Global relationships in tree functional traits.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide.

The Protective Role of Bark and Bark Fibers of the Giant Sequoia (Sequoiadendron giganteum) during High-Energy Impacts.

The influences of (1) a high fiber content, (2) the arrangement of fibers in fiber groups, and (3) a layered hierarchical composition of the bark of the giant sequoia (Sequoiadendron giganteum) on its energy dissipation capability are analyzed and discussed regarding the relevance for an application in bioinspired components in civil engineering. The giant sequoia is native to the Sierra Nevada (USA), a region with regular rockfalls. It is thus regularly exposed to high-energy impacts, with its bark playing a major protective role, as can be seen in the wild and has been proven in laboratory experiments. The authors quantify the fundamental biomechanical properties of the bark at various length scales, taking into account its hierarchical setup ranging from the integral level (whole bark) down to single bark fibers. Microtensile tests on single fibers and fiber pairs give insights into the properties of single fibers as well as the benefits of the strong longitudinal interconnection between single fibers arranged in pairs. Going beyond the level of single fibers or fiber pairs, towards the integral level, quasistatic compression tests and dynamic impact tests are performed on samples comprising the whole bark (inner and outer bark). These tests elucidate the deformation behavior under quasistatic compression and dynamic impact relevant for the high energy dissipation and impact-damping behavior of the bark. The remarkable energy dissipation capability of the bark at the abovementioned hierarchical levels are linked to the layered and fibrous structure of the bark structurally analyzed by thin sections and SEM and µCT scans.

The role of bud protection and bark density in frost resistance of savanna trees.

Frost events occur with a significant frequency in savannas of the Southern Hemisphere, especially in the Cerrados of Brazil. One of the main strategies to deal with such events is to invest in thick and dense bark, which can insulate internal branch tissues and protect buds, essential to ensure resprouting if frost damage causes plant canopy die-back. Such strategies may be fundamental to determine the persistence of savanna species in regions where low temperatures and frost events are recurrent. Here we describe bud protection and bark strategies of 53 woody species growing in typical savanna vegetation of central Brazil. In addition, we used an experimental approach exposing branches to 0 °C to measure temperature variation in internal branch tissue and test its relationship to bud protection and bark properties. We found that the majority of species (69%) showed medium to high bud protection against extreme temperatures; however, the degree of bud protection was not clearly related to bark properties, such as bark thickness and density. Bark density is a fundamental trait in determining protection against low temperatures (0 °C), since species with low bark density showed lower temperature variation in their internal branch tissues, independently of the bud protection degree. Bark properties and bud protection are two different (albeit related) strategies for the protection and persistence of savanna trees under extreme environmental temperatures and can explain ecological observations related to savanna tree responses after frost events.

Tree defence and bark beetles in a drying world: carbon partitioning, functioning and modelling.

Drought has promoted large-scale, insect-induced tree mortality in recent years, with severe consequences for ecosystem function, atmospheric processes, sustainable resources and global biogeochemical cycles. However, the physiological linkages among drought, tree defences, and insect outbreaks are still uncertain, hindering our ability to accurately predict tree mortality under on-going climate change. Here we propose an interdisciplinary research agenda for addressing these crucial knowledge gaps. Our framework includes field manipulations, laboratory experiments, and modelling of insect and vegetation dynamics, and focuses on how drought affects interactions between conifer trees and bark beetles. We build upon existing theory and examine several key assumptions: (1) there is a trade-off in tree carbon investment between primary and secondary metabolites (e.g. growth vs defence); (2) secondary metabolites are one of the main component of tree defence against bark beetles and associated microbes; and (3) implementing conifer-bark beetle interactions in current models improves predictions of forest disturbance in a changing climate. Our framework provides guidance for addressing a major shortcoming in current implementations of large-scale vegetation models, the under-representation of insect-induced tree mortality.

Measurement of Inner Bark and Leaf Osmolality.

