The fiber bundles from the lignified leaf sheath of windmill palm (Trachycarpus fortunei) are widely used as natural fibers for various products, and exhibit excellent durability. In this study, the cell wall of windmill palm fibers was characterized using transmission electron microscopy, high resolution field emission scanning electron microscopy, and polarized light microscopy, and chemical analysis to measure lignin content. It was found that (1) the secondary wall was composed of just two layers, outer (equivalent to S1, 0.65 ± 0.12 μm) and inner (equivalent to S2, 1.28 ± 0.30 μm) ones, with a high ratio of S1 to the whole cell wall thickness; (2) the microfibrils of the S1 are orientated in an S-helix (MFA, 127.0° ± 2.0), and those of the S2 in a Z-helix (MFA, 43.7° ± 2.2); and (3) the Klason lignin content of fiber bundles was very high (nearly 40%). It is suggested that these structural and chemical features of windmill palm fibers are involved in their mechanical properties such as high flexibility and elasticity, and also related to their high durability.
This study presents anatomical characteristics, mechanical properties, microfibril angles (MFAs) and Klason lignin contents of leaf-sheath fibrovascular bundles from 14 palm genera (18 species). Observed by light microscopy, all fibrovascular bundles consisted equally of thick-walled sclerenchyma fibers and vascular tissue, while the shape and localization of vascular tissues on the transverse sections varied among species. It was possible to group these fibrovascular bundles into 3 types based on the vascular tissue’s differences: type A – rounded in the central region; type B – angular in the marginal region; and type C – aliform in the central region. These three anatomical types of fibrovascular bundles showed some correlation with a current phylogenetic classification of palm species. Through mechanical tests, this research confirmed the correlation between diameter and mechanical properties of the fibrovascular bundles of palms; tensile strength and Young’s modulus showed a decreasing trend with increasing diameter. We clarified that this trend was due to a marked increase in the proportion of transverse sectional area comprised by vascular tissue with increasing diameter of fibrovascular bundles. The MFAs of fibrovascular bundles ranged from 10.3º to 47.1º, which were generally larger than those of non-woody plants, conifers, and broad-leaved trees. The Klason lignin contents of palm species were also high, ranging from 18.3% to 37.8%, with a mean value of 29.6%. These large MFAs and high lignin contents could lead to the long-term plastic deformation and relatively low tensile strength of palm fibrovascular bundles.
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.
Agarwoods such as Aquilaria spp. and Gyrinops spp. (Thymelaeaceae) produce interxylary phloem in their secondary xylem and intraxylary phloem at the periphery of the pith, facing the primary xylem. We studied young shoots of Aquilaria sinensis and characterized the development of its intraxylary phloem. It was initiated by the division of parenchyma cells localized in the outer parts of the ground meristem immediately following the maturation of first-formed primary xylem. Its nascent sieve plates bore donut-like structures, the individual pores of which were so small (less than 0.1 μm) that they were hardly visible under FE-SEM. Intraxylary phloem developed into mature tissue by means of the division and proliferation of parenchyma cells. During the shoots’ active growth period, the sieve pore sizes were 0.1–0.5 μm, with tubular elements passing through them. In the maturation stage, large clusters of sieve tubes continued to be differentiated in the intraxylary phloem. In the partial senescence stage observed in a three-centimeter-diameter branch, intraxylary phloem cells in the adaxial part became crushed, and sieve plates had pores over 1–2 μm in diameter without any callose deposition. Before and after the differentiation of interxylary phloem in the first and second internodes, callose staining detected more than twice as many sieve tubes in intraxylary phloem than in external phloem. However, after differentiation of interxylary phloem in the eleventh internode, more sieve tubes were found in interxylary phloem than in intraxylary and external phloem. This suggests that prior to the initiation of interxylary phloem intraxylary phloem acts as the principal phloem. After its differentiation, however, interxylary phloem takes over the role of principal phloem. Interxylary phloem thus acts as the predominant phloem in the translocation of photosynthates in Aquilaria sinensis.
Variation in fiber diameter and wall thickness was analyzed, with respect to the distance from xylem vessels in tangential and radial directions, using images containing the largest diameter along individual fibers from serial sections. For the diameter of the fibers around the vessel, it was often difficult to make a measurement in the radial and tangential direction, because of deformation. Then, around a vessel, we measured the fiber lumen diameter along the radial and tangential axes of the adjacent vessel. These are respectively referred as direction perpendicular to vessel enlargement (PnVE) and parallel with vessel enlargement (PrVE), as the enlarging vessel apparently accounts for the directionality of this fiber deformation. The change in cell wall thickness of individual fibers was also measured from the serial cross sections. Fibers adjacent to vessels were significantly wider in diameter and had thicker cell walls in the PrVE than those that were more distant from vessels in both radial and tangential directions. Wall thickness along the fiber length was found to vary in both fibers adjacent to and distant from the vessel, whereby wall thickness was greatest at the center, and gradually decreased towards the tip of the fiber.
New observations of radial sieve tubes in the secondary xylem of two genera and four species of agarwood — Aquilaria sinensis, A. crasna, A. malaccensis and Gyrinops versteeghii (Thymelaeaceae) — are presented in this study. The earliest radial sieve tubes in Gyrinops are formed in the secondary xylem adjacent to the pith. The radial sieve tubes originate from the vascular cambium and develop in both uniseriate and multiseriate ray tissue. In addition to sieve plates in lateral and end walls, scattered or clustered minute sieve pores are localized in the lateral wall of radial sieve tubes. There is a direct connection between radial sieve tubes in ray tissue and axial sieve tubes in interxylary phloem strands (IP), such as (i) connection by bending of radial sieve tube strands, (ii) connection of two IP strands by an oblique bridge, and (iii) connection of two IP strands at a right angle. The average number of radial sieve tubes and interxylary phloem was found to be 1.7 per mm3 and 9.1 per mm2 in the secondary xylem. Considering the higher frequency of radial sieve tubes with the increasing thickness of the secondary xylem, the direct connections between radial and axial sieve tubes could play a significant role in assisting the translocation of metabolites in Aquilaria and Gyrinops.