Search

Resilience to drought represents an important focus for trees during climate change, with the aim of predicting the resistance and recovery of species worldwide. Previous studies mainly linked tree growth resilience to plant functional traits that are related to resource acquisition and investment. Here, we investigate a potential link between resilience and the amount of parenchyma tissue in wood, a multifunctional tissue that may provide various physiological benefits to drought‐related mechanisms.We compiled a global tree‐ring data set to evaluate the relationship between growth resistance, resilience or recovery from drought and xylem parenchyma tissue fractions of 50 angiosperms, which was complemented with a local study of nine species from Mt. Tiantong in China. We also assessed the influence of climate and phylogeny on these relationships.We found that growth recovery after drought was positively related to the fractions of total parenchyma locally and globally. This association remained statistically significant when accounting for the effects of climate and phylogeny. No other associations between parenchyma fractions and growth resilience metrics were statistically significant.Our results suggest that drought recovery of angiosperm trees is affected by the amount of parenchyma. Incorporating xylem parenchyma fraction with other traits, such as hydraulic traits, could therefore enhance our comprehension of how various angiosperm tree species will respond to future droughts. Further studies should focus on unravelling the physiological roles of xylem parenchyma fraction. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Due to land use change, green alder (Alnus alnobetula), formerly restricted to moist slopes, is now expanding to drier sun-exposed sites at the alpine treeline. The highly productive shrub is forming closed thickets, establishing nitrogen-saturated species poor shrublands. To evaluate wood anatomical adaptations to changing environmental conditions, we analyzed vessel characteristics (mean vessel area, MVA; vessel density, VD; and theoretic conductive area, TCA) and axial parenchyma abundance, as well as their distribution in the annual ring at a moist north-facing and a dry south-facing site at the alpine treeline on Mt. Patscherkofel (Central European Alps, Austria). Results revealed that lower soil water availability and enhanced evaporative demand did not affect MVA while VD and TCA were significantly reduced at the dry south-facing site. This suggests that in green alder, vessel size is a static trait whereas vessel number responds plastic. Limited water availability also triggered a significant increase in axial parenchyma, confirming the important role of xylem parenchyma in water relations. Harsh environmental conditions at the distributional limit of green alder may have affected xylogenesis, leading to a near semi-ring-porous distribution of vessels and an accumulation of parenchyma in the late growing season. We conclude that in a warmer and drier climate, growth limitation and physiological stress may set limits to the distribution of Alnus alnobetula at drought-prone sites in the alpine treeline ecotone. [ABSTRACT FROM AUTHOR]

Karst forests are habitats in which access to soil water can be challenging for plants. Therefore, safe and efficient xylem water transport and large internal water storage may benefit tree growth. In this study, we selected 22 tree species from a primary subtropical karst forest in southern China and measured their xylem anatomical traits, saturated water content (SWC), hydraulic conductivity (Ks) and embolism resistance (P 50). Additionally, we monitored growth of diameter at breast height (DBH) in 440 individual trees of various sizes over three consecutive years. Our objective was to analyze the relationships between xylem structure, hydraulic efficiency, safety, water storage and growth of karst tree species. The results showed significant differences in structure but not in hydraulic traits between deciduous and evergreen species. Larger vessel diameter, paratracheal parenchyma and higher SWC were correlated with higher Ks. Embolism resistance was not correlated with the studied anatomical traits, and no tradeoff with Ks was observed. In small trees (5–15 cm DBH), diameter growth rate (DGR) was independent of hydraulic traits. In large trees (>15 cm DBH), higher Ks and more negative P 50 accounted for higher DGR. From lower to greater embolism resistance, the size–growth relationship shifted from growth deceleration to acceleration with increasing tree size in eight of the 22 species. Our study highlights the vital contributions of xylem hydraulic efficiency and safety to growth rate and dynamics in karst tree species; therefore, we strongly recommend their integration into trait-based forest dynamic models. [ABSTRACT FROM AUTHOR]

