of shade needles than sun needles in drought‐stressed trees, the difference between the two needle types was reduced by 25% in the drought‐stressed trees compared to the control trees. The observed changes in needle anatomy provide new understanding of how Norway spruce adapts to drought stress and may improve predictions of how forests will respond to global climate change.

Root Structure: In Situ Studies Through Sap Flow Research

Sap flow research highlights new perspectives to study in situ root structure of large trees. Several examples demonstrate the ability of the Heat Field Deformation method, HFD, to do this under natural and experimental conditions. Within the latter, localized irrigation, sink- or source-severing trigger sap flow responses that help us to understand the belowground parts of a tree, such as the presence of anastomoses between roots of different trees. The vertical profile of root density, as well as root size around a tree, can be derived from the stem sap flow radial profile. Increase of stem flow due to localized irrigation may be used to distinguish root locations near the corresponding stem sector. Responses of root or stem sap flow when exposing roots using an air-spade or following the severing of roots or branches help us to understand the relationships between different sapwood conducting layers and paths of water between sources and sinks.

Quantifying in situ phenotypic variability in the hydraulic properties of four tree species across their distribution range in Europe

Many studies have reported that hydraulic properties vary considerably between tree species, but little is known about their intraspecific variation and, therefore, their capacity to adapt to a warmer and drier climate. Here, we quantify phenotypic divergence and clinal variation for embolism resistance, hydraulic conductivity and branch growth, in four tree species, two angiosperms (Betula pendula, Populus tremula) and two conifers (Picea abies, Pinus sylvestris), across their latitudinal distribution in Europe. Growth and hydraulic efficiency varied widely within species and between populations. The variability of embolism resistance was in general weaker than that of growth and hydraulic efficiency, and very low for all species but Populus tremula. In addition, no and weak support for a safety vs. efficiency trade-off was observed for the angiosperm and conifer species, respectively. The limited variability of embolism resistance observed here for all species except Populus tremula, suggests that forest populations will unlikely be able to adapt hydraulically to drier conditions through the evolution of embolism resistance.

Norway Spruce Fine Roots and Fungal Hyphae Grow Deeper in Forest Soils After Extended Drought

Global warming will most likely lead to increased drought stress in forest trees. We wanted to describe the adaptive responses of fine roots and fungal hyphae, at different soil depths, in a Norway spruce stand to long-term drought stress induced by precipitation exclusion over two growing seasons. We used soil cores, minirhizotrons and nylon meshes to estimate growth, biomass and distribution of fine roots and fungal hyphae at different soil depths. In control plots fine roots proliferated in upper soil layers, whereas in drought plots there was no fine root growth in upper soil layers and roots mostly occupied deeper soil layers. Fungal hyphae followed the same pattern as fine roots, with the highest biomass in deeper soil layers in drought plots. We conclude that both fine roots and fungal hyphae respond to long-term drought stress by growing into deeper soil layers.