Analysis of circular bordered pit function I. Angiosperm vessels with homogenous pit membranes

A model predicted pit and vessel conductivity, the air-seed pressure for cavitation, and the implosion pressure causing vessel collapse. Predictions were based on measurements from 27 angiosperm species with circular bordered pits and air-seed pressures of 0.2-11.3 MPa. Vessel implosion pressure exceeded air-seed pressure by a safety factor of 1.8 achieved by the increase in vessel wall thickness per vessel diameter with air-seed pressure. Intervessel pitting reduced the implosion pressure by 20 to 40%. Pit hydraulic conductivity decreased by 30-fold from low (<1 MPa) to high (>10 MPa) air-seed pressure primarily because of decreasing pit membrane conductivity. Vessel conductivity (per length and wall area) increased with vessel length as higher lumen conductivity overcame low pit conductivity. At the "saturating vessel length," vessel conductivity maximized at the Hagen-Poiseuille value for the lumen per wall area. Saturated vessel conductivity declined by sixfold with increasing air-seed pressure because of increased wall thickness associated with increased implosion resistance. The saturated vessel length is likely the optimal length because: (a) shorter vessels have lower conductivities, (b) longer vessels do not increase conductivity when functional yet decrease it more when cavitated, (c) observed pit structure most closely optimized vessel conductivity at the saturated length, and (d) saturated lengths were similar to measured lengths.     

两种麻黄根与茎次生木质部的解剖与进化和干旱环境的关系

用光镜及扫描电镜对两种麻黄根、茎次生木质部进行了解剖研究,结果表明：轴向系统主要由导管和管胞组成。横向系统由细胞壁木质化了的射线薄壁组织细胞组成。导管直径甚小,多孔式穿孔板,并存在导管与管胞之间的管状分子类型,推断麻黄属是裸子植物中最早出现导管的类群;管胞中有一些两头尖、胞腔小、具缘纹孔含纹孔塞的长分子,可视作纤维状管胞,使管胞的输导作用被削弱,而支持功能被加强;射线异型多列,已不具备裸子植物具较窄射线的特点。导管与管胞并存,恰好起到了一般沙生被子植物具宽窄两种类型的导管、复孔率高等典型的对干旱环境的适应特征的作用,茎中导管分子的长度和宽度均小于根,这与茎部需要较强的机械支持力相一致。     

Bordered pit structure and function determine spatial patterns of air-seeding thresholds in xylem of Douglas-fir (Pseudotsuga menziesii; Pinaceae) trees

The air-seeding hypothesis predicts that xylem embolism resistance is linked directly to bordered pit functioning. We tested this prediction in trunks, roots, and branches at different vertical and radial locations in young and old trees of Pseudotsuga menziesii. Dimensions of bordered pits were measured from light and scanning electron micrographs, and physiological data were from published values. Consistent with observations, calculations showed that earlywood tracheids were more resistant to embolism than latewood tracheids, mainly from earlywood having stretchier pit membranes that can distend and cover the pit aperture. Air seeding that occurs in earlywood appears to happen through gaps between the torus edge and pit border, as shown by the similar calculated pressures required to stretch the membrane over the pit aperture and to cause embolism. Although bordered pit functioning was correlated with tracheid hydraulic diameter, pit pore size and above all pit aperture constrained conductivity the most. From roots to branches and from the trunk base to higher on the trunk, hydraulic resistance of the earlywood pit membrane increased significantly because of a decrease in the size of the pit aperture and size and number of margo pores. Moreover, overall wood conductivity decreased, in part due to lower pit conductivity and a decrease in size and frequency of pits. Structural and functional constraints leading to the trade-off of efficiency against safety of water transport were also demonstrated at the individual pit level, with a positive correlation between pit membrane resistance on an area basis and the pressure differential required to cause membrane stretching, a characteristic that is essential for pit aspiration.     

