Tracheid Walls Research Articles

Bacterial decay in waterlogged archaeological compression wood varies with severity of compression wood

Abstract Bacterial decay in compression wood (CW) tracheids of waterlogged archaeological wood (WAW) was investigated using light microscopy, confocal laser scanning microscopy, transmission electron microscopy (TEM), and TEM immunogold labeling. Erosion bacteria were identified as the main degraders, and the extent of cell wall degradation differed depending on the severity of CW tracheids (mild vs. severe). Mild CW tracheids showed preferential decay in the inner S2 layer, with the locally degraded and/or fragmented S3 layer remaining. In contrast, severe CW tracheids revealed gradual degradation of the cell wall from the erosion progressing from exposed faces of the cell wall as decay progressed. The overall decay was more extensive in mild than in severe CW tracheids, and degradation of the highly lignified outer S2 layer (S2L) was only detected in mild CW tracheids. TEM immunogold labeling of 1,4-β-galactan, homogalacturonan (HG), heteroxylan, and heteromannan epitopes showed that there was no preferential degradation of pectins and hemicelluloses by action of diffusible enzymes and/or agents through the un-decayed cell wall during bacterial decay, in both mild and severe CW tracheids. Inter-CW tracheid bordered pit membranes showed higher decay resistance than CW tracheid walls. Degradation of HG and heteromannan epitopes was suppressed in pit membranes.

Metal-organic frameworks-derived carbon modified wood carbon monoliths as three-dimensional self-supported electrodes with boosted electrochemical energy storage performance

Wood-derived carbon monoliths, in recent years, have attracted tremendous interest in the field of energy storage, but their electrochemical characteristics are still far from satisfactory. Here, we report a universal and efficient approach for the preparation of structure-engineered, heteroatom-functionalized and property-boosted wood carbons. A two-step ion-exchange process greatly enriches the nucleation sites of ZIF-8 on the inner wall of wood tracheids, hence leading to a unique carbon/carbon heterostructure after carbonizing and acid-washing. Particularly, the prepared NPCM-900 with a large specific surface area of 708.2 m2 g−1, a hierarchical porous architecture and a suitable N content of 2.3% delivers an ultrahigh area-normalized specific capacitance of 23.7 F cm−2 at 5 mA cm−2, which stands for a new capacitive record among the wood-based binder-free electrodes. The NPCM-900//NPCM-900 all-solid-state supercapacitor has an admirable energy density of 9.3 Wh m−2 at 24.9 W m−2 and a large power density of 248.3 W m−2 at 4.8 Wh m−2, while the NPCM-900 based Zn-ion hybrid supercapacitor (NPCM-900//Zn) exhibits a superior energy density of 12.7 Wh m−2. Furthermore, the NPCM-900//NPCM-900 and NPCM-900//Zn present great stabilities with capacitance retentions of 87% and 85%, respectively, after 5000 cycles. These parameters notably outperform those of most wood-based supercapacitors, endowing the NPCM-900 with extensive prospects for practical use.

Effects of the combination of compression and impregnation with phenolic resin on the dimensional stability in the multiscale wood structure of Chinese fir

Chinese fir wood was pretreated by phenol–formaldehyde (PF) resin and followed by the compression along the radial direction at the compression ratio of 10%, 20% and 30%, respectively. The microscopic images of the control and modified wood at different humidity were then collected to evaluate the dimensional stability of wood cellular structures. The interactions between the cell wall and resin were also observed by SEM and Raman spectroscopy to reveal the fixation effects of resin on the compressed wood at cell-level. Results indicated that the number of wood cells filled with PF resin increased significantly with the increase of impregnation pressure and processing time, which contributed to an obvious decrease in the linear and volumetric expansion ratio of wood at high humidity, in particular at the RH of 90%. The filling of PF resin in the wood cell lumen and the open pits on the tracheid walls not only inhibited the entry of water molecules but also had a chemical interaction with cell-wall, which could significantly reduce the set-recovery of the compressed wood. The radial expansion ratio of wood at a compression ratio of 30% has been reduced by about 65% at the RH of 90% after PF resin impregnation.