Sugar transport in the phloem is driven by turgor pressure gradients which are created by osmotic gradients resulting from sugars loaded to the phloem at the source tissue and unloaded at the sink tissue. Therefore, osmolality is a key parameter that can be used to evaluate sugar status and get an indication of the driving force for phloem transport. Here we describe how osmotic concentration measurements from inner bark (practically, the phloem) and needles of trees can be measured. This protocol presents the procedure used by Lintunen et al. (Front Plant Sci 7:726, 2016) and Paljakka et al. (Plant Cell Environ 40:2160-2173, 2017), extended by practical advice and discussion of potential errors and caveats. We describe how to implement this procedure for gymnosperm as well as angiosperm trees. This method uses mechanical sap extraction with a centrifuge from inner bark and leaf samples, which have gone through a deep freeze treatment and thawing. The osmotic potential of these samples is then analyzed with a freezing point or vapor pressure osmometer. The aim of these measurements is to study the spatial and temporal dynamics of phloem function.

Bark in Woody Plants: Understanding the Diversity of a Multifunctional Structure.

Most biological structures carry out multiple functions. Focusing on only one function to make adaptive inferences overlooks that manifold selection pressures and tradeoffs shape the characteristics of a multifunctional structure. Focusing on single functions can only lead to a partial picture of the causes underlying diversity and the evolutionary origin of the structure in question. I illustrate this discussion using bark as a study case. Bark comprises all the tissues surrounding the xylem in woody plants. Broadly, bark includes an inner and mostly living region and an outer, dead one. Of all plant structures, bark has the most complex anatomical structure and ontogenetic origin involving two (and often three) different meristems. Traditionally, the wide diversity in bark traits, mainly bark thickness, has been interpreted as the result of the selective pressures imposed by fire regime. However, recent research has shown that explanations based on fire regime cannot account for salient patterns of bark variation globally including the very strong inner bark thickness-stem diameter scaling, which is likely due to metabolic needs, and the very high intracommunity variation in total, inner, and outer bark thickness, and in inner:outer proportions. Moreover, explanations based on fire disregard that in addition to fire protection, bark carries out several other crucial functions for plants including translocation of photosynthates; storage of starch, soluble sugars, water, and other compounds; protection from herbivores, pathogens, and high temperatures; wound closure, as well as mechanical support, photosynthesis, and likely being involved in xylem embolism repair. All these functions are crucial for plant performance and are involved in synergistic (e.g., storage of water and insulation) and trade-off relationships (e.g., protection from fire vs photosynthetic activity). Focusing on only one of these functions, protection from fire has provided an incomplete picture of the selective forces shaping bark diversity and has severely hindered our incipient understanding of the functional ecology of this crucial region of woody stems. Applying a multifunctional perspective to the study of bark will allow us to address why we observe such high intracommunity variation in bark traits, why some bark trait combinations are ontogenetically impossible or penalized by selection, how bark is coordinated functionally with other plant parts, and as a result, to understand how bark contributes to the vast diversity of plant ecological strategies across the globe.

The formation and accumulation of protein-networks by physical interactions in the rapid occlusion of laticifer cells in rubber tree undergoing successive mechanical wounding.

BACKGROUND: Although the wound response of plants has been extensively studied, little is known of the rapid occlusion of wounded cell itself. The laticifer in rubber tree is a specific type of tissue for natural rubber biosynthesis and storage. In natural rubber production, tapping is used to harvest the latex which flows out from the severed laticifer in the bark. Therefore, study of the rapid wound-occlusion of severed laticifer cells is important for understanding the rubber tree being protected from the continuously mechanical wounding. RESULTS: Using cytological and biochemical techniques, we revealed a biochemical mechanism for the rapid occlusion of severed laticifer cells. A protein-network appeared rapidly after tapping and accumulated gradually along with the latex loss at the severed site of laticifer cells. Triple immunofluorescence histochemical localization showed that the primary components of the protein-network were chitinase, ß-1,3-glucanase and hevein together with pro-hevein (ProH) and its carboxyl-terminal part. Molecular sieve chromatography showed that the physical interactions among these proteins occurred under the condition of neutral pH. The interaction of ß-1,3-glucanase respectively with hevein, chitinase and ProH was testified by surface plasmon resonance (SPR). The interaction between actin and ß-1,3-glucanase out of the protein inclusions of lutoids was revealed by pull-down. This interaction was pharmacologically verified by cytochalasin B-caused significant prolongation of the duration of latex flow in the field. CONCLUSIONS: The formation of protein-network by interactions of the proteins with anti-pathogen activity released from lutoids and accumulation of protein-network by binding to the cytoskeleton are crucial for the rapid occlusion of laticifer cells in rubber tree. The protein-network at the wounded site of laticifer cells provides not only a physical barrier but also a biochemical barrier to protect the wounded laticifer cells from pathogen invasion.