Halophytes accumulate and sequester high concentrations of salt in vacuoles while maintaining lower levels of salt in the cytoplasm. The current data on cellular and subcellular partitioning of salt in halophytes are, however, limited to only a few dicotyledonous C3 species. Using cryo‐scanning electron microscopy X‐ray microanalysis, we assessed the concentrations of Na, Cl, K, Ca, Mg, P and S in various cell types within the leaf‐blades of a monocotyledonous C4 halophyte, Rhodes grass (Chloris gayana). We also linked, for the first time, elemental concentrations in chloroplasts of mesophyll and bundle sheath cells to their ultrastructure and photosynthetic performance of plants grown in nonsaline and saline (200 mM NaCl) conditions. Na and Cl accumulated to the highest levels in xylem parenchyma and epidermal cells, but were maintained at lower concentrations in photosynthetically active mesophyll and bundle sheath cells. Concentrations of Na and Cl in chloroplasts of mesophyll and bundle sheath cells were lower than in their respective vacuoles. No ultrastructural changes were observed in either mesophyll or bundle sheath chloroplasts, and photosynthetic activity was maintained in saline conditions. Salinity tolerance in Rhodes grass is related to specific cellular Na and Cl distributions in leaf tissues, and the ability to regulate Na and Cl concentrations in chloroplasts. [ABSTRACT FROM AUTHOR]

Summary: Parenchyma cells in the xylem store nonstructural carbohydrates (NSC), providing reserves of energy that fuel woody perennials through periods of stress and/or limitations to photosynthesis. If the capacity for storage is subject to selection, then the fraction of wood occupied by living parenchyma should increase towards stressful environments.Ray parenchyma fraction (RPF) and seasonal NSC dynamics were quantified for 12 conifers and three oaks along a transect spanning warm dry foothills (500 m above sea level) to cold wet treeline (3250 m asl) in California's central Sierra Nevada.Mean RPF was lower for both conifer and oak species with warmer dryer ranges. RPF variability increased with elevation or in relation to associated climatic variables in conifers – treeline‐dominant Pinus albicaulis had the lowest mean RPF measured (c. 3.7%), but the highest environmentally standardized variability index. Conifer RPF variability was explained by environment, increasing predominantly towards cooler wetter range edges. In oaks, NSC was explained by environment – values increasing for evergreen and decreasing for deciduous oaks with elevation. Lastly, all species surveyed appear to prioritize filling available RPF with sugar to achieve molarities that balance reasonable tensions over starch to maximize stored carbon.RPF responds to environment but is unlikely to spatially constrain NSC storage. [ABSTRACT FROM AUTHOR]

Plant adaptive potential is critically dependent on efficient communication and coordination of resource allocation and signalling between above and below-ground plant parts. Control of xylem ion loading plays an important role in this process. This review focuses on the molecular identity, tissue-specific expression patterns, and transcriptional and post-translational regulation of transporters mediating xylem loading of Na+, K+, and Cl- in plants grown under abiotic stress conditions such as drought, salinity, and soil flooding. The data are discussed in the context of breeding crops for stress resilience, which remains one of the highest priorities for dealing with the global food security challenge. This resilience was present in wild ancestors, but has been lost during domestication of crop species and exacerbated by the selection for higher yielding cultivars over the last 100 years. Thus, progress in the field requires a major rethink of current paradigms in crop breeding and the targeting of previously unexplored genes and traits. We argue that control of xylem ion loading is one of these traits and represents an unexplored opportunity for genetic improvement of plant germplasms. [ABSTRACT FROM AUTHOR]