New Evidence for Palaeo-wildfire in the Early Permian (Artinskian) of Gondwana from Wardha Valley Coalfield, India

Abstract Macroscopic fossil charcoal fragments have been recovered from the borehole AK-19 (at a depth of 120.00 m) drilled at the Astona-Kothurna coal block, Wardha valley coalfield. In general, the occurrence of charcoal in sediments is accepted as a direct indicator for palaeo-wildfires. The charcoal fragments exhibits well preserved homogenized cell walls as well as anatomical details such as uniseriate and biseriate pitting patterns on tracheid walls under Field Emission Scanning Electronic Microscope (FESEM), suggesting a gymnospermous wood affinity of the studied material. The excellent preservation, their large size and almost unabraded edges of charcoal fragments suggest a par-autochthonous origin. These findings add further evidence for the widespread occurrence of such wildfires during the Early Permian not only in India, but on the entire Gondwana continent.

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.

Two new petrified gymnosperms with solenoid piths from the Pedra de Fogo Formation, Permian of Maranhão, Brazil

Solenoid piths are formed by systems of secretory ducts or canals, and they are relatively common in petrified gymnosperms from the Permian of Gondwana. We described herein two new taxa, Polycanaloxylon merlottii gen. et sp. nov. and Europoxylon garapensis sp. nov., from the Pedra de Fogo Formation (Cisuralian), northeastern Brazil. In both species the pith is non-septate and solenoid, primary xylem bundles have endarch maturation, rays are short and uniseriate, pits on the radial walls of tracheids are commonly uni to biseriate, cross-field pitting is cupressoid and wood is pycnoxylic. Whereas P. merlottii has a solenoid pith with a parenchyma sheath and sclerenchyma cells, E. garapensis is distinguished by a pith that only has secretory canals and parenchyma cells. In addition to expanding the distribution area of a typical Euramerican taxon into Parnaíba Basin, the report of Europoxylon reinforces a connection between these areas in the Late Pennsylvanian, thus enabling the dispersion of floristic elements. Solenoid forms are especially common in Kungurian sedimentary deposits of southern Brazil, and possibly India. However, since the occurrence of them seems to be linked to specific habitats, it should be proven that such environments are indeed contemporaneous in order to use these stems as a tool for relative dating in Gondwanan localities.

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.

Megaporoxylon sinensis sp. nov., a new coniferous trunk from the Upper Triassic of northern Bogda Mountains, northwestern China

A coniferous trunk, Megaporoxylon sinensis sp. nov., is described from the Carnian–Norian (Triassic) Huangshanjie Formation in the Dalongkou section, Jimsar County, Xinjiang, northwestern China. The trunk is composed of pith, primary and secondary xylem. The pith is solid, circular, heterocellular, with pitted parenchyma and secretory cells and canals. The primary xylem is endarch, with spiral and scalariform thickenings on tracheidal walls. The secondary xylem is pycnoxylic, consisting of tracheids and rays. Radial tracheidal pitting is araucarian. Tangential pitting and axial parenchyma are absent. Uniseriate pits with oval apertures are commonly present on radial tracheid walls. Biseriate pitting occurs only on radial walls of the first to fifth tracheids outward the primary xylem. Rays are parenchymatous, uniseriate, and 1–7 cells high. Cross-field pitting is window-like or phyllocladoid. Previous records of species of Megaporoxylon Kräusel were limited to the Southern Hemisphere during the Carboniferous to Triassic. Its occurrence from the Upper Triassic of northern Xinjiang is the first report in the Northern Hemisphere and suggests a floristic exchange between Gondwana and Laurasia. The new trunk and two previously described fossil stems, including Medulloprotaxodioxylon triassicum Wan et al. and Xenoxylon junggarensis Wan et al., demonstrate that conifers were important elements of Carnian–Norian terrestrial ecosystems and had a higher diversity in northern China than previous thought. The growth ring pattern of the trunk and palaeogeographic reconstruction of the research area at high latitudes suggest that the growth of the trees was limited by the seasonality of the light regime.