Xylem of early angiosperms: Nuphar (Nymphaeaceae) has novel tracheid microstructure1

SEM studies of xylem of stems of Nuphar reveal a novel feature, not previously reported for any angiosperm. Pit membranes of tracheid end walls are composed of coarse fibrils, densest on the distal (outside surface, facing the pit of an adjacent cell) surface of the pit membrane of a tracheid, thinner, and disposed at various levels on the lumen side of a pit membrane. The fibrils tend to be randomly oriented on the distal face of the pit membrane; the innermost fibrils facing the lumen take the form of longitudinally oriented strands. Where most abundantly present, the fibrils tend to be disposed in a spongiform, three-dimensional pattern. Pores that interconnect tracheids are present within the fibrillar meshwork. Pit membranes on lateral walls of stem tracheids bear variously diminished versions of this pattern. Pits of root tracheids are unlike those of stems in that the lumen side of pit membranes bears a reticulum revealed on the outer surface of the tracheid after most of the thickness of a pit membrane is shaved away by the sectioning process. No fibrillar texturing is visible on the root tracheid pits when they are viewed from the inside of a tracheid. Tracheid end walls of roots do contain pores of various sizes in pit membranes. These root and stem patterns were seen in six species representing the two sections of Nuphar, plus one intersectional hybrid, as well as in one collection of Nymphaea, included for purposes of comparison. Differences between root and stem tracheids with respect to microstructure are consistent in all species studied. Microstructural patterns reported here for stem tracheid pits of Nymphaeaceae are not like those of Chloranthaceae, Illiciaceae, or other basal angiosperms. They are not referable to any of the patterns reported for early vascular plants. The adaptational nature of the pit membrane structure in these tracheids is not apparent; microstructure of pit membranes in basal angiosperms is more diverse than thought prior to study with SEM.     

Modelling the hydrodynamic resistance of bordered pits

Previous studies of the hydrodynamics of plant stems have shown that resistance to flow through bordered pits on the side walls of tracheids makes up a significant proportion of their total resistance, and that this proportion increases with tracheid diameter. This suggests a possible reason why tracheids with a diameter above around 100 microm have failed to evolve. This possibility has been investigated by obtaining an estimate for the resistance of a single pit, and incorporating it into analytical models of tracheid resistance and wood resistivity. The hydrodynamic resistance of the bordered pits of Tsuga canadensis was investigated using large-scale physical models. The importance of individual components of the pit were investigated by comparing the resistance of models with different pore sizes in their pit membrane, and with or without the torus and border. The estimate for the resistance of a real bordered pit was 1.70x10(15) Pa s m(-3). Resistance of pits varied with morphology as might be predicted; the resistance was inversely proportional to the pore size to the power of 0.715; removing the torus reduced resistance by 28%, while removal of the torus and border together reduced it by 72%. It was estimated that in a 'typical tracheid' pit resistance should account for 29% of the total. Incorporating the results into the model for the resistivity of wood showed that resistivity should fall as tracheid diameter increases. However, to minimize resistance wider tracheids would also need to be proportionally much longer. It is suggested that the diameter of tracheids in conifers is limited by upper limits to cell length or cell volume. This limitation is avoided by angiosperms because they can digest away the ends of their cells to produce long, wide vessels composed of many short cells.     

水青树特殊管胞的分布位置及其形态特征的研究

针对水青树（Tetracentron sinense）中一类特殊管胞进行较为全面的观察研究,判断细胞种类并分析维管组织输导机理及树木进化过程中的细胞演化规律。通过切片和解离技术,借助光学显微镜和电子显微镜对34年生水青树特殊管胞的分布位置和形态特征进行观察。结果表明：(1)特殊管胞在树木水平方向自内向外径向呈串排列,并贯穿年轮界限,多为一列,少数两列,且较为稀见。每个特殊管胞弦向左右两侧或单侧均与木射线细胞相连通。纵向上,特殊管胞单独或数个上下端接相连。(2)特殊管胞主要有以下3种类型：无端壁的纺锤形,有一个倾斜端壁,以及有两个倾斜端壁。特殊管胞的平均长度为286.44μm;横切面为四边形,平均弦向宽度为55.22μm,其平均壁厚为1.53μm。(3)特殊管胞两端封闭,无穿孔。(4)特殊管胞侧面壁上的纹孔数量较多且纹孔膜明显可见,具体表现为：弦面壁上布满特殊管胞之间的具缘纹孔,呈对列、互列偶见梯状排列;径面壁上存在与射线细胞间的具狭缘单纹孔,呈大圆形至椭圆形,每区域多为2～10个纹孔,呈1～4排横列;径面壁上与正常管胞间几无纹孔。水青树特殊管胞分布有一定规律,其长度远小于水青树正常管胞...     