Trees experience low apoplastic water potential frequently in most environments. Low apoplastic water potential increases the risk of embolism formation in xylem conduits and creates dehydration stress for the living cells. We studied the magnitude and rate of xylem diameter change in response to decreasing apoplastic water potential and the role of living parenchyma cells in it to better understand xylem diameter changes in different environmental conditions. We compared responses of control and heatinjured xylem of Pinus sylvestris (L.) and Populus tremula (L.) branches to decreasing apoplastic water potential created by osmotic stress, desiccation and freezing. It was shown that xylem in control branches shrank more in response to decreasing apoplastic water potential in comparison with the samples that were preheated to damage living xylem parenchyma. By manipulating the osmotic pressure of the xylem sap, we observed xylem shrinkage due to decreasing apoplastic water potential even in the absence of water tension within the conduits. These results indicate that decreasing apoplastic water potential led to withdrawal of intracellular water from the xylem parenchyma, causing tissue shrinkage. The amount of xylem shrinkage per decrease in apoplastic water potential was higher during osmotic stress or desiccation compared with freezing. During desiccation, xylem diameter shrinkage involved both dehydration-related shrinkage of xylem parenchyma and water tension-induced shrinkage of conduits, whereas dehydration-related shrinkage of xylem parenchyma was accompanied by swelling of apoplastic ice during freezing. It was also shown that the exchange of water between symplast and apoplast within xylem is clearly faster than previously reported between the phloem and the xylem. Time constant of xylem shrinkage was 40 and 2 times higher during osmotic stress than during freezing stress in P. sylvestris and P. tremula, respectively. Finally, it was concluded that the amount of water stored in the xylem parenchyma is an important reservoir for trees to buffer daily fluctuations in water relations. [ABSTRACT FROM AUTHOR]

Main conclusion: Cell wall fluorescence and immunocytochemistry demonstrate that xylem parenchyma cell walls do not show changes in structure and composition related to gravitropic response comparable to those of tracheids, even when they have lignified secondary cell walls. Tracheid cell walls in compression wood have altered composition and structure which generates the strain responsible for correction of stem lean as part of the gravitropic response of woody plants. Xylem parenchyma cell walls vary among conifer species and can be lignified secondary walls (spruce) or unlignified primary walls (pine). It can be expected that xylem parenchyma with lignified secondary cell walls might show features of compression wood comparable to those of tracheids that have a similar type of cell wall. A comparison of xylem parenchyma cell walls in normal and compression wood in species with lignified and non-lignified parenchyma cell walls provides a unique opportunity to understand the process of reaction wood formation in conifers. Using both UV/visible fluorescence microscopy of cell wall fluorophores and immunocytochemistry of galactan and mannan epitopes, we demonstrate that xylem parenchyma cell walls do not show the changes in composition and structure typical of compression wood tracheids. Adjacent cells of different types but with similar cell wall structure can undergo cell wall developmental changes related to support or defence functions independent of their neighbours. Tracheids are sensitive to gravitropic signals while xylem parenchyma cells are not. [ABSTRACT FROM AUTHOR]

Ultrasonic acoustic emission (UAE) methods have been applied for the detection of freeze-thaw-induced embolism formation in water conduits of tree species. Until now, however, the exact source(s) of UAE has not been identified especially in angiosperm species, in which xylem tissues are composed of diverse types of cells. In this study, UAE was recorded from excised branches of walnut (Juglans regia cv. Franquette) during freeze-thaw cycles, and attempts were made to characterize UAEs generated by cavitation events leading to embolism formation according to their properties. During freeze-thaw cycles, a large number of UAEs were generated from the sample segments. However, the cumulative numbers of total UAE during freeze-thawing were not correlated with the percentage loss of hydraulic conductivity after thawing, suggesting that the sources of UAE were not only cavitation leading to embolism formation in vessels. Among the UAEs, cumulative numbers of UAEs with absolute energy >10.0 fJ strongly correlated with the increase in percentage loss of hydraulic conductivity. The high absolute energy of the UAEs might reflect the formation of large bubbles in the large lumen of vessels. Therefore, UAEs generated by cavitation events in vessels during freeze-thawing might be distinguished from other signals according to their magnitudes of absolute energy. On the other hand, the freezing of xylem parenchyma cells was followed by a certain number of UAEs. These results indicate the possibility that UAE methods can be applied to the detection of both freeze-thaw-induced embolism and supercooling breakdown in parenchyma cells in xylem. [ABSTRACT FROM AUTHOR]

An introduction is presented in which the editor discusses various reports within the issue on topics including the role of living cell in making new wood and in their water transport system, the functions of living cells in angiosperms, and the lifespan of vessels.