Nickel-cobalt layered double hydroxide nanosheets anchored to the inner wall of wood carbon tracheids by nitrogen-doped atoms for high-performance supercapacitors

In this paper, a novel type of electrode material for high-performance hybrid supercapacitors is designed. The electrode mainly uses nitrogen-doped atoms to anchor the nickel–cobalt layered double hydroxide on the inner wall of wood-derived carbon tracheids from Chinese fir wood scraps. The specific capacity of the composite single electrode is 14.26 mAh cm−2 at 10 mA cm−2. The hybrid supercapacitor with a composite electrode cathode and nitrogen-doped wood-derived monolithic carbon materials as the anode has a high specific capacitance of 4.74F cm−2 at 5 mA cm−2, and the capacitance retention rate is 93.15% after 8000 charge–discharge cycles. The highest energy density and power density reach 1.48 mWh cm−2 and 22.40 mW cm−2, respectively. After doping with nitrogen, the combination with the nickel–cobalt layered double hydroxide is more uniform and stable, and the capacitance and cycling stability are significantly improved.

Bordered Pit Formation in Cell Walls of Spruce Tracheids.

The process of pit formation in plants still has various questions unaddressed and unknown, which opens up many interesting and new research opportunities. The aim of this work was elucidation of the mechanism for the formation of bordered pits of the spruce (Picea abies (L.) Karst.) tracheid with exosomes participation and mechanical deformation of the cell wall. Sample sections were prepared from spruce stem samples after cryomechanical destruction with liquid nitrogen. The study methods included scanning electron microscopy and enzymatic treatment. Enzymatic treatment of the elements of the bordered pit made it possible to clarify the localization of cellulose and pectin. SEM images of intermediate stages of bordered pit formation in the radial and tangential directions were obtained. An asynchronous mechanism of formation of bordered-pit pairs in tracheids is proposed. The formation of the pit pair begins from the side of the initiator cell and is associated with enzymatic hydrolysis of the secondary cell wall and subsequent mechanical deformation of the primary cell walls. Enzymatic hydrolysis of the S1 layer of the secondary cell wall is carried out by exosome-delivered endoglucanases.

New petrified calamitaleans from the Permian of the Parnaíba Basin, central-north Brazil, part III, with some concerns regarding anatomical features of Paleozoic petrified sphenophytes

A new petrified calamitalean is described from the Permian Motuca Formation, Parnaíba Basin, central-north Brazil. In contrast to all other three-dimensionally preserved Brazilian species, it represents a semi-self-supporting woody plant characterized by a distinctive branch system and cellular anatomy. Arthropitys buritiranensis sp. nov. is proposed to accommodate the new material. The branch architecture is composed of at least three types of branches: (i) principal woody side branches, (ii) up to three secondary woody branches per node, scattered along the stem or the principal side branches, and (iii) 8–11 leafy twigs per whorl, at every third, rarely the second node, without secondary growth. The wood is segmented into fascicular wedges and interfascicular rays, clearly visible only within the inner wood of the stem and woody branches. The tracheid walls of the secondary xylem have exclusively scalariform pits. Ray parenchyma cells have pitted horizontal walls. The new calamitalean was part of the riparian vegetation of tree ferns, calamitaleans and gymnosperms recorded in a distal fluvial plain setting. The wood shows irregular growth interruptions and, therefore, points to a seasonally dry climate that shaped even low latitude paleofloras of tropical Pangaea.