SEM studies on vessels in ferns. 11. Ophioglossum

With scanning electron microscopy (SEM), the nature of metaxylem vessel elements and tracheids was examined in Ophioglossum crotalophomides, 0. pendulum subsp. falcatum , and 0. vulgatum roots and rhizomes. Vessels were identified in all species. End walls of vessel elements, which bear perforations, are like lateral wall pitting of those elements in the secondary wall framework and differ only in absence of pit membranes or presence of pit membrane remnants. Some of the perforations contain pit membrane remnants that have large pores, small porosities, or are threadlike or weblike in structure. Dimorphic perforations were found in some vessel elements of rhizomes of 0. pendulum subsp. falcatum. Tracheids are very likely present in addition to vessels in all three species. The secondary wall framework of both tracheids and vessels is basically scalariform, although deviations in pattern are present. Vessel elements of Ophiglossum are entirely comparable to those of leptosporangiate ferns.     

A model of bubble growth leading to xylem conduit embolism

The dynamics of a gas bubble inside a water conduit after a cavitation event was modeled. A distinction was made between a typical angiosperm conduit with a homogeneous pit membrane and a typical gymnosperm conduit with a torus-margo pit membrane structure. For conduits with torus-margo type pits pit membrane deflection was also modeled and pit aspiration, the displacement of the pit membrane to the low pressure side of the pit chamber, was found to be possible while the emboli was still small. Concurrent with pit aspiration, the high resistance to water flow out of the conduit through the cell walls or aspirated pits will make the embolism process slow. In case of no pit aspiration and always for conduits with homogeneous pit membranes, embolism growth is more rapid but still much slower than bubble growth in bulk water under similar water tension. The time needed for the embolism to fill a whole conduit was found to be dependent on pit and cell wall conductance, conduit radius, xylem water tension, pressure rise in adjacent conduits due to water freed from the embolising conduit, and the rigidity and structure of the pits in the case of margo-torus type pit membrane. The water pressure in the conduit hosting the bubble was found to occur almost immediately after bubble induction inside a conduit, creating a sudden tension release in the conduit, which can be detected by acoustic and ultra-acoustic monitoring of xylem cavitation.     

What the Penaeaceae alliance (Myrtales) tells us about the nature of vestured pits in xylem

Molecular studies indicate that Penaeaceae, Oliniaceae, and the monospecific families Alzateaceae and Rhynchocalycaceae form a clade of Myrtales. Of these four families, Penaeaceae have tracheids with vestured pits, whereas the others have septate fibers lacking vestures; all have vestured pits in vessels. Tracheid presence in Penaeaceae may be related to the arid South African habitats of the family. Presence of vestures on tracheids in families with vestured vessel pits is one indication that imperforate elements are tracheids and are conductive cells, whereas fiber-tracheids and libriform fibers are non-conductive. Tracheids occur widely in angiosperms and may be plesiomorphies or apomorphies. Combretaceae, the first branch of the Myrtales clade, has a great diversity of vesture features in vessels compared to the Penaeaceae alliance families. Alzatea has vestures that spread over the inside of the vessels, whereas in most taxa of the alliance, vestures are confined to the pit cavities and pit apertures. Vestures in the alliance tend to be globular in shape, and are bridged together by strands of wall material. Lignotubers and roots in Penaeaceae have vestures much like those in stems. Only a few species and genera (notably Alzatea) of the alliance have vesture features the pattern of which correlates with the current taxonomic system. Vestured pits should be viewed from the inside surface of vessels as well as the outer surface, and although sectional views of vestured pits are infrequent, they are very informative. Studies that explore diversity from one order or family to another are needed and offer opportunities for understanding the evolutionary significance of this feature.     

国产对囊蕨亚科（蹄盖蕨科）植物的管状分子

利用扫描电镜观察了国产蹄盖蕨科（Athyriaceae）对囊蕨亚科（Deparioideae）10种植物及双盖蕨属（Diplazium Sw．）3种植物根状茎的管状分子。结果显示,这些管状分子端壁和侧壁的形态及结构分别相同且侧壁具有穿孔板（多穿孔板）。根据穿孔板的形态特征,将该亚科的管状分子分为5种类型：（1）梯状穿孔板,无穿孔的二型性现象：（2）梯状穿孔板,有穿孔的二型性现象：（3）网状穿孔板：（4）梯状-网状混合的穿孔板：（5）大孔状穿孔板。按照纹孔膜残留的程度又可分为3种：部分区域有完整的纹孔膜、残留呈网状或线状以及很少或无纹孔膜残留。结合前人的研究资料,发现蕨类植物的管状分子与被子植物的导管分子在形态和输导机理上存在明显差异,管胞和导管分子不能仅仅根据纹孔膜的存在与否来确定,而应根据穿孔板存在于端壁还是侧壁进行判断,即穿孔板仅存在于端壁的管状分子为导管分子：端壁和侧壁形态及结构分别相同,有或无穿孔板的管状分子为管胞。由此可以推测蕨类植物和裸子植物中输导水分和矿物质的管状分子主要为管胞。单叶双盖蕨属（Triblemma（J．Sm．）Ching）与双盖蕨属管状分子的特征并不相似,显示了将单叶双盖蕨属从双盖蕨属独立出来归人对囊蕨亚科的合理性。根据管状分子的特征,推测假蹄盖蕨属（Athyriopsis Ching）和蛾眉蕨属（Lunathyrium Koidz．）可能是比较进化的属,而介蕨属（Dryoathyrium Ching）相对比较原始,单叶双盖蕨属的系统位置应介于假蹄盖蕨属与介蕨属之间。     