The morphological variation and structure-function relationships of xylem parenchyma still remain open to discussion. We analyzed the three-dimensional structure of a poorly known type of xylem parenchyma with disjunctive walls in the tropical hardwood Okoubaka aubrevillei (Santalaceae). Disjunctive cells occurred among the apotracheal parenchyma cells and at connections between axial and ray parenchyma cells. The disjunctive cells were partly detached one from another, but their tubular structures connected them into a continuous network of axial and ray parenchyma. The connecting tubules had thick secondary walls and simple pits with plasmodesmata at the points where one cell contacted a tubule of another cell. The imperforate Iracheary elements of the ground tissue were seven times longer than the axial parenchyma strands, a fact that supports a hypothesis that parenchyma cells develop disjunctive walls because they are pulled apart and partly separated during the intrusive growth of fibers. We discuss unresolved details of the formation of disjunctive cell walls and the possible biomechanical advantage of the wood with disjunctive parenchyma: the proportion of tissue that improves mechanical strength is increased by the intrusive elongation of fibers (thickwalled tracheids), whereas the symplastic continuum of the parenchyma is maintained through formation of disjunctive cells. [ABSTRACT FROM AUTHOR]

The structure and transport properties of pit membranes at the interface between the metaxylem and xylem parenchyma cells and the possible role of these pit membranes in solute transfer to the phloem were investigated. Electron microscopy revealed a fibrillar, almost tubular matrix within the pit membrane structure between the xylem vessels and xylem parenchyma of leaf blade bundles in rice (Oryza sativa). These pits are involved primarily with regulating water flux to the surrounding xylem parenchyma cells. Vascular parenchyma cells contain large mitochondrial populations, numerous dictyosomes, endomembrane complexes, and vesicles in close proximity to the pit membrane. Taken collectively, this suggests that endocytosis may occur at this interface. A weak solution of 5,6-carboxyfluorescein diacetate (5,6-CFDA) was applied to cut ends of leaves and, after a minimum of 30 min, the distribution of the fluorescent cleavage product, 5,6-carboxyfluorescein (5,6-CF), was observed using confocal microscopy. Cleavage of 5,6-CFDA occurred within the xylem parenchyma cells, and the non-polar 5,6-CF was then symplasmically transported to other parenchyma elements and ultimately, via numerous pore plasmodesmata, to adjacent thick-walled sieve tubes. Application of Lucifer Yellow, and, separately, Texas Red-labelled dextran (10 kDa) to the transpiration stream, confirmed that these membrane-impermeant probes could only have been offloaded from the xylem via the xylem vessel–xylem parenchyma pit membranes, suggesting endocytotic transmembrane transfer of these membrane-impermeant fluorophores. Accumulation within the thick-walled sieve tubes, but not in thin-walled sieve tubes, confirms the presence of a symplasmic phloem loading pathway, via pore plasmodesmata between xylem parenchyma and thick-walled sieve tubes, but not thin-walled sieve tubes. [ABSTRACT FROM PUBLISHER]

Oxygen levels as low as 1–5% (gaseous mole fraction) occur in secondary xylem, but it is not known if there is a consistent pattern of decline in O2 from the cambium toward the pith, or whether parenchyma cells experience hypoxic conditions deep within the stem. We developed a system for repeatedin situmeasurement of O2 at different depths within stems ofAcer rubrum,Fraxinus americana,Tsuga canadensis, andQuercus rubra. In summer during active transpiration, O2 declined from the cambium toward the heartwood boundary inF. americana,T. canadensisandQ. rubra, but remained constant inA. rubrum. Average sapwood O2 was about 10%, with the lowest values observed in the innermost sapwood around 3–5%. Before spring leaf flush, O2 content in the outer sapwood was reduced inQ. rubraandT. canadensisrelative to summer, and was occasionally lower than in the inner sapwood. Sapwood respiration inT.canadensiswas constant above 5% O2, but reduced by about 65% at 1% O2. InF.americana, sapwood respiration was constant above 10% O2 but reduced by 25% at 5% O2, and by 75% at 1% O2, the most extreme inhibition observed. However, when prolonged (72 h) exposure to 1%, 5% and 10% O2 was followed by re-equilibration to 10% O2, no inhibition was found. Given the minor (and reversible) effect of low O2 on parenchyma metabolism at levels common in the inner sapwood, it is unlikely that O2 content severely limits parenchyma respiration or leads to parenchyma cell death during sapwood senescence. Within-stem O2 levels may instead be most relevant to metabolism in the cambial zone and phloem, for which sapwood could serve as a significant source of O2. [ABSTRACT FROM AUTHOR]