BIOSOCIAL DIVERSITY OF SCOTS PINE (PINUS SYLVESTRIS L.) IN A TREE STAND IN RELATION TO CHOSEN HYDRAULIC CONDUCTIVITY INDICATORS OF THE STEM

The research focused in determining the lignification indicator of fresh needled springs and the mass of fresh needles in reference to the lignin content in tracheid walls of peripheral area of the stem (MFT/LC and MFN/LC) of Scots pine differentiated as far as its biosocial position within the community expressed by Kraft’s classification. The material for the analysis came from mature pine stands growing on North European Plain, on the territory of Poland. Chemical and structural analyses of wood encompassed the area of mature sapwood, i.e. thickness of the last 10 annual rings located at 1.3 m (DBH). It seems that the noticed differences values of both indicators (MFT/LC and MFN/LC) in pines belonging to the first three Kraft’s biological classes are connected with physiological, physical and structural conditionings of water transport with minerals in xylem and are closely connected with competition for sunlight, water, nutrients and living space.

A new Protophyllocladoxylon stem from the Xishanyao Formation (Middle Jurassic) in the Santanghu Basin, Xinjiang, Northwest China

A new permineralized stem, Protophyllocladoxylon yiwuense Gou et Feng sp. nov., is described from the Middle Jurassic Xishanyao Formation in Naomaohu Town of Yiwu County, Hami City, Xinjiang Uygur Autonomous Region, Northwest China. The stem is only preserved with secondary xylem, which is pycnoxylic and exclusively composed of tracheids and parenchymatous rays. The tracheids have uniseriate or biseriate, alternately arranged bordered pits on their radial walls, and uniseriate bordered pits on the tangential walls. The rays are uniseriate or partially biseriate, up to 34 cells high. There are one or two large, simple pits in each cross field. A branch trace horizontally passes through the secondary xylem showing anatomical structures identical to the secondary xylem. Analyses of the growth-ring width and mean sensitivity values indicate that P. yiwuense had long growing seasons with readily available water supplies and favorable paleoclimatic conditions. White-rot decay features are commonly recognized in the stem, including pocket-like cavities, complete removal of the middle lamellae, thickened corners and separation of the secondary walls of tracheids, as well as septa-like structures in the tracheid lumens due to the cell's reactions to fungal activity. Our study sheds new light into the plant diversity and paleoenvironmental and paleoecological conditions of the Middle Jurassic.

Mapping variation of handsheet properties within loblolly pine trees

Abstract Within-tree variation of four handsheet properties (burst index, sheet density, STFI short-span compression strength (STFI) and tensile index) was mapped for loblolly pine trees aged 13 and 22 years using NIR predicted handsheet property data (representing an average of 18 trees for each age). All within-tree maps were similar demonstrating a radial decrease in handsheet properties at all heights, with sheet density and tensile index having the greatest within-tree variation. The corewood zone had the highest values for all properties, while the lowest values were observed in a region consistent with juvenile and transitional outerwood as defined by Burdon et al. (2004). The maps are also similar to, but the inverse of, maps reported in prior studies for density and tracheid coarseness and wall thickness. Relationships amongst the examined handsheet properties and wood and tracheid properties explains the overall similarities of the different maps. The maps provide a representation of within-tree variation of important paper properties at two different ages. An understanding of how these properties vary within loblolly pine trees can aid in better utilization of forest resources.

Zhuotingoxylon liaoi gen. et sp. nov., a silicified coniferous trunk from the Changhsingian (Permian) of southern Bogda Mountains, northwestern China

A silicified trunk, Zhuotingoxylon liaoi Wan, Yang, Wang, Liu et Wang gen. et sp. nov., is described from the uppermost part of Guodikeng Formation in South Taodonggou section, Turpan–Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. It is characterized by a solid pith, endarch primary xylem and pycnoxylic wood. The pith is composed of parenchyma and sclereids. Radial walls of primary xylem tracheids have spiral and scalariform thickenings. Secondary xylem consists of thick‐walled tracheids and parenchymatous rays. Uniseriate rounded pits with oval apertures are distributed on radial tracheidal walls separately. Cell walls of rays are homogeneous and smooth. Rays are 1–10 cells high in tangential section. Cross‐field pits are cupressoid. There are 1–4 bordered pits with slit‐like to oval apertures in each cross‐field. Based on the anatomical features of the pith and xylems, it is proposed that the new stem has a coniferous affinity. The new fossil stem adds to the knowledge of vascular plant diversity close to the Permian–Triassic boundary.

