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% towards northern Europe and 38% towards 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 towards 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 towards 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.

This article is a broad review focused on dragon trees—one of the most famous groups of trees in the world, well known from ancient times. These tertiary relicts are severely endangered in most of the area where they grow. The characteristic features of the dragon tree group are described and the species belonging to this group are listed. This review gathers together current knowledge regarding the taxonomy, evolution, anatomy and morphology, physiology, and ontogeny of arborescent dragon tree species. Attention is also paid to the composition, harvesting, medicinal, and ethnobotanical use of the resin (dragons’ blood). An evaluation of population structure, distribution, ecology, threats, and nature conservation forms the final part of the review. In the conclusions we recommend further avenues of research that will be needed to effectively protect all dragon tree species.

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.

Spring drought is becoming a frequently occurring stress factor in temperate forests. However, the understanding of tree resistance and resilience to the spring drought remains insufficient. In this study, European beech (Fagus sylvatica L.) seedlings at the early stage of leaf development were moderately and severely drought stressed for 1 month and then subjected to a 2-week recovery period after rewatering. The study aimed to disentangle the complex relationships between leaf gas exchange, vascular anatomy, tree morphology and patterns of biomass allocation. Stomatal conductance decreased by 80 and 85% upon moderate and severe drought stress, respectively, which brought about a decline in net photosynthesis. However, drought did not affect the indices of slow chlorophyll fluorescence, indicating no permanent damage to the light part of the photosynthetic apparatus. Stem hydraulic conductivity decreased by more than 92% at both drought levels. Consequently, the cambial activity of stressed seedlings declined, which led to lower stem biomass, reduced tree ring width and a lower number of vessels in the current tree ring, these latter also with smaller dimensions. In contrast, the petiole structure was not affected, but at the cost of reduced leaf biomass. Root biomass was reduced only by severe drought. After rewatering, the recovery of gas exchange and regrowth of the current tree ring were observed, all delayed by several days and by lower magnitudes in severely stressed seedlings. The reduced stem hydraulic conductivity inhibited the recovery of gas exchange, but xylem function started to recover by regrowth and refilling of embolized vessels. Despite the damage to conductive xylem, no mortality occurred. These results suggest the low resistance but high resilience of European beech to spring drought. Nevertheless, beech resilience could be weakened if the period between drought events is short, as the recovery of severely stressed seedlings took longer than 14 days.

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.