国产对囊蕨亚科(蹄盖蕨科)植物的管状分子

利用扫描电镜观察了国产蹄盖蕨科(Athyriaceae)对囊蕨亚科(Deparioideae)10种植物及双盖蕨属(Diplazium Sw.)3种植物根状茎的管状分子。结果显示, 这些管状分子端壁和侧壁的形态及结构分别相同且侧壁具有穿孔板(多穿孔板)。根据穿孔板的形态特征, 将该亚科的管状分子分为5种类型: (1)梯状穿孔板, 无穿孔的二型性现象; (2)梯状穿孔板, 有穿孔的二型性现象; (3)网状穿孔板; (4)梯状-网状混合的穿孔板; (5)大孔状穿孔板。按照纹孔膜残留的程度又可分为3种: 部分区域有完整的纹孔膜、残留呈网状或线状以及很少或无纹孔膜残留。结合前人的研究资料, 发现蕨类植物的管状分子与被子植物的导管分子在形态和输导机理上存在明显差异, 管胞和导管分子不能仅仅根据纹孔膜的存在与否来确定, 而应根据穿孔板存在于端壁还是侧壁进行判断, 即穿孔板仅存在于端壁的管状分子为导管分子; 端壁和侧壁形态及结构分别相同, 有或无穿孔板的管状分子为管胞。由此可以推测蕨类植物和裸子植物中输导水分和矿物质的管状分子主要为管胞。单叶双盖蕨属(Triblemma(J. Sm.) Ching)与双盖蕨属管状分子的特征并不相似, 显示了将单叶双盖蕨属从双盖蕨属独立出来归入对囊蕨亚科的合理性。根据管状分子的特征, 推测假蹄盖蕨属(Athyriopsis Ching)和蛾眉蕨属(Lunathyrium Koidz.)可能是比较进化的属, 而介蕨属 (Dryoathyrium Ching)相对比较原始, 单叶双盖蕨属的系统位置应介于假蹄盖蕨属与介蕨属之间。     

Inter-tracheid pitting and the hydraulic efficiency of conifer wood: the role of tracheid allometry and cavitation protection

Plant xylem must balance efficient delivery of water to the canopy against protection from air entry into the conduits via air-seeding. We investigated the relationship between tracheid allometry, end wall pitting, safety from air-seeding, and the hydraulic efficiency of conifer wood in order to better understand the trade-offs between effective transport and protection against air entry. Root and stem wood were sampled from conifers belonging to the Pinaceae, Cupressaceae, Podocarpaceae, and Araucariaceae. Hydraulic resistivity of tracheids decreased with increasing tracheid diameter and width, with 64 ± 4% residing in the end wall pitting regardless of tracheid size or phylogenetic affinity. This end-wall percentage was consistent with a near-optimal scaling between tracheid diameter and length that minimized flow resistance for a given tracheid length. There was no evidence that tracheid size and hydraulic efficiency were constrained by the role of the pits in protecting against cavitation by air-seeding. An increase in pit area resistance with safety from cavitation was observed only for species of the northern hemisphere (Pinaceae and Cupressaceae), but this variable was independent of tracheid size, and the increase in pit resistance did not significantly influence tracheid resistance. In contrast to recent work on angiosperm vessels, protection against air-seeding in conifer tracheids appears to be uncoupled from conduit size and conducting efficiency.     