Localised laccase activity modulates distribution of lignin polymers in gymnosperm compression wood.

Summary The woody stems of coniferous gymnosperms produce specialised compression wood to adjust the stem growth orientation in response to gravitropic stimulation. During this process, tracheids develop a compression‐wood‐specific S2L cell wall layer with lignins highly enriched with p‐hydroxyphenyl (H)‐type units derived from H‐type monolignol, whereas lignins produced in the cell walls of normal wood tracheids are exclusively composed of guaiacyl (G)‐type units from G‐type monolignol with a trace amount of H‐type units. We show that laccases, a class of lignin polymerisation enzymes, play a crucial role in the spatially organised polymerisation of H‐type and G‐type monolignols during compression wood formation in Japanese cypress (Chamaecyparis obtusa).We performed a series of chemical‐probe‐aided imaging analysis on C. obtusa compression wood cell walls, together with gene expression, protein localisation and enzymatic assays of C. obtusa laccases.Our data indicated that CoLac1 and CoLac3 with differential oxidation activities towards H‐type and G‐type monolignols were precisely localised to distinct cell wall layers in which H‐type and G‐type lignin units were preferentially produced during the development of compression wood tracheids.We propose that, not only the spatial localisation of laccases, but also their biochemical characteristics dictate the spatial patterning of lignin polymerisation in gymnosperm compression wood.

Palaeobotanical and biomarker evidence for Early Permian (Artinskian) wildfire in the Rajmahal Basin, India

This study provides a combined analysis on the palynology, fossil charcoal and biomarkers of the subsurface coal deposits from a borehole RMB #2 drilled at the Dhulia Coal Block, Rajmahal Basin, India, in attempts to establish the chronology of sedimentation and to propose palaeobotanical as well as geochemical evidence for the occurrence of wildfires in these sediments. The palynological investigation suggests a Scheuringipollenites barakarensis palynoassemblage from the lower Barakar Formation, dated as Artinskian (Early Permian) in age. This assemblage reveals the dominance of Glossopteridales and sub-dominance of taxa belonging to Cordaitales and Coniferales. Fossil charcoal in sediments is usually recognized as a direct indicator for the occurrence of palaeo-wildfires. More data involving the anatomical features of fossil charcoal analyzed by Field Emission Scanning Electronic Microscope broaden our knowledge on Early Permian wildfires from the peninsula of India. The studied macroscopic charcoal fragments exhibit anatomical details such as homogenized cell walls, uniseriate simple and biseriate alternate pitting on tracheid walls and rays of varying heights pointing to a gymnospermous wood affinitity. The excellent preservation of charcoal fragments, shown by their large sizes and almost unabraded edges, suggests a parautochthonous origin. The embedded biomarker study performed for charcoal sediments and its characterization demonstrate the presence of n-alkanes, isoprenoids, terpenoids and aromatic compounds. A bimodal distribution pattern of n-alkanes with a Cmax at n-C25 is identified. Diterpenoids and pentacyclic terpenoids are identified, indicating the input of an early conifer vegetation and bacterial activity, respectively. The identified polyaromatic hydrocarbon (PAH) compounds, together with the charcoal fragments, clearly suggest that repeated wildfire events occurred during the deposition of these Artinskian sediments in the Rajmahal Basin.

