PRESENCE OF VESSELS IN WOOD OF SARCANDRA (CHLORANTHACEAE); COMMENTS ON VESSEL ORIGINS IN ANGIOSPERMS

Sarcandra is the only genus of Chloranthaceae hitherto thought to be vesselless. Study of liquid-preserved material of S. glabra revealed that in root secondary xylem some tracheary elements are wider in diameter and have markedly scalariform end walls combined with circular pits on lateral walls. Examination of these wider tracheary elements with scanning electron microscope (SEM) demonstrated various degrees of pit membrane absence in the end walls. Commonly a few threadlike fibrils traverse the pits (perforations); these as well as intact nature of pit membranes in pits at ends of some perforation plates are evidence that lack of pit membranes does not result from damage during processing. Some perforations lack any remnants of pit membranes. Although perforation plates and therefore vessels are present in Sarcandra roots, no perforations were observed in tracheary elements of stems or lignotubers. Further, stem tracheids do not have the prominently scalariform end walls that the vessel elements in roots do. Presence of vessels in Sarcandra removes at least one (probably several) hypothetical events of vessel origin that must be postulated to account for known patterns of vessel distribution in angiosperms, assuming that they are primitively vesselless. Seven (perhaps fewer) vessel origin events in angiosperms could account for these patterns; two of those events (Nelumbo and monocotyledons) are different from the others in nature. Widely accepted data on trends of vessel specialization in woody dicotyledons yield an unappreciated implication: vessel specialization has happened in a highly polyphyletic manner in dicotyledons, and therefore multiple vessel origins represent a logical extension backward in time. If a group of vesselless dictyoledons ancestral to other angiosperms existed, they can be hypothesized to have had a relatively homogeneous floral plan now that Sarcandra-like plants no longer need be imagined within that group. Sarcandra and other Chloranthaceae show that the borderline between vessel absence and presence is less sharp than generally appreciated.     

PRIMARY XYLEM ELEMENTS AND ELEMENT ASSOCIATIONS OF ANGIOSPERMS

Observations based on a study of more than 1350 species distributed among 165 families of angiosperms are presented. The tracheary elements which mature later than the helical ones in the protoxylem-metaxylem transition are described in terms of a 2-phase wall deposition process. These elements have a helical framework (first-order secondary wall) between the gyres of which is deposited additional secondary wall material in the form of sheets or strands or both (second-order framework). This is indicated both by the sequence of mature elements throughout the primary xylem and by the ontogeny of later maturing elements. Elements in which the second-order system of the secondary wall is deposited more or less synchronously and in which the sheets or strands are restricted to cell edges, i.e., lines of intersection of adjacent cell faces, are interpreted as primitive. Elements in which the second-order sheets and strands appear nonsynchronously and with less regard for cell edges are interpreted as advanced. Alternate pitting results from the appearance of oblique second-order strands with subsequent wall deposition maintaining strand orientation such that pit axes are tilted. In certain elements second-order strands are deposited before, and wall deposition continues after, the cessation of cell elongation. This results in an alternate pit pattern and may explain certain irregular patterns. Branching of the first-order helix seems to be relatively insignificant in the development of more elaborate wall patterns. It is more significant in perforation plate elaboration. “Open pits” occur in a number of dicotyledons. These are pit-like openings which are extended laterally as the thin areas between the gyres of a helix or comparable openings in a reticulum. They constitute a conspicuous feature of the entire protoxylem-metaxylem transition in certain species. The simple perforation plate of only certain angiosperms seems to be the result of bar breakdown in a multipored plate. Reduction in pore number is also the result of fusions in the first-order framework lateral to a multipored plate. In dicots this trend rarely culminates in a simple perforation plate, but it frequently does so in monocots. This type of pore number reduction and enlargement frequently accompanies bar breakdown in dicots and certain monocots. The perforation plate is often simple as the result of a terminalization on the cell, in which case the pore does not intersect the first-order framework. This type of perforation plate occurs in species with and without more obliquely oriented simple perforation plates subject to a breakdown interpretation. Complex multiperforate plates are interpreted as falling into 3 categories : Plates in which a reticulum has resulted from introduction of second-order secondary wall strands at various orientations and with variable amounts of distortion following deposition; plates in which a reticulum has resulted from ramification and fusion of the strands of the first-order framework ; and plates which are multiperforate as the result of the presence of a number of separate loci of breakdown within a single pore membrane. Possible ontogenetic complexities in the development of perforation plates subject to breakdown interpretation are discussed. Protoxylem-metaxylem transitions are described in terms of the sequence of types of perforation plates. Most sequences with various types of plates support the concept of progressively earlier ontogenetic expression of specialized features with progressive elimination of primitive ones. The concept is contradicted by those species in which nearly all perforation plates are simple. Non-simple plates in these species are found from early protoxylem through mid-metaxylem but not in the earliest protoxylem. If non-simple plates are uncommon in a species, they may have a different ontogenetic history in terms of procambial divisions and apical cell growth. In the monocots with a variety of perforation plate types, the probability that a given element will be imperforate or perforate in one way or another will depend on its diameter, not on its position within the sequence. The occurrence of vessels of very limited extent is discussed. It has been calculated that vessels in the stem of Scleria average from 2 to nearly 50 cells in length depending on their diameter.     