Size and function in conifer tracheids and angiosperm vessels

The wide size range of conifer tracheids and angiosperm vessels has important consequences for function. In both conduit types, bigger is better for conducting efficiency. The gain in efficiency with size is maximized by the control of conduit shape, which balances end-wall and lumen resistances. Although vessels are an order of magnitude longer than tracheids of the same diameter, they are not necessarily more efficient because they lack the low end-wall resistance of tracheids with torus-margo pits. Instead, vessels gain conducting efficiency over tracheids by achieving wider maximum diameters. End-walls contributed 56-64% to total xylem resistance in both conduit types, indicating that length limits conducting efficiency. Tracheid dimensions may be more limited by unicellularity and the need to supply strength to homoxylous wood than by the need to protect against cavitation. In contrast, the greater size of the multicellular vessel is facilitated by fibers that strengthen heteroxylous wood. Vessel dimensions may be most limited by the need to restrict intervessel pitting and cavitation by air-seeding. Stressful habitats that promote narrow vessels should favor coexistence of conifers and angiosperms. The evolution of vessels in angiosperm wood may have required early angiosperms to survive a phase of mechanic and hydraulic instability.     

Recalcitrant vulnerability curves: methods of analysis and the concept of fibre bridges for enhanced cavitation resistance

Vulnerability curves (VCs) generally can be fitted to the Weibull equation; however, a growing number of VCs appear to be recalcitrant, that is, deviate from a Weibull but seem to fit dual Weibull curves. We hypothesize that dual Weibull curves in Hippophae rhamnoides L. are due to different vessel diameter classes, inter‐vessel hydraulic connections or vessels versus fibre tracheids. We used dye staining techniques, hydraulic measurements and quantitative anatomy measurements to test these hypotheses. The fibres contribute 1.3% of the total stem conductivity, which eliminates the hypothesis that fibre tracheids account for the second Weibull curve. Nevertheless, the staining pattern of vessels and fibre tracheids suggested that fibres might function as a hydraulic bridge between adjacent vessels. We also argue that fibre bridges are safer than vessel‐to‐vessel pits and put forward the concept as a new paradigm. Hence, we tentatively propose that the first Weibull curve may be accounted by vessels connected to each other directly by pit fields, while the second Weibull curve is associated with vessels that are connected almost exclusively by fibre bridges. Further research is needed to test the concept of fibre bridge safety in species that have recalcitrant or normal Weibull curves.     

Vertical and radial profiles in tracheid characteristics along the trunk of Douglas-fir trees with implications for water transport

The main stems of three young Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirbel) Franco) trees were dissected to obtain samples of secondary xylem from internodes axially along the trunk and radially within each internode. From these samples, measurements were obtained of tracheid diameter, length, the number of inter-tracheid pits per tracheid, and the diameter of the pit membranes. In addition, samples were obtained along the trunks of three old growth trees and also a small sample of roots for measurement of tracheid diameter. A gradient was apparent in all measured anatomical characters vertically along a sequence among the outer growth rings. These gradients arose not because of a gradient vertically along the internodes, but because of the strong gradients present at each internode among growth rings out from the pith. Tracheid characteristics were correlated: wider and longer tracheids had more numerous pits and wider pits, such that total pit area was about 6% of tracheid wall area independent of tracheid size. A stem model combining growth rings in parallel and internodes in series allowed for estimates of whole trunk conductance as a function of tree age. Conductance of the stem (xylem area specific conductivity) declined during the early growth of the trees, but appeared to approach a stable value as the trees aged.     

Direct measurements of intervessel pit membrane hydraulic resistance in two angiosperm tree species

The hydraulic resistance of pit membranes was measured directly in earlywood vessels of Fraxinus americana and Ulmus americana. The area-specific resistance of pit membranes (r(mem)) was higher than modeled or measured values obtained previously for hardwood species, with r(mem) of 5.24 × 10(3) MPa·s·m(-1) for Fraxinus and 2.56 × 10(3) MPa·s·m(-1) for Ulmus. The calculated resistance of pit canals was three orders of magnitude below total pit resistance indicating that pit membranes contributed the majority of resistance. Scanning electron microscopy indicated that pit membranes of Ulmus were thinner and more porous than those of Fraxinus, consistent with the difference in r(mem) between the species. Measurements of average vessel diameter and length and area of wall overlap with neighboring vessels were used to partition the vascular resistance between vessel lumen and pit membrane components. Pit membrane resistance accounted for 80% of the total resistance in Fraxinus and 87% in Ulmus in 2-yr-old branch sections. However, measurements of vessel dimensions in the trunk suggest that the division of resistance between pit membrane and lumen components would be closer to co-limiting in older regions of the tree. Thus, pit membrane resistance may be of greater relative importance in small branches than in older regions of mature trees.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF COSTA RICAN BLECHNUM FERNS

