Conifer Xylem Research Articles

Insights into asynchronous changes of cell wall polymers accumulated in different cell types during conifer xylem differentiation

An improved understanding of the events involved in cell wall polymers deposition during xylem development could provide new scientific ways for molecular regulation and biomass utilization. Axial and radial cells are spatially heterogeneous and have highly cross-correlated developmental behavior, whereas the deposition of corresponding cell wall polymers during xylem differentiation is less studied. To clarify our hypothesis that cell wall polymers of two cell types accumulated asynchronously, we performed hierarchical visualization, including label-free in situ spectral imaging of different polymer compositions during the development of Pinus bungeana. In axial tracheids, the deposition of cellulose and glucomannan was observed on earlier stages of secondary wall thickening than that of xylan and lignin, while xylan distribution was strongly related to spatial distribution of lignin during differentiation. The content of lignin and polysaccharides increased by over 130 % and 60 % respectively when the S3 layer was formed, compared to the S2 stage. In ray cells, the deposition of crystalline cellulose, xylan, and lignin was generally lagged compared to that in corresponding axial tracheids, although the process followed a similar order. The concentration of lignin and polysaccharides in ray cells was only approximately 50 % of that in the axial tracheids during secondary wall thickening.

Cradle for the newborn Monochamus saltuarius: Microbial associates to ward off entomopathogens and disarm plant defense.

The Japanese pine sawyer, Monochamus saltuarius, as a beetle vector of Bursaphelenchus xylophilus (pine wood nematode), is an economically important forest pest in Eurasia. To feed on the phloem and xylem of conifers, M.saltuarius needs to overcome various stress factors, including coping with entomopathogenic bacteria and also various plant secondary compounds (PSCs). As an important adaptation strategy to colonize host trees, M.saltuarius deposit eggs in oviposition pits to shield their progeny. These pits harbor bacterial communities that are involved in the host adaptation of M.saltuarius to the conifers. However, the composition, origin, and functions of these oviposition pit bacteria are rarely understood. In this study, we investigated the bacterial community associated with M.saltuarius oviposition pits and their ability to degrade PSCs. Results showed that the bacterial community structure of M.saltuarius oviposition pits significantly differed from that of uninfected phloem. Also, the oviposition pit bacteria were predicted to be enriched in PSC degradation pathways. The microbial community also harbored a lethal strain of Serratia, which was significantly inhibited. Meanwhile, metatranscriptome analysis indicated that genes involved in PSCs degradation were expressed complementarily among the microbial communities of oviposition pits and secretions. Invitro degradation showed that bacteria cultured from oviposition pits degraded more monoterpenes and flavonoids than bacteria cultured from uninfected phloem isolates. Disinfection of oviposition pits increased the mortality of newly hatched larvae and resulted in a significant decrease in body weight in the early stages. Overall, our results reveal that M.saltuarius construct oviposition pits that harbor a diverse microbial community, with stronger PSCs degradation abilities and a low abundance of entomopathogenic bacteria, resulting in the increased fitness of newly hatched larvae.

Variation in Tracheid Dimensions of Conifer Xylem Reveals Evidence of Adaptation to Environmental Conditions.

Globally distributed extant conifer species must adapt to various environmental conditions, which would be reflected in their xylem structure, especially in the tracheid characteristics of earlywood and latewood. With an anatomical trait dataset of 78 conifer species growing throughout China, an interspecific study within a phylogenetic context was conducted to quantify variance of tracheid dimensions and their response to climatic and soil conditions. There was a significant difference in tracheid diameter between earlywood and latewood while no significant difference was detected in tracheid wall thickness through a phylogenetically paired t-test. Through a phylogenetic principle component analysis, Pinaceae species were found to be strongly divergent in their tracheid structure in contrast to a conservative tracheid structure in species of Cupressaceae, Taxaceae, and Podocarpaceae. Tracheid wall thickness decreased from high to low latitudes in both earlywood and latewood, with tracheid diameter decreasing for latewood only. According to the most parsimonious phylogenetic general least square models, environment and phylogeny together could explain about 21∼56% of tracheid structure variance. Our results provide insights into the effects of climate and soil on the xylem structure of conifer species thus furthering our understanding of the trees’ response to global change.