The maintenance of Mg2+ homeostasis in the plant is essential for viability. This review describes Mg2+ functions and balancing in plants, with special focus on the existing knowledge of the involved transport mechanisms. Mg2+ is essential for the function of many cellular enzymes and for the aggregation of ribosomes. Mg2+ concentrations also modulate ionic currents across the chloroplast and the vacuolar membranes, and might thus regulate ion balance in the cell and stomatal opening. The significance of Mg2+ homeostasis has been particularly established with regard to Mg2+'s role in photosynthesis. Mg2+ is the central atom of the chlorophyll molecule, and fluctuations in its levels in the chloroplast regulate the activity of key photosynthetic enzymes. Relatively little is known of the proteins mediating Mg2+ uptake and transport in plants. The plant vacuole seem to play a key role in Mg2+ homeostasis in plant cells. Physiological and molecular evidence indicate that Mg2+ entry to the vacuole is mediated by Mg2+/H+ exchangers. The Arabidopsis vacuolar Mg2+/H+ exchanger, AtMHX, is highly transcribed at the vascular tissue, apparently most abundantly at the xylem parenchyma. Inclusion of Mg2+ ions into the vacuoles of this tissue may determine their partitioning between the various plant organs. Impacts of Mg2+ imbalance are described with respect for both plant physiology and for its nutritional value to animal and human. [ABSTRACT FROM AUTHOR]

Rapidly developing tumours at hypocotyls of Ricinus communis, induced by Agrobacterium tumefaciens strain C58, were characterized by strong differentiation of vascular bundles and their functional connection to the host bundles. The stem/tumour interface showed increased xylem, with numerous vessels accompanied by multiseriate unlignified rays. To know how nutrients efficiently accumulate in the tumour sink tissue, cell electropotentials (Em) in cross‐sections were mapped. The measured cells were identified by injected Lucifer Yellow. Xylem and phloem parenchyma cells and stem/tumour‐located rays hyperpolarized to Em values of about −170 mV, which suggest high plasma membrane proton pump activities. Rapidly dividing cells of cambia or small tumour parenchyma cells had low Em. The tumour aerenchyma and the stem cortex cells displayed values close to the energy‐independent diffusion potential. The lowest values were recorded in stem pith cells. Cell K+ concentrations largely matched the respective Em. The pattern of individual cell electropotentials was supplemented by whole organ voltage measurements. The voltage differences between the tumour surface and the xylem perfusion solution in stems attached to the tumours, the trans‐tumour electropotentials (TTP), confirm the findings of respiration‐dependent and phytohormone‐stimulated high plasma membrane proton pump activity in intact tumours, mainly in the xylem and phloem parenchyma and ray cells. TTPs were inhibited by addition of NaN3, CN− plus SHAM or N2 gas in the xylem perfusion solution and by external N2 flushing. The data provide functional evidence for the structural basis of priority over the host shoot in nutrient flow from the stem to the tumour. [ABSTRACT FROM PUBLISHER]

Cellular functions require adequate homeostasis of several divalent metal cations, including Mg2+ and Zn2+. Mg2+, the most abundant free divalent cytoplasmic cation, is essential for many enzymatic reactions, while Zn2+ is a structural constituent of various enzymes. Multicellular organisms have to balance not only the intake of Mg2+ and Zn2+, but also the distribution of these ions to various organs. To date, genes encoding Mg2+ transport proteins have not been cloned from any multicellular organism. We report here the cloning and characterization of an Arabidopsis thaliana transporter, designated AtMHX, which is localized in the vacuolar membrane and functions as an electrogenic exchanger of protons with M2+ and Zn2+ ions. Functional homologs of AtMHX have not been cloned from any organism. Ectopic overexpression of AtMHX in transgenic tobacco plants render them sensitive to growth on media containing elevated levels of Mg2+ or Zn2+, but does not affect the total amounts of these minerals in shoots of the transgenic plants. AtMHX mRNA is mainly found at the vascular cylinder, and a large proportion of the mRNA is localized in close association with the xylem tracheary elements. This localization suggests that AtMHX may control The partitioning of Mg2+ and Zn2+ between the various plant organs. [ABSTRACT FROM AUTHOR]