FLORA OF THE LOWER CRETACEOUS CEDAR MOUNTAIN FORMATION OF UTAH AND COLORADO. PART III: ICACINOXYLON PITTIENSE N. SP

Icacinoxylon pittiense, a new species of angiospermous wood from the Lower Cretaceous Cedar Mountain Formation of Utah is described and compared with similar fossil and modem woods. It is distinguished from other species of Icacinoxylon by its thick-walled fiber-tracheids with their walls making up at least 50% of the total diameter of the cells, conspicuous bordered pits with obliquely crossing extended apertures on both the tangential and radial walls of its fiber-tracheids, scalariform perforation plates with as few as four or greater than 30 bars, transitional opposite to scalariform pitting on its vessel walls, thick-walled ray cells, and distinct sheath or border cells in its rays. Icacinoxylon pittiense is the first species of this genus to be reported from Cretaceous sediments. This wood is of special interest because very few angiosperm woods have been reported from lower Cretaceous strata.     

Vessel dimensions in liana and tree species of Gnetum (Gnetales)

Maximum vessel diameters were examined in the secondary xylem of stems of Gnetum of various sizes. One tree (G. gnemon) and 13 liana species were compared. In three species, vessel length distributions were determined by the latex paint method, and showed many short and fewer long vessels. Latex and compressed air methods, used to find the maximum vessel lengths, showed that maximum vessel lengths were similar for three species of Gnetum. In old stems, mean and maximum vessel diameters tended to be greater in lianas than in the tree species. The skewed distribution of vessel lengths and the trend of wider vessels in lianas as compared to trees were similar to those distributions and trends described previously for angiosperms. In random samples of macerated wood of three species, simple perforation plates were most common in vessel members of all species. Foraminate and modified foraminate perforations were less frequent. Average diameter of vessel members with either foraminate or modified foraminate perforations was less than for those with simple perforations. The resemblance of Gnetum vessels to those of angiosperm trees and vines is most likely a case of convergent evolution (homoplasy) in xylem characteristics.     

PIT MEMBRANE REMNANTS IN PERFORATION PLATES OF PRIMITIVE DICOTYLEDONS AND THEIR SIGNIFICANCE

Perforations of vessel elements characteristically retain remnants of pit membranes (primary walls) in woods of species of more than 30 families of dicotyledons. Scanning electron microscopy is necessary to demonstrate presence and type of membrane remnant. Species with these remnants in perforations given in earlier literature as well as those newly reported here are listed. Perforation membrane remnants may take the form of flakes, strands, or webs, and particular types may characterize particular families (e.g., strands or bands in Illiciaceae). Some families have abundant perforation membrane remnants (e.g., Chloranthaceae, Illiciaceae). Where membranes are nearly intact, they are porose and closely resemble the porose pit membranes on end walls of Tetracentron tracheids. In Tetracentron, however, tracheary elements are monomorphic, so vessel origin cannot yet be said to have occurred. Membrane remnants in perforations are regarded as a relictual primitive feature that should be added to the list of primitive character states claimed for vessel elements in angiosperms; alternative hypotheses are considered and discussed, and evidence from DNA phylogenies is needed. In vessel-bearing dicotyledons with membrane remnants in perforations, many perforations are relatively clear, but an appreciable proportion of perforation plates do have membrane remnants.     