Hydraulic Function Analysis of Conifer Xylem Based on a Model Incorporating Tracheids, Bordered Pits, and Cross-Field Pits

Wood has a highly complex and anisotropic structure. Its xylem characteristics are key in determining the hydraulic properties of plants to transport water efficiently and safely, as well as the permeability in the process of wood impregnation modification. Previous studies on the relationship between the xylem structure and hydraulic conductivity of conifer have mainly focused on tracheids and bordered pits, with only a few focusing on the conduction model of cross-field pits which connect tracheids and rays. This study takes the xylem structure of conifer as an example, drawing an analogy between water flow under tension and electric current, and extends the model to the tissue scale, including cross-field pits by establishing isometric scaling. The structure parameters were collected by scanning electron microscopy and transmission electron microscopy. The improved model can quantify the important hydraulic functional characteristics of xylem only by measuring the more easily obtained tracheid section size. Then, this model was applied to quantify the relationship between the xylem anatomical structure and hydraulic properties in the pine (Pinus sylvestris L. var. mongholica Litv.) and the spruce (Picea koraiensis Nakai), and also to evaluate the effects of the number and size of cross-field pits on xylem conduction. The results showed that the growth ring conduction value of the pine was more than twice that of the spruce for the two tree species with similar growth widths in this study. The tracheid wall resistance of the pine reflected the result of the interaction of the size and number of cross-field pits, in comparison, the wall resistance of the spruce was more sensitive to the number of cross-field pits. Finally, the calculation output of the new model was cross-validated with the literature, which verified the accuracy and effectiveness of the model. This study provides an effective and complete solution for xylem conductivity measurement and the study of wood ecophysiological diversity and processing.

Non-linear Response to Cell Number Revealed and Eliminated From Long-Term Tracheid Measurements of Scots Pine in Southern Siberia.

Dendroclimatic research offers insight into tree growth–climate response as a solution to the forward problem and provides reconstructions of climatic variables as products of the reverse problem. Methodological developments in dendroclimatology have led to the inclusion of a variety of tree growth parameters in this field. Tree-ring traits developed during short time intervals of a growing season can potentially provide a finer temporal scale of both dendroclimatic applications and offer a better understanding of the mechanisms of tree growth reaction to climatic variations. Furthermore, the transition from classical dendroclimatic studies based on a single integral variable (tree-ring width) to the modern multitude of quantitative variables (e.g., wood anatomical structure) adds a lot of complexity, which mainly arises from intrinsic feedbacks between wood traits and muddles seasonality of registered climatic signal. This study utilized life-long wood anatomical measurements of 150- to 280-year-old trees of Pinus sylvestris L. growing in a moisture-sensitive habitat of the forest-steppe of Southern Siberia (Russia) to investigate and eliminate legacy effect from cell production in tracheid traits. Anatomical parameters were calculated to describe the results of the three main subsequent stages of conifer xylem tracheid development, namely, cell number per radial file in the ring, mean and maximum cell radial diameter, and mean and maximum cell-wall thickness. Although tree-ring width was almost directly proportional to cell number, non-linear relationships with cell number were revealed in tracheid measurements. They exhibited a stronger relationship in the areas of narrow rings and stable anatomical structure in wider rings. The exponential models proposed in this study demonstrated these relationships in numerical terms with morphometric meaning. The ratio of anatomical measurements to their modeled values was used to develop long-term anatomical chronologies, which proved to retain information about climatic fluctuations independent of tree-ring width (cell number), despite decreased common signal.

Analysis of the Structure and Hydraulic Function of Bordered Pits Using the Lattice Boltzman Method

Fluid flow between adjacent tracheids is realized through bordered pits in the xylem of conifers. The pit has an extremely small size and a highly complex structure. This paper presents a mesoscopic analytical method for the relationship between the pit structure and its hydraulic characteristics through mathematical modeling using the lattice Boltzmann method (LBM) and curved boundary treatment. Mongolian Scots pine were selected as the research subject of this study, and the bordered pit structure parameters was collected by scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and the original geometric features were maintained for direct modeling analysis. The model revealed the relationship between various components of the bordered pit and liquid flow velocity/resistance, indicating that margo is the main factor affecting flow resistance. Further anatomical investigation separately analyzed the influence of change in a single factor, including pit diameter, pit aperture diameter, pit depth, torus diameter, and margo thickness, on the overall flow and pressure drop to confirm the importance of various factors in this relationship. Additionally, the influence of pore size and pore location distribution in the margo on the flow rate and pressure drop was further analyzed quantitatively. The results showed that the flow rate through individual pores is the result of the combined effect of pore area and radial position of the pore in the margo. Our study promotes the research and application of the mesoscopic model LBM in simulating flow conditions in the complex flow field of pits, which realizes the numerical analysis of the flow field model based on individualized real bordered pits. In comparison with the classical macroscopic model, the accuracy and effectiveness of the proposed model are proved. This research can provide a promising method for analyzing the physiological and ecological functions of conifer and realizing the efficient utilization of wood resources.