Effects of Intra-Seasonal Drought on Kinetics of Tracheid Differentiation and Seasonal Growth Dynamics of Norway Spruce along an Elevational Gradient

Research Highlights: Our results provide novel perspectives on the effectiveness and collapse of compensatory mechanisms of tracheid development of Norway spruce during intra-seasonal drought and the environmental control of intra-annual density fluctuations. Background and Objectives: This study aimed to compare and integrate complementary methods for investigating intra-annual wood formation dynamics to gain a better understanding of the endogenous and environmental control of tree-ring development and the impact of anticipated climatic changes on forest growth and productivity. Materials and Methods: We performed an integrated analysis of xylogenesis observations, quantitative wood anatomy, and point-dendrometer measurements of Norway spruce (Picea abies (L.) Karst.) trees growing along an elevational gradient in South-western Germany during a growing season with an anomalous dry June followed by an extraordinary humid July. Results: Strong endogenous control of tree-ring formation was suggested at the highest elevation where the decreasing rates of tracheid enlargement and wall thickening during drought were effectively compensated by increased cell differentiation duration. A shift to environmental control of tree-ring formation during drought was indicated at the lowest elevation, where we detected absence of compensatory mechanisms, eventually stimulating the formation of an intra-annual density fluctuation. Transient drought stress in June also led to bimodal patterns and decreasing daily rates of stem radial displacement, radial xylem growth, and woody biomass production. Comparing xylogenesis data with dendrometer measurements showed ambivalent results and it appears that, with decreasing daily rates of radial xylem growth, the signal-to-noise ratio in dendrometer time series between growth and fluctuations of tree water status becomes increasingly detrimental. Conclusions: Our study provides new perspectives into the complex interplay between rates and durations of tracheid development during dry-wet cycles, and, thereby, contributes to an improved and mechanistic understanding of the environmental control of wood formation processes, leading to the formation of intra-annual density fluctuations in tree-rings of Norway spruce.

Flexural anisotropy of rift-sawn softwood boards induced by the end-grain orientation

Inspired by the use of rift-sawn softwood board for covering curved surfaces in Kokerabuki, a traditional Japanese roofing method, we investigated the flexural anisotropy of wood caused by its end-grain orientation. We measured the flexural displacement of softwood species, Chamaecyparis obtusa and Cryptomeria japonica, and hardwood species, Populus suaveolens and Cerasus serrulata. For the softwood species, this was approximately five times longer for the rift-sawn specimens than for the other grain patterns. Using the replica method to measure the softwood tracheid deformation with different flexural displacements, we confirmed the different deformation mechanisms of the tracheid in the flat- and quarter-sawn specimens, and rift-sawn specimens. In the flat- and quarter-sawn woods, on-axis loading was generated, in which the stress was concentrated on the radial and tangential cell walls parallel to the direction of tension or compression. By contrast, in the rift-sawn wood, off-axis loading was generated, in which the stress was evenly distributed throughout the corner cell walls without the wall directly resisting the tensile and compressive forces. We also concluded that the tapered shape of the tracheid walls contributes to the excellent flexibility of rift-sawn softwood.

A new conifer stem, Ductoagathoxylon wangii from the Middle Jurassic of the Santanghu Basin, Xinjiang, Northwest China

A new permineralized stem, Ductoagathoxylon wangii Gou et Feng, is described from the Middle Jurassic Xishanyao Formation in Naomaohu Town of Yiwu County, Hami City, Xinjiang Uygur Autonomous Region, Northwest China. The fossil stem is preserved with pith, primary xylem, and secondary xylem. The solid pith is heterocellular, and characterized by the presence of regularly arranged clusters of secretory cells at the periphery. The primary xylem is endarch to mesarch, showing annular, helical, and scalariform thickenings on the tracheid walls from the protoxylem to the metaxylem. The secondary xylem is pycnoxylic and exclusively composed of tracheids and parenchymatous rays. The secondary xylem tracheids have mostly uniseriate, partially biseriate, and rarely triseriate alternately arranged contiguous bordered pits on their radial walls. The rays are uniseriate and up to ten cells high. There are five to nine cupressoid pits in each cross-field, arranged in the araucarioid type. Growth rings are distinct and are characterized by a narrow band of latewood. This is the third fossil stem described from the Middle Jurassic of China that is preserved with anatomical structures of pith, primary, and secondary xylem, thus shedding new light on the diversity and evolution of conifers. The occurrence of a traumatic scar in the stem indicates that the tree was probably damaged by an abiotic cause, but it successfully survived through physiological responses to the wound.