PITTING IN PSILOPHYTON DAWSONII,AN EARLY DEVONIAN TRIMEROPHYTE

Tracheids of Psilophyton dawsonii Banks, Leclercq, and Hueber 1975 from calcareous pebbles of late Early Devonian age on the Gaspé Peninsula, eastern Canada, are shown to have scalariform bordered and circular bordered pits. Macerated tracheids embedded in Spurr epoxy resin and sectioned at 2 μm; peels, ground sections, SEM and petrographic observations all indicate that interconnections between the scalariform bars are composed of secondary wall material and that the patterns produced are not artifacts of preservation. The interconnections produce a wide range of patterns from simple vertical strands, to reticula, to more extensive interconnections that outline circular openings. The circular openings result in scalariform bordered pits that are considered to be multiaperturate. P. dawsonii is believed to illustrate the most complex pitting yet demonstrated conclusively in Early Devonian time. Among trimerophytes P. forbesii, tracheids of P. charientos, and Hostinella from Röragen, Norway lack only the circular bordered pits.     

Single‐vessel flow measurements indicate scalariform perforation plates confer higher flow resistance than previously estimated

During vessel evolution in angiosperms, scalariform perforation plates with many slit‐like openings transformed into simple plates with a single circular opening. The transition is hypothesized to have resulted from selection for decreased hydraulic resistance. Previously, additional resistivity of scalariform plates was estimated to be small – generally 10% or less above lumen resistivity – based on numerical and physical models. Here, using the single‐vessel technique, we directly measured the hydraulic resistance of individual xylem vessels. The resistivity of simple‐plated lumens was not significantly different from the Hagen–Poiseuille (HP) prediction (+6 ± 3.3% mean deviation). In the 13 scalariform‐plated species measured, plate resistivity averaged 99 ± 13.7% higher than HP lumen resistivity. Scalariform species also showed higher resistivity than simple species at the whole vessel (+340%) and sapwood (+580%) levels. The strongest predictor of scalariform plate resistance was vessel diameter (r² = 0.84), followed by plate angle (r² = 0.60). An equation based on laminar flow through periodic slits predicted single‐vessel measurements reasonably well (r² = 0.79) and indicated that Baileyan trends in scalariform plate evolution maintain an approximate balance between lumen and plate resistances. In summary, we found scalariform plates of diverse morphology essentially double lumen flow resistance, impeding xylem flow much more than previously estimated.     

鹅掌楸属(木兰科)次生木质部导管穿孔板的比较研究

利用扫描电子显微镜对鹅掌楸属仅存的2个自然种鹅掌楸和北美鹅掌楸的次生木质部导管穿孔板特征进行了详细的研究。结果显示,鹅掌楸属的2个种均以梯状穿孔板为主,同时存在网状-梯状混合穿孔板。鹅掌楸和北美鹅掌楸的导管穿孔板具有明显差异:(1)鹅掌楸具网状穿孔板,而北美鹅掌楸没有观察到;(2)北美鹅掌楸具单穿孔板,而鹅掌楸在该实验中未发现;(3)北美鹅掌楸具有横隔较粗的梯状穿孔板且横隔数目较多;(4)鹅掌楸的导管穿孔板多数横隔较少;(5)北美鹅掌楸的穿孔板倾斜角度较大;(6)北美鹅掌楸具麻黄式穿孔板的存在,且有纹孔膜残留存在。研究认为,北美鹅掌楸导管分子穿孔板分化较鹅掌楸更为剧烈。     

Pit membrane remnants in perforation plates of Hydrangeales with comments on pit membrane remnant occurrence, physiological significance and phylogenetic distribution in dicotyledons