Jan J. Lyczakowski.

Jan J. Lyczakowski.

Fossil charcoals from the Lower Jurassic challenge assumptions about charcoal morphology and identification

AbstractCharcoal morphometry is increasingly employed in the analysis of Quaternary sediments, but has not been applied in studies of earlier sediments. We present findings relating to charcoal morphometry and morphotypes in a Toarcian sequence, which contains a preponderance of highly elongate forms, and morphometric values well outside expected ranges. SEM imaging reveals elongate particles of two distinct kinds. One originates in conifer xylem, and may be associated with formation at high levels of heating. It is proposed that the other may derive from tree ferns, or from plants with similar growth forms. Our results show the importance of recognizing the wide morphological variation that exists in sedimentary charcoal. Failing to do so may result in serious errors in its identification, interpretation and quantification.

針葉樹木部からの氷核形成阻害物質の検出(平成17年度 第51回低温低物工学会研究報告)

針葉樹木部からの氷核形成阻害物質の検出(平成17年度 第51回低温低物工学会研究報告)

NorWood: a gene expression resource for evo-devo studies of conifer wood development.

Summary The secondary xylem of conifers is composed mainly of tracheids that differ anatomically and chemically from angiosperm xylem cells. There is currently no high‐spatial‐resolution data available profiling gene expression during wood formation for any coniferous species, which limits insight into tracheid development.RNA‐sequencing data from replicated, high‐spatial‐resolution section series throughout the cambial and woody tissues of Picea abies were used to generate the NorWood.conGenIE.org web resource, which facilitates exploration of the associated gene expression profiles and co‐expression networks.Integration within PlantGenIE.org enabled a comparative regulomics analysis, revealing divergent co‐expression networks between P. abies and the two angiosperm species Arabidopsis thaliana and Populus tremula for the secondary cell wall (SCW) master regulator NAC Class IIB transcription factors. The SCW cellulose synthase genes (CesAs) were located in the neighbourhoods of the NAC factors in A. thaliana and P. tremula, but not in P. abies. The NorWood co‐expression network enabled identification of potential SCW CesA regulators in P. abies.The NorWood web resource represents a powerful community tool for generating evo‐devo insights into the divergence of wood formation between angiosperms and gymnosperms and for advancing understanding of the regulation of wood development in P. abies.

Tree physiology: Tracking tree carbon gain.

Wood formation drives the storage of carbon in trees and other woody plants. A detailed analysis of conifer xylem tissue formation reveals how the intra-annual dynamics of biomass production are recorded in wood.

Lipid-content-normalized polycyclic aromatic hydrocarbons (PAHs) in the xylem of conifers can indicate historical changes in regional airborne PAHs

The temporal variation of polycyclic aromatic hydrocarbons (PAHs) concentrations as well as the lipid content in the xylem of Masson pine trees sampled from the same site were determined and compared with the days of haze occurrence and with the historical PAHs reported in sedimentary cores. The patterns of the lipid content as well as the PAH concentrations based on the xylem dry weight (PAHs-DW) decreased from the heartwood to the sapwood. The trajectories of PAHs normalized by xylem lipid content (PAHs-LC) coincided well with the number of haze-occurred days and were partly similar with the historical changes in airborne PAHs recorded in the sedimentary cores. The results indicated that PAHs-LC in the xylem of conifers might reliably reflect the historical changes in airborne PAHs at a regional scale. The species-specificity should be addressed in the utility and application of dendrochemical monitoring on historical and comparative studies of airborne PAHs.

Evidence for Air-Seeding: Watching the Formation of Embolism in Conifer Xylem.

Water transport in plants is based on a metastable system as the xylem "works" at negative water potentials (ψ). At critically low ψ, water columns can break and cause embolism. According to the air-seeding hypothesis, this occurs by air entry via the pits. We studied the formation of embolism in dehydrating xylem sections of Juniperus virginiana (Cupressaceae), which were monitored microscopically and via ultrasonic emission analyses. After replacement of water by air in outer tracheid layers, a complex movement of air-water menisci into tracheids was found. With decreasing ψ, pits started to aspirate and the speed of menisci movements increased. In one experiment, an airseeding event could be detected at a pit. The onset of ultrasonic activity was observed when pits started to close, and ultrasonic emission ceased at intense dehydration. Experiments clearly indicated that predictions of the air-seeding hypothesis are correct: At low ψ, pit mechanisms to prevent air entry failed and air spread into tracheids. ψ fluctuations caused complex movements of air-water menisci and pits, and at low ψ, air-seeding caused ultrasonic emissions. Main insights are presented in a video.