Hydraulic conductivities of stems, stipes, and elongate leaf stipes were determined for greenhouse-grown Blechnum (B. fraxineum, B. fragile, B. buchtienii, B. sprucei) and Salpichlaena (S. volubilis) plants collected in tropical rain forests of Costa Rica. Organ conductivity was examined in relation to morphology and tracheid characteristics in order to gain an understanding of factors influencing water flow. Hydraulic conductivity of plant organs was determined by measurement of transpiration rates, leaf areas, and water potential gradients. Erect stemmed species develop larger whole plant water potential gradients than elongate stemmed species for a similar transpiration rate. Elongate leaves develop even smaller water potential gradients for a given transpiration rate. Stems have larger hydraulic conductivities but smaller leaf-specific conductivities (LSCs) than stipes. Small conductivities and small LSCs are associated with short, erect stems. Elongate structures have large conductivities and large LSCs. Of the tracheid characteristics examined, the most important characteristics determining the magnitude of organ hydraulic conductivity are diameter, pit aperture area between tracheids, taper length, and cell length. Large conductivities of S. volubilis climbing leaf stipes are associated with very large-diameter tracheids (some > 200 μm), large tracheid number, exceptionally long tracheids (some > 4 cm), large pit aperture area between tracheids, short tracheid taper, and smooth tracheid lumen walls. Hagen-Poiseuille estimates of hydraulic conductivity range from 1.1 to 3.3 times the measured values. Conductivity of stipes is highly correlated with leaf area supplied by stipes. Conductivities of stems and elongate leaf stipes also correlate with leaf area supplied by these structures. Estimated hydraulic conductivities of field-grown Blechnum and Salpichlaena demonstrate that larger conductivities are associated with larger plants. This study contributes toward our knowledge of fern water relations and extends previous growth form/hydraulic architecture characterizations by providing a more comprehensive comparison of closely related species. In addition, this study provides evidence for the relative importance of tracheid characteristics in determining the magnitude of organ hydraulic conductivity.     

Plasmodesmata and pit development in secondary xylem elements

Developing pit membranes of secondary xylem elements in Drimys winteri, Fagus sylvatica, Quercus robur, Sorbus aucuparia, Tilia vulgaris and Trochodendron aralioides have been examined by transmission electron microscopy. Absence of plasmodesmata from the membranes of vessel elements and tracheids indicates that their pits develop independently of these structures. On the other hand, plasmodesmata are abundant in pit membranes between fibres, parenchyma cells, and combinations of these cell types in Fagus, Quercus and Tilia. In each case the plasmodesmata pass right through the developing pit membrane. In the case of Sorbus fibres, however, plasmodesmata were absent from the majority of pit membrane profiles seen in sections. Occasionally they were observed in large numbers associated with a swollen region on one side of the pit membrane between fibres and between fibres and parenchyma, radiating from a small area of the middle lamella. In the case of fibre to parenchyma pitting, this swelling was always found on the fibre side of the membrane, while on the other side a small number of plasmodesmata were present completing communication with the parenchyma cytoplasm. These observations are discussed with regard to the role of plasmodesmata in pit formation, and in the differentiation of the various cell types in secondary xylem. The significance their distribution may have for our understanding of xylem evolution is also discussed.     

Occurrence of plasmodesmata between differentiating vessels and other xylem cells inSorbus torminalis L. Crantz and their fate during xylem maturation

Summary The development of pit-pairs between differentiating xylem cells has been examined by transmission electron microscopy in young shoots ofSorbus torminalis. In some vessel-to-tracheid pits, as well as in previously studied intertracheid pits, a thickening of the pit membrane containing branched plasmodesmata was observed. A secondary wall-like cap was deposited over the thickening prior to cytoplasmic autolysis; some plasmodesmata, parallel to the plane of section, appeared to perforate the cap. At the end of the cell maturation stage, the central part of the primary wall thickening was hydrolysed, while the cap, including plasmodesmata remnants, appeared unaltered. In half-bordered pit-pairs between a parenchyma cell and a vessel or a tracheid, similar structures could be observed beside the conducting elements. When the vessel or tracheid matured, sealing of the pit membrane plasmodesmata resulted from the formation of a protective layer on the parenchyma-side rather than from the deposition of a cap on the conducting cell-side. These observations provide the first information on the presence of symplasmic connections in pits between differentiating vessels and neighbouring xylem cells. InS. torminalis, xylem differentiation is probably highly coordinated within a symplasmic domain; the persistence of such connections may account for the lack of specialization ofSorbus wood.