Perforation plates from ten species of seven genera of Hydrangeales sensu Thorne were studied using scanning electron microscopy (SEM). The presence of pit membranes in perforations ranges from abundant, as in Carpenteria and Hydrangea, to minimal, as in Deutzia, Escallonia and Philadelphus. Abnormally great pit membrane presence may result from the presence of secondary compounds that inhibit lysis, as in Quintinia serrata; such interference with the natural lysis process may or may not be evident from coarseness and irregularity of pit membrane surface and of threads composing the pit membrane remnants. The presence of pit membrane remnants in perforation plates is hypothesized to be a symplesiomorphy, found in a fraction of dicotyledons with scalariform perforation plates (but still in an appreciable number of species). Pit membrane remnant presence may represent incomplete lysis of primary wall material (cellulose microfibrils) in species that occupy highly mesic habitats, where such impedance in the conductive stream does not have an appreciable negative selective value. This physiological interpretation of pit membrane remnants in perforations is enhanced by the phylogenetic distribution as well as the strongly mesic ecological preferences of species that exemplify this phenomenon in dicotyledons at large. Families with pit membrane presence in perforations are scattered throughout phylogenetic trees, but they occur most often in basal branches of major clades (superorders) or as basal branches of orders within the major clades. Further study will doubtless reveal other families and genera in which this phenomenon occurs, although it is readily detected only with SEM. Phylogenetic stages in the disappearance of pit membrane remnants from perforation plates are described, ranging from intact pit membranes except for presence of pores of various sizes, to presence of membrane remnants only at lateral ends of perforations and in one or two perforations (arguably pits) at the transition between a perforation plate and subadjacent lateral wall pitting. Developmental study of the mechanism and timing of lysis of pit membranes in perforations, and assessment of the role of the conductive stream in their removal, are needed to enhance present understanding of vessel evolution. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146 , 41–51.     

领春木茎次生木质部中导管穿孔板的变异

领春木Euptelea pleiosperma Hook. f. &; Thoms.隶属领春木科Eupteleaceae。该科为东亚特有的单型科,其系统位置一直颇有争议。本文对中国产领春木茎次生木质部中导管穿孔板的变异进行了观察,以期为它的系统位置提供进一步的解剖学证据。结果表明,领春木茎次生木质部中包括无明显穿孔板的管胞状导管和典型的导管两种类型。在无明显穿孔板的导管中,穿孔中的纹孔膜全部或部分消失,但穿孔无规则排列或聚集,不形成具典型的形态特征的穿孔板;在典型的导管中,穿孔板形态变异较大,包括几个类型:网状穿孔板(含麻黄式穿孔板)、网状和梯状混合型穿孔板、梯状穿孔板、梯状穿孔板向单穿孔板的过渡。在上述导管穿孔板类型中,只有梯状穿孔板的穿孔中可以观察到纹孔膜的残余。在领春木次生木质部中也观察到了端壁多穿孔板及侧壁穿孔板。根据观察结果,我们认为领春木次生木质部导管穿孔板的许多特征说明该科可能处于毛茛目中比较原始的系统位置。     

WOOD ANATOMY AND RELATIONSHIPS OF LACTORIDACEAE

Mature wood of Lactoris, not previously available for study, reveals ten distinctive characters: vessels with simple perforation plates; vessels in pore multiples; vessel-to-axial parenchyma pits scalariform or transitional, vessel-to-vessel pits alternate; fiber-tracheids with vestigial pits; fiber-tracheids, vessels, and axial parenchyma storied; axial parenchyma vasicentric scanty; axial parenchyma either not subdivided or, if subdivided, with thin nonlignified walls between the cells (like the septa in septate fibers); rays wide and tall, little altered during ontogeny; ray cells upright; and ray cells taller adjacent to fascicular areas. All of these features occur in woods of Piper and other Piperaceae. The systematic position of Lactoris is therefore reassessed. Evidence available to date is consonant with placement of Lactoridaceae in Piperales, in which it would be more primitive than Piperaceae or Saururaceae. Features cited as evidence for alternative placements of Lactoridaceae are reviewed.     