Cell wall ultrastructure of palm leaf fibers

The cell wall organization of leaf sheath fibers in different palm species was studied with polarized light microscopy (PLM) and transmission electron microscopy (TEM). The secondary wall of the fibers consisted of only two layers, S1 and S2. The thickness of the S1 layer in leaf sheath fibers from the different palm species ranged from 0.31 to 0.90 μm, with a mean value of 0.57 μm, which was thicker than that of tracheids and fibers in secondary xylem of conifers and dicotyledons. The thickness of the S2 layer ranged from 0.44 to 3.43 μm, with a mean value of 1.86 μm. The ratio of S1 thickness to the whole cell wall thickness in palm fibers appears to be higher than in secondary xylem fibers and tracheids. The lignin in the fiber walls is very electron dense which makes it difficult to obtain high contrast of the different layers in the secondary wall. To clarify the cell wall layering with cellulose microfibrils in different orientations, the fibrovascular bundles of the windmill palm (Trachycarpus fortunei) were delignified with different reaction time intervals. The treated fibers were surveyed using attenuated total reflection Fourier transform infrared (ATR-FTIR) spectroscopy analysis and TEM. The secondary fiber walls of windmill palm clearly showed only two layers at different reaction intervals with different lignin contents, even after almost all lignin was removed. We suggest that the two-layered structure in the secondary wall of palm leaf fibers, which presumably also applies to the homologous fibers in palm stems, is a specific character different from the fibers in other monocotyledons (such as bamboo and rattan) and dicot wood.

Structure-function constraints of tracheid-based xylem: a comparison of conifers and ferns

The ferns comprise one of the most ancient tracheophytic plant lineages, and occupy habitats ranging from tundra to deserts and the equatorial tropics. Like their nearest relatives the conifers, modern ferns possess tracheid-based xylem but the structure-function relationships of fern xylem are poorly understood. Here, we sampled the fronds (megaphylls) of 16 species across the fern phylogeny, and examined the relationships among hydraulic transport, drought-induced cavitation resistance, the xylem anatomy of the stipe, and the gas-exchange response of the pinnae. For comparison, the results are presented alongside a similar suite of conifer data. Fern xylem is as resistant to cavitation as conifer xylem, but exhibits none of the hydraulic or structural trade-offs associated with resistance to cavitation. On a conduit diameter basis, fern xylem can exhibit greater hydraulic efficiency than conifer and angiosperm xylem. In ferns, wide and long tracheids compensate in part for the lack of secondary xylem and allow ferns to exhibit transport rates on a par with those of conifers. We suspect that it is the arrangement of the primary xylem, in addition to the intrinsic traits of the conduits themselves, that may help explain the broad range of cavitation resistance in ferns.

Ultrasonic emissions from conifer xylem exposed to repeated freezing

Ultrasonic emission measurements enable the analysis of xylem cavitation induced by drought and freeze–thaw events. Several studies have indicated that ultrasonic acoustic emissions (UAE) in conifers occur upon freezing and not upon thawing, although classical theory has postulated gas bubble formation during freezing and cavitation during thawing. We analyzed the pattern and quality of freeze–thaw-induced UAE in seven conifers ( Abies alba, Larix decidua, Juniperus communis, Picea abies, Pinus cembra, Pinus mugo, Pinus sylvestris). Axes samples dehydrated to different water potentials were exposed to repeated frost cycles. The number, amplitude and energy of UAE signals were registered and related to water potential, temperature course and wood characteristics (wood density, tracheid diameter). For P. abies, ultrasonic emission analysis was also performed on bark samples, xylem samples without bark, as well as on stems of young potted trees. In all conifers, UAE were registered in water-stressed samples but not in saturated or dehydrated samples. No signals were emitted by the bark of P. abies. Ultrasonic activity occurred only upon freezing, and identical patterns were observed in axes samples and stems of potted P. abies trees. A weak positive relationship between tracheid diameter and UAE energy was observed, indicating wide tracheids to emit signals with higher energy. The classical bubble formation hypothesis cannot sufficiently explain the occurrence of UAE during freezing and upon repeated temperature cycles, as demonstrated in this study. We suggest that the low water potential of ice induces air-seeding near the ice-water interface, and consequently, causes UAE.