Mechanical contribution of secondary phloem to postural control in trees: the bark side of the force.

To grow straight, plants need a motor system that controls posture by generating forces to offset gravity. This motor function in trees was long thought to be only controlled by internal forces induced in wood. Here we provide evidence that bark is involved in the generation of mechanical stresses in several tree species. Saplings of nine tropical species were grown tilted and staked in a shadehouse and the change in curvature of the stem was measured after releasing from the pole and after removing the bark. This first experiment evidenced the contribution of bark in the up-righting movement of tree stems. Combined mechanical measurements of released strains on adult trees and microstructural observations in both transverse and longitudinal/tangential plane enabled us to identify the mechanism responsible for the development of asymmetric mechanical stress in the bark of stems of these species. This mechanism does not result from cell wall maturation like in wood, or from the direct action of turgor pressure like in unlignified organs, but is the consequence of the interaction between wood radial pressure and a smartly organized trellis structure in the inner bark.

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.

Impact of warming, moderate nitrogen addition and bark herbivory on BVOC emissions and growth of Scots pine (Pinus sylvestris L.) seedlings.

The changing climate will expose boreal forests to rising temperatures, increasing soil nitrogen (N) levels and an increasing risk of herbivory. The single and interaction effects of warming (+2 °C increase), moderate N addition (30 kg ha-1 year-1) and bark herbivory by large pine weevil (Hylobius abietis L.) on growth and emissions of biogenic volatile organic compounds (BVOCs) from shoots of Scots pine (Pinus sylvestris L.) seedlings were studied in growth chambers over 175 days. In addition, warming and N addition effects on shoot net photosynthesis (Pn) were measured. Nitrogen addition increased both shoot and root dry weights, whereas warming, in combination with herbivory, reduced stem height growth. Warming together with N addition increased current-year shoot Pn, whereas N effects on previous-year shoot Pn were variable over time. Warming decreased non-oxygenated monoterpene (MT) emissions in June and increased them in July. Of individual MT compounds, α-pinene, δ-3-carene, γ-terpinene and terpinolene were among the most frequently responsive compounds in warming treatments in the May-July period. Sesquiterpene emissions were observed only from warming treatments in July. Moderate N addition increased oxygenated monoterpenes in May, and MTs in June and September. However, N addition effect on MTs in June was clearer without warming than with warming. Bark herbivory tended to increase MT emissions in combination with warming and N addition 3 weeks after the damage caused by weevils. Of individual compounds in other BVOC blends, herbivory increased the emissions of methyl-benzene, benzene and hexanal in July. Hence, though both warming and N addition have a potential to change BVOC emissions from Scots pines, the N effect may also be partly cancelled by warming. Furthermore, herbivory pressure in combination with climate warming and N addition may, at least periodically, increase BVOC release to the atmosphere from young Scots pine seedlings.

Bark Colonization of Kiln-Dried Wood by the Walnut Twig Beetle: Effect of Wood Location and Pheromone Presence.

The walnut twig beetle (Pityophthorus juglandis Blackman) (Coleoptera: Curculionidae) is a regulated pest in the United States due to its causal role in thousand cankers disease of walnut trees, including the commercially valuable eastern black walnut (Juglans nigra L.). Several state quarantines designed to limit spread of P. juglandis regulate movement of kiln-dried walnut lumber that contains bark. Previous research demonstrated that P. juglandis will enter and re-emerge from bark of kiln-dried, J. nigra slabs subjected to extreme beetle pressure (baited with a pheromone lure and hung in infested J. nigra trees). This study evaluated P. juglandis bark colonization of both kiln-dried and fresh J. nigra slabs, varying the presence of aggregation pheromone and relative proximity to a beetle source. Wood treatment, slab location, and pheromone presence all significantly affected P. juglandis colonization, as assessed by subsequent beetle emergence. When placed on the ground directly beneath infested trees, kiln-dried slabs were not colonized, and fresh slabs were colonized only when baited with the pheromone lure (6/14 replicates). When placed in crowns of infested trees, kiln-dried slabs were colonized only when baited with pheromone (3/14 replicates), whereas fresh slabs were colonized with and without pheromone (14/14 and 1/13 replicates, respectively). Timing of emergence indicated that beetles did not reproduce in kiln-dried bark. Results suggest that the risk of kiln-dried walnut bark becoming colonized by the P. juglandis during movement of commercial wood products is very low. This information may be useful to government agencies that administer quarantines regulating the transport of walnut lumber.