Distribution of scalariform and simple perforation plates within the vessel network in secondary xylem of Araliaceae and its implications for wood evolution

The distribution of scalariform and simple perforation plates along vessels in Arthrophyllum otopyrenum, Meryta tenuifolia, and Polyscias multijuga (Araliaceae) is examined. In all three species, most vessels bear simple perforation plates only, but the combination of simple and scalariform perforation plates in variable ratios also occurs. Aggregated arrangement of scalariform perforation plates along the vessels was statistically confirmed in some vessel portions. The scalariform perforation plates occur mostly in narrow vessels that are grouped in multiples. Within the clade represented by Polyscias and Arthrophyllum, the evolutionary transition from scalariform to simple perforation plates is realized as the gradual elimination of vessels or vessel portions with scalariform perforation plates, but is not accompanied by a gradual decrease of the number of bars per perforation plate. The narrow vessels that are grouped in vessel multiples are likely to retain the ability to develop scalariform plates, which could promote the evolution from simple to scalariform perforation plates as is the case within Meryta.     

八属木兰科植物木材导管分子的比较解剖

本文比较描述了我国木兰科木莲属、华盖木属,长蕊木兰属,拟单性木兰属,合果木属,观光木属,香木兰属和鹅掌楸属等8属22种植物次生质部的导管分子,这些导管分子的长度和宽度都有差异,木莲属的所有种中都有具梯状穿孔板的导管分子,但在少数种中偶然可见到单穿孔,在其它属中,也都有具梯状穿孔板的导管分子,仅在观光木属中偶见的单穿孔,木莲属大多数种均无螺纹加厚,仅桂南木莲一种除外,另外,除了合果木属和鹅掌楸无螺纹加厚外,其余均有此种加厚,各属的导管分子还存在其它一些差异。     

SEM studies on vessels in ferns. 17. Psilotaceae

Perforation plates are reported in aerial and subaerial axes of Psilotum nudum and in aerial axes of Tmesipteris obliqua. In Psilotum, both perforations lacking pit membranes and perforations with pit membrane remnants were observed. Perforation plates in Psilotum may consist wholly of one type or the other. In Tmespteris, perforations have threadlike pit membranes or consist of porose pit membranes. Wide perforations alternating with narrow pits, a conformation observed in various ferns, were observed in Psilotum (subaerial axes). In Psilotum, perforations are more common in metaxylem than in protoxylem; perforations in protoxylem consist of primary wall areas containing small circular porosities or relatively large circular to oval perforations. There are no modifications in the secondary wall framework of protoxylem or metaxylem in Psilotum or Tmesipteris that would permit one to distinguish presence of perforations or perforation plates with light microscopy, and scanning electron microscopy (SEM) is required for demonstration of porose walls or perforations. The tracheary elements of the Psilotaceae studied have no features not also observed in other ferns with SEM.     

蜈蚣草导管分子的观察

对蜈蚣草(Pteris vittata)叶轴中导管分子进行了观察。结果表明,其导管分子十分细长,均具长、且十分倾斜的梯状穿孔板,穿孔没有纹孔膜的残余,与侧壁上的梯纹纹孔有着明显的差异。     

蜈蚣草导管分子的观察

对蜈蚣草(Pteris vittata)叶轴中导管分子进行了观察。结果表明,其导管分子十分细长,均具长、且十分倾斜的梯状穿孔板,穿孔没有纹孔膜的残余,与侧壁上的梯纹纹孔有着明显的差异。     

STRUCTURE OF TRACHEIDS IN THREE SPECIES OF LYCOPODIUM

The structure of tracheids in Lycopodium lucidulum, L. clavatum, and L. tristachyum was studied with the light microscope. Protoxylem development is at least sometimes and possibly always mesarch in indeterminate axes of all three species. Centrifugally formed protoxylem elements are reticulate and discontinuities in the secondary walls of these elements are sometimes conspicuously bordered. Wall thickenings of first formed protoxylem elements consist mainly of indirectly connected rings. Late centripetally formed protoxylem elements and transitional elements have a reticulate secondary wall pattern. The narrowest metaxylem elements have circular bordered pits while in wider metaxylem elements pits are bordered and may vary from circular to scalariform. Pitting is uniseriate to triseriate in tracheids of all three species, and intermittent tetraseriate pitting was occasionally observed in L. lucidulum. Crassulae occur in tracheids of the three species, and in L. clavatum an additional framework, probably representing thickened compound middle lamella, is also present. Pits often appear helically arranged, and in all three species pits are connected by thin areas in the secondary wall. Macrofibrils approximately 0.5 μ wide were observed in tracheids of the three species. In L. clavatum the arrangement of macro-fibrils was predominantly bidirectional.     