Summary: Caffeic acid 3-O-methyltransferase (COMT) and caffeoyl CoA 3-O-methyltransferase (CCOMT) catalyze parallel reactions that are believed to be involved in the biosynthesis of lignin monomers. Antisera specific for alfalfa (Medicago sativa L.) COMT or CCOMT were raised against the enzymes expressed inEscherichia coli, and were used for immunolocalization studies in lignifying alfalfa stem tissue. Both COMT and CCOMT were localized to xylem parenchyma cells, as assessed by light microscopy and immunocytochemistry. Electron microscopy revealed that both enzymes were located in the cytoplasm of xylem parenchyma cells, and to a lesser extent, in the cytoplasm of phloem cells. There was no significant difference in the localization pattern of COMT and CCOMT, suggesting that the two enzymes may be part of a metabolic grid leading to production of lignin monomers in lignifying tissue of mature alfalfa stem internodes. [ABSTRACT FROM AUTHOR]

In order to study the mechanism and regulation of K+ resorption from the xylem by the cells that border the xylem vessels (the xylem parenchyma cells), K+ inward-rectifying channels (KIRCs) in the plasma membrane of xylem parenchyma cells from Hordeum vulgare L. cv. Apex were studied using the patch-clamp technique. In the inside-out configuration, three different types of K+ channel and a further K+ conductance could be identified. Two of these channels, named KIRC1 and KIRC2, were activated by guanosine 5′-[β,γ-imido]triphosphate (Gpp(NH)p; 150 μM), a non-hydrolyzable derivative of GTP, indicating that channel activity was up-regulated by G-proteins; modulation of channel activity occurred via a membrane-delimited pathway, since the effect could be demonstrated in cell-free patches. At 100 mM external K+, KIRC1 had a conductance of 8 pS. There was no effect of ATP on channel activity. Likewise, addition of 150 μM guanosine 5′-[β-thio]diphosphate (GDPβS) or adenosine 5′-[γ-thio]triphosphate (ATPγS) failed to activate KIRC1, indicating nucleotide specificity of the effect. A second K+ channel, activated by Gpp(NH)p (KIRC2) with gating properties clearly different from the first one was less frequently observed. Four different substates could be identified; the main level had a conductance of about 2 pS. Gating below the Nernst potential of K+ (EK) was voltage-independent. The channel closed at potentials more positive than EK. A third, hyperpolarization-activated K+ channel, KIRC3, with a low open probability was encountered in inside-out patches. It had a conductance of 45 pS in 100 mM K+. Channel activity was not affected by the addition of G-protein modulators. Moreover, slowly activating inward currents carried by K+ were recorded in several patches that are ascribed to a `subpicosiemens conductance'. Neither GDPβS nor Gpp(NH)p appeared to have an effect on the currents. Whole-cell measurements with these G-protein modulators included in the pipette solution were in general agreement with the results obtained on cell-free patches. A statistical evaluation revealed that time-dependent inward currents were larger when the G-protein activator Gpp(NH)p was included in the pipette medium compared to measurements with the inhibitor GDPβS. With the GTP analogue, an additional instantaneous component was elicited that was ascribed to KIRC2 activity. Data are discussed with respect to the putative role of G-proteins in conveying hormonal signals. Regulation by G-protein may either serve to fine-tune K+ uptake by xylem parenchyma cells or to initiate depolarization, followed by salt-efflux through depolarization-activated cation and anion channels. [ABSTRACT FROM AUTHOR]