Winter at the alpine timberline causes complex within‐tree patterns of water potential and embolism in Picea abies

Winter temperatures at the alpine timberline cause ice formation in the xylem of conifers blocking water uptake as well as water shifts within the axes system. This amplifies drought stress that, in combination with freeze-thaw events, causes embolism. This study focussed on within-tree patterns of water potential (psi) and embolism in Norway spruce [Picea abies (L.) Karst.]. At five sampling dates in midwinter, psi was determined at numerous positions in the crown of three trees, and at the end of March, the extent of embolism in representative sections of the axes system was analysed. Until 14 March, mean psi decreased to -3.77 +/- 0.11 MPa with less negative psi in exposed crown parts. On 30 March, psi was -1.60 +/- 0.06 MPa, while loss of conductivity reached up to 100%. Conductivity losses increased with exposition and were highest in the smallest tree. The observed complex within-tree patterns of psi and embolism were caused by ice blockages and differences in stress intensities within the xylem. High conductivity losses despite moderate psi in exposed crown parts indicated freeze-thaw events to be a major inducer of winter embolism. Tree size may play a critical role for winter water relations as trees profit from water stored in the stem and in crown parts below the snow cover.

Tensile and compressive stresses in tracheids are induced by swelling based on geometrical constraints of the wood cell

Plants are able to pre-stress their tissues in order to actuate their organs. Here, we demonstrate with two tissue types of the secondary xylem of conifers (normal wood and compression wood of spruce (Picea abies)) that either tensile or compressive stresses can develop in the longitudinal direction during the swelling of the cell wall. This dramatic difference appears to be due mostly to differences in cell geometry and cellulose fibril orientation. A mechanical model was developed to demonstrate swelling experiments with the help of sodium iodide experiments. The reversal of longitudinal extension can be predicted, based on the orientation of the (nearly inextensible) cellulose fibrils and the shape of the cell.

Mechanical reinforcement of tracheids compromises the hydraulic efficiency of conifer xylem

Wood structure and function of juvenile wood from 18 conifer species from four conifer families (Araucariaceae, Cupressaceae, Pinaceae and Podocarpaceae) were examined for a trade-off between wood reinforcement and hydraulic efficiency. Wood density and tracheid 'thickness-to-span' ratio were used as anatomical proxies for mechanical properties. The thickness:span represented the ratio of tracheid double wall thickness to lumen diameter. Hydraulic resistivity (R) of tracheids on a cross-sectional area basis (RCA) increased over 50-fold with increasing density and thickness:span, implying a strength versus efficiency conflict. The conflict arose because density and thickness:span were increased by narrowing tracheid diameter rather than by thickening walls, which may be developmentally difficult. In the Pinaceae and Cupressaceae species, density and thickness:span correlated strongly with protection from drought-induced embolism, suggesting that mechanical strength was required in part to withstand tracheid collapse by negative sap pressure. These species showed a corresponding trade-off between increasing RCA and embolism protection. In contrast, species of Podocarpaceae and Araucariaceae were overbuilt for their embolism protection and were hydraulically inefficient, having greater density, thickness:span and RCA, none of which were correlated with vulnerability to embolism.

No Correlation between Latewood Formation and Leader Growth in Douglas-Fir Saplings

The width of earlywood and latewood in conifer xylem may have a profound effect on water transport and storage, vulnerability to embolism, and wood strength, yet the controls over the timing of latewood formation are unclear. Tracheids differentiating in the cambial zone are influenced by IAA, indole-3 acetic acid, the radial concentration gradient of which appears to either increase cell expansion (earlywood) or increase cell wall deposition (latewood). There are suggestive data that latewood begins to form when the growth of the leader stops, but definitive results are lacking. Height growth was measured in 14 Douglas-fir (Pseudotsuga menziesii) saplings at 10 dates between May and August, from the beginning of the growing season until after height growth had ceased. The cambium was also pinned six times between June and July, to induce xylem scarring at known dates. After height growth ceased, saplings were harvested and transverse sections of the wood were made at the pin insertion points. The date at which 95% of the height growth had occurred and the date at which latewood formation had begun were estimated. Analysis showed no correlation of these data, suggesting that the two phenomena may occur around the same time, but that one is not causal of the other.