Water uptake can occur through woody portions of roots and facilitates localized embolism repair in grapevine.

Water acquisition is thought to be limited to the unsuberized surface located close to root tips. However, there are recurring periods when the unsuberized surfaces are limited in woody root systems, and radial water uptake across the bark of woody roots might play an important physiological role in hydraulic functioning. Using X-ray microcomputed tomography (microCT) and hydraulic conductivity measurements (Lpr ), we examined water uptake capacity of suberized woody roots in vivo and in excised samples. Bark hydration in grapevine woody roots occurred quickly upon exposure to water (c. 4 h). Lpr measurements through the bark of woody roots showed that it is permeable to water and becomes more so upon wetting. After bark hydration, microCT analysis showed that absorbed water was utilized to remove embolism locally, where c. 20% of root xylem vessels refilled completely within 15 h. Embolism removal did not occur in control roots without water. Water uptake through the bark of woody roots probably plays an important role when unsuberized tissue is scarce/absent, and would be particularly relevant following large irrigation events or in late winter when soils are saturated, re-establishing hydraulic functionality before bud break.

Gradients and dynamics of inner bark and needle osmotic potentials in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst).

Preconditions of phloem transport in conifers are relatively unknown. We studied the variation of needle and inner bark axial osmotic gradients and xylem water potential in Scots pine and Norway spruce by measuring needle and inner bark osmolality in saplings and mature trees over several periods within a growing season. The needle and inner bark osmolality was strongly related to xylem water potential in all studied trees. Sugar concentrations were measured in Scots pine, and they had similar dynamics to inner bark osmolality. The sucrose quantity remained fairly constant over time and position, whereas the other sugars exhibited a larger change with time and position. A small osmotic gradient existed from branch to stem base under pre-dawn conditions, and the osmotic gradient between upper stem and stem base was close to zero. The turgor in branches was significantly driven by xylem water potential, and the turgor loss point in branches was relatively close to daily minimum needle water potentials typically reported for Scots pine. Our results imply that xylem water potential considerably impacts the turgor pressure gradient driving phloem transport and that gravitation has a relatively large role in phloem transport in the stems of mature Scots pine trees.

Exploring the bark thickness-stem diameter relationship: clues from lianas, successive cambia, monocots and gymnosperms.

Bark thickness is ecologically crucial, affecting functions from fire protection to photosynthesis. Bark thickness scales predictably with stem diameter, but there is little consensus on whether this scaling is a passive consequence of growth or an important adaptive phenomenon requiring explanation. With a comparative study across 913 species, we test the expectation that, if bark thickness-stem diameter scaling is adaptive, it should be possible to find ecological situations in which scaling is predictably altered, in this case between species with different types and deployments of phloem. 'Dicots' with successive cambia and monocots, which have phloem-free bark, had predictably thinner inner (mostly living) bark than plants with single cambia. Lianas, which supply large leaf areas with limited stem area, had much thicker inner bark than self-supporting plants. Gymnosperms had thicker outer bark than angiosperms. Inner bark probably scales with plant metabolic demands, for example with leaf area. Outer bark scales with stem diameter less predictably, probably reflecting diverse adaptive factors; for example, it tends to be thicker in fire-prone species and very thin when bark photosynthesis is favored. Predictable bark thickness-stem diameter scaling across plants with different photosynthate translocation demands and modes strongly supports the idea that this relationship is functionally important and adaptively significant.

A signature of tree health? Shifts in the microbiome and the ecological drivers of horse chestnut bleeding canker disease.

Host susceptibility to pathogens can be shaped by genetic, ecological, and evolutionary factors. The ability to predict the spread of disease therefore requires an integrated understanding of these factors, including effects of pests on pathogen growth and competition between pathogens and commensal microbiota for host resources. We examined interactions between the leaf-mining moth Cameraria ohridella, the bacterial causal agent of bleeding canker disease Pseudomonas syringae pv aesculi, and the bark-associated microbiota of horse chestnut (Aesculus hippocastanum) trees. Through surveys of > 900 trees from 60 sites in the UK, we tested for ecological or life history predictors of leaf miner infestation, bleeding canker, or coinfection. Using culture-independent sequencing, we then compared the bark microbiomes from 46 trees to measure the association between microbiome composition and key ecological variables, including the severity of disease. Both pest and pathogen were found to respond to tree characteristics, but neither explained damage inflicted by the other. However, we found a clear loss of microbial diversity and associated shift in microbiome composition of trees as a function of disease. These results show a link between bark-associated microbiota and tree health that introduces the intriguing possibility that tree microbiota play key roles in the spread of disease.