ANATOMY AND AFFINITIES OF PENTHORUM

The genus Penthorum L. consists of two species of perennial herbs, P. sedoides of eastern North America and P. chinense of eastern Asia. Penthorum has long been considered intermediate between Crassulaceae and Saxifragaceae. An anatomical study of both species was undertaken to contribute to a better understanding of the relationships of these plants. Prominent anatomical features of Penthorum include: an aerenchymatous cortex and closely-spaced collateral vascular bundles of stems; one-trace unilacunar nodes; brochidodromous venation, rosoid teeth bearing hydathodes, and anomocytic stomata of leaves; angular vessel elements with many-barred scalariform perforation plates and alternate to scattered intervascular pits; thin-walled non-septate fiber-tracheids; abundant homocellular erect uniseriate and biseriate rays; and absence of axial xylem parenchyma. In general, Penthorum possesses neither the morphological nor the anatomical synapomorphies which define Crassulaceae, and features shared with Saxifragaceae are largely symplesiomorphous. Thus Penthorum is probably best classified in the monogeneric Penthoraceae.     

Resistances to fluid flow of model xylem vessels with simple and scalariform perforation plates

The resistances of xylem vessel walls and perforation plates has been investigated using 18 large-scale physical models made of plastic tubing into which scale models of plates were inserted. Flow of water through vessels was modelled using glycerol instead of water to keep the Reynolds number below 0.1. The technique proved easy, cheap and reliable.Results showed that perforation plate resistance is low compared with the resistance of the walls, whatever the plate morphology; plates only provided 0.6-18.6% extra resistance. Simple plates provided less resistance than scalariform plates, but because they are arranged closer together in vessels, resistance values (1.7-5.1%) overlap with those of scalariform plates. The resistance of scalariform plates varied in a systematic way with their morphology. For a given plate angle, increasing the number of bars increased resistance. For a given bar number, increasing the angle of the plate to the vessel axis also increased the resistance. However, for a given gap between bars, increasing the angle of the plate to the vessel axis decreased resistance. These results are discussed in the light of theories about the function of perforation plates.Keywords: Flow, xylem vessel, perforation plate, models.      

Scalariform-to-simple transition in vessel perforation plates triggered by differences in climate during the evolution of Adoxaceae

Background and Aims Angiosperms with simple vessel perforations have evolved many times independently of species having scalariform perforations, but detailed studies to understand why these transitions in wood evolution have happened are lacking. We focus on the striking difference in wood anatomy between two closely related genera of Adoxaceae, Viburnum and Sambucus, and link the anatomical divergence with climatic and physiological insights.Methods After performing wood anatomical observations, we used a molecular phylogenetic framework to estimate divergence times for 127 Adoxaceae species. The conditions under which the genera diversified were estimated using ancestral area reconstruction and optimization of ancestral climates, and xylem-specific conductivity measurements were performed.Key Results Viburnum, characterized by scalariform vessel perforations (ancestral), diversified earlier than Sambucus, having simple perforations (derived). Ancestral climate reconstruction analyses point to cold temperate preference for Viburnum and warm temperate for Sambucus. This is reflected in the xylem-specific conductivity rates of the co-occurring species investigated, showing that Viburnum lantana has rates much lower than Sambucus nigra.Conclusions The lack of selective pressure for high conductive efficiency during early diversification of Viburnum and the potentially adaptive value of scalariform perforations in frost-prone cold temperate climates have led to retention of the ancestral vessel perforation type, while higher temperatures during early diversification of Sambucus have triggered the evolution of simple vessel perforations, allowing more efficient long-distance water transport.