Treatment of stem sections of peach ( Prunus persica (L.) Batsch) and flowering dogwood ( Cornus florida L.) with macerase, an enzyme mixture rich in pectinase, for 24-48 h resulted in a complete flattening of the low-temperature exotherm (LTE) as determined by differential thermal analysis (DTA). Ultrastructural analysis of macerase-treated tissue demonstrated a nearly complete digestion of the pit membrane (black cap and primary cell-wall) of nearly 100% of the xylem-parenchyma cells examined after 48 h of exposure to the enzyme. Additionally, the underlying amorphous layer was partially degraded in up to 57% of the cells examined. The macerase treatment had no visible effect on secondary cell-walls of xylem tissue. In contrast, treatment of stem tissue with cellulysin (mostly cellulase) resulted in a shift of the LTE to warmer temperatures as determined by DTA, and a digestion of only the outermost layer of the pit membrane in nearly 100% of the cells examined, with little or no effect on the underlying layers. Treatment of tissue with 25 mM sodiumphosphate buffer also resulted in a shift of the LTE to warmer temperatures but the shift was not as great as in cellulysin-treated tissue. The shift was associated with a partial degradation of the outermost layer of the pit membrane in dogwood (33-45% of the cells examined) but not in peach (3-7% of the cells). Collectively, the data indicate that pectins may be an integral structural element of the pit membrane and that this portion of the cell-wall, along with the underlying amorphous layer, play a major role in forming a barrier to water movement and growth of ice crystals. This barrier allows xylem parenchyma of some species of woody plants to undergo deep supercooling. [ABSTRACT FROM AUTHOR]

• Scots pines of different canopy status was studied. • The amount of wood rays was assessed in the outermost 30 tree-rings. • The amount of wood rays was highly variable among individuals. • Dimensions of wood rays differed between earlywood, latewood and canopy status. • The inter-annual variation of wood rays was related to prior-year conditions. Non-structural carbohydrates (NSC) reserves are crucial for trees to cope with weather extremes, thus to ensure their survival and ecological plasticity. The NSC reserves can depend on social status, suggesting uneven plasticity of trees at the stand level. In stemwood of Scots pine (Pinus sylvestris L.), which is a widespread and important species, NSC reserves are stored in parenchyma in wood rays (WR). The quantity of WRs is adjusted intra-annually, allowing retrospective analysis of factors affecting their formation. Accordingly, the differences in WR quantity in stemwood of dominant and intermediate (canopy trees with reduced and narrow crowns) maturing Scots pine were assessed by quantitative wood anatomy. Tangential cuts from the outermost 30 tree-rings were analysed. The relative ray area was intermediate, i.e. covering about 5% of the tangential cut, yet expressed high individuality among the trees. The size and amount of WR mainly differed between the earlywood and latewood; WRs in latewood were higher although narrower in comparison to earlywood, yet their total amount was higher in earlywood. Canopy status had only a slight effect, as quantity and height of WR tended to be higher for the intermediate trees, particularly in earlywood. The size and quantity of WR expressed inter-annual variation, which was mainly related to the meteorological conditions prior to the formation of the tree-ring (previous summer and autumn) indicating legacy effects of climatic factors on NSC and susceptibility of trees to cumulative effects of weather extremes. However, the climatic signals in the inter-annual variation of WR were weaker than observed before, likely due to location of the studied stand in the mid-part of the species range. Nevertheless, the observed differences in mean values and inter-annual variation of WR suggested a within-species diversity of carbon allocation patterns, supporting adaptability of the species. [ABSTRACT FROM AUTHOR]

Woody plants are characterised by a highly complex vascular system, wherein the secondary xylem (wood) is responsible for the axial transport of water and various substances. Previous studies have focused on the dead conductive elements in this heterogeneous tissue. However, the living xylem parenchyma cells, which constitute a significant functional fraction of the wood tissue, have been strongly neglected in studies on tree biology. Although there has recently been increased research interest in xylem parenchyma cells, the mechanisms that operate in these cells are poorly understood. Therefore, the present review focuses on selected roles of xylem parenchyma and its relevance in wood functioning. In addition, to elucidate the importance of xylem parenchyma, we have compiled evidence supporting the hypothesis on the significance of parenchyma cells in tree functioning and identified the key unaddressed questions in the field. [ABSTRACT FROM AUTHOR]