Retention of stored water enables tropical tree saplings to survive extreme drought conditions.

Trees generally maintain a small safety margin between the stem water potential (Ψstem) reached during seasonal droughts and the Ψstem associated with their mortality. This pattern may indicate that species face similar mortality risk during extreme droughts. However, if tree species vary in their ability to regulate Ψstem, then safety margins would poorly predict drought mortality. To explore variation among species in Ψstem regulation, I subjected potted saplings of six tropical tree species to extreme drought and compared their responses with well-watered plants and pretreatment reference plants. In the drought treatment, soil water potential reached <-10 MPa, yet three species, Bursera simaruba (L.) Sarg., Cavanillesia platanifolia (Bonpl.) Kunth and Cedrela odorata L. had 100% survival and maintained Ψstem near -1 MPa (i.e., desiccation-avoiding species). Three other species, Cojoba rufescens (Benth.) Britton and Rose, Genipa americana L. and Hymenaea courbaril L. had 50%, 0% and 25% survival, respectively, and survivors had Ψstem <-6 MPa (i.e., desiccation-susceptible species). The desiccation-avoiding species had lower relative water content (RWC) in all organs and tissues (root, stem, bark and xylem) in the drought treatment than in the reference plants (means 72.0-90.4% vs 86.9-97.9%), but the survivors of the desiccation-susceptible C. rufescens had much lower RWC in the drought treatment (44.5-72.1%). Among the reference plants, the desiccation-avoiding species had lower tissue density, leaf-mass fraction and lateral-root surface area (LRA) than the desiccation-susceptible species. Additionally, C. platanifolia and C. odorata had reduced LRA in the drought treatment, which may slow water loss into dry soil. Together, these results suggest that the ability to regulate Ψstem during extreme drought is associated with functional traits that favor retention of stored water and that safety margins during seasonal drought poorly predict survival during extreme drought.

Colonization Dynamics of Subcortical Insects on Forest Sites With Relatively Stressed and Unstressed Loblolly Pine Trees.

Loblolly pine (Pinus taeda L.) is the most important commercial tree species in the southeastern United States. Since the 1950s, there have been reports of loblolly pines showing reduced growth and increased mortality, particularly in central Alabama and western Georgia, United States; the phenomenon is termed as southern pine decline (SPD). Recently, the role of rhizophagous (root-feeding) insects in loblolly pine health within the context of SPD has come under greater scrutiny. We investigated the impacts of subcortical insects, particularly rhizophagous weevils (Coleoptera: Curculionidae), on loblolly pine health in northeastern Georgia. We created plots-representing a gradient of increased relative tree stress-from ungirdled trees, ungirdled trees baited with ethanol and turpentine (ungirdled-baited), and girdled trees. In total, 10,795 subcortical insects from four families (Buprestidae, Cerambycidae, Curculionidae, and Siricidae) and >82 species were trapped in two years. Almost half of the insects trapped (46% of individuals and 11% of species) were nonnative to North America. Insect captures in plots with girdled trees were 61 and 187% greater than those with ungirdled-baited and ungirdled trees, respectively. Tree treatment impacted captures of native, but not nonnative insects. Relative feeding area by the rhizophagous weevils Hylobius pales (Herbst) and Pachylobius picivorus (Germar) on pine twigs placed in pitfall traps was 1, 17, and 82% in plots with ungirdled, ungirdled-baited, and girdled trees, respectively. Hence, there was a strong association of native subcortical insects, especially rhizophagous weevils, with relatively highly stressed trees, confirming that they are secondary instead of primary pine colonizers.

Response of Organ Structure and Physiology to Autotetraploidization in Early Development of Energy Willow Salix viminalis.

The biomass productivity of the energy willow Salix viminalis as a short-rotation woody crop depends on organ structure and functions that are under the control of genome size. Colchicine treatment of axillary buds resulted in a set of autotetraploid S. viminalis var. Energo genotypes (polyploid Energo [PP-E]; 2n = 4x = 76) with variation in the green pixel-based shoot surface area. In cases where increased shoot biomass was observed, it was primarily derived from larger leaf size and wider stem diameter. Autotetraploidy slowed primary growth and increased shoot diameter (a parameter of secondary growth). The duplicated genome size enlarged bark and wood layers in twigs sampled in the field. The PP-E plants developed wider leaves with thicker midrib and enlarged palisade parenchyma cells. Autotetraploid leaves contained significantly increased amounts of active gibberellins, cytokinins, salicylic acid, and jasmonate compared with diploid individuals. Greater net photosynthetic CO2 uptake was detected in leaves of PP-E plants with increased chlorophyll and carotenoid contents. Improved photosynthetic functions in tetraploids were also shown by more efficient electron transport rates of photosystems I and II. Autotetraploidization increased the biomass of the root system of PP-E plants relative to diploids. Sections of tetraploid roots showed thickening with enlarged cortex cells. Elevated amounts of indole acetic acid, active cytokinins, active gibberellin, and salicylic acid were detected in the root tips of these plants. The presented variation in traits of tetraploid willow genotypes provides a basis to use autopolyploidization as a chromosome engineering technique to alter the organ development of energy plants in order to improve biomass productivity.

Bark water uptake promotes localized hydraulic recovery in coastal redwood crown.

Coastal redwood (Sequoia sempervirens), the world's tallest tree species, rehydrates leaves via foliar water uptake during fog/rain events. Here we examine if bark also permits water uptake in redwood branches, exploring potential flow mechanisms and biological significance. Using isotopic labelling and microCT imaging, we observed that water entered the xylem via bark and reduced tracheid embolization. Moreover, prolonged bark wetting (16 h) partially restored xylem hydraulic conductivity in isolated branch segments and whole branches. Partial hydraulic recovery coincided with an increase in branch water potential from about -5.5 ± 0.4 to -4.2 ± 0.3 MPa, suggesting localized recovery and possibly hydraulic isolation. As bark water uptake rate correlated with xylem osmotic potential (R(2) = 0.88), we suspect a symplastic role in transferring water from bark to xylem. Using historical weather data from typical redwood habitat, we estimated that bark and leaves are wet more than 1000 h per year on average, with over 30 events being sufficiently long (>24 h) to allow for bark-assisted hydraulic recovery. The capacity to uptake biologically meaningful volumes of water via bark and leaves for localized hydraulic recovery throughout the crown during rain/fog events might be physiologically advantageous, allowing for relatively constant transpiration.

Genotype variation in bark texture drives lichen community assembly across multiple environments.

A major goal of community genetics is to understand the influence of genetic variation within a species on ecological communities. Although well-documented for some organisms, additional research is necessary to understand the relative and interactive effects of genotype and environment on biodiversity, identify mechanisms through which tree genotype influences communities, and connect this emerging field with existing themes in ecology. We employ an underutilized but ecologically significant group of organisms, epiphytic bark lichens, to understand the relative importance of Populus angustifolia (narrowleaf cottonwood) genotype and environment on associated organisms within the context of community assembly and host ontogeny. Several key findings emerged. (1) In a single common garden, tree genotype explained 18-33% and 51% of the variation in lichen community variables and rough bark cover, respectively. (2) Across replicated common gardens, tree genotype affected lichen species richness, total lichen cover, lichen species composition, and rough bark cover, whereas environment only influenced composition and there were no genotype by environment interactions. (3) Rough bark cover was positively correlated with total lichen cover and richness, and was associated with a shift in species composition; these patterns occurred with variation in rough bark cover among tree genotypes of the same age in common gardens and with increasing rough bark cover along a -40 year tree age gradient in a natural riparian stand. (4) In a common garden, 20-year-old parent trees with smooth bark had poorly developed lichen communities, similar to their 10-year-old ramets (root suckers) growing in close proximity, while parent trees with high rough bark cover had more developed communities than their ramets. These findings indicate that epiphytic lichens are influenced by host genotype, an effect that is robust across divergent environments. Furthermore, the response to tree genotype is likely the result of genetic variation in the timing of the ontogenetic shift from smooth to rough bark allowing communities on some genotypes to assemble faster than those on other genotypes. Organisms outside the typical sphere of community genetics, such as lichens, can help address critical issues and connect plant genotype effects to long-established streams of biological research, such as ontogeny and community assembly.