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.     

Comparative Anatomical Studies on the Vessel Elements of Chinese Magnolia and Michelia

This paper deals mainly with comparative studies on elements of secondary xylem in 39 species of Chinese Magnolia and Michelia. The former plants present longer and slender vessels. Most of them have simple perforation and only a few scalariform perforation plates. In contrast, most of the Michelia species possess scalariform perforation plates and only one with simple perforation. The scalariform perforation plates of Magnolia species, if present, exhibit more bars than that of Michelia With few exceptions, there is no spiral thickening on the vessel wall in Magnolia, whereas in Michelia it is lust the opposite. In addition, some other differences in vessel elements between these two genera are also discussed.     

国产五味子科植物导管分子的比较解剖

对五味子科 2属 1 9种植物导管分子的结构进行了比较观察。结果表明 :2属均以具缘纹孔导管为主 ,少有梯纹和螺纹导管 ,南五味子属多数种还有梯孔纹导管 ;2属导管长度和宽度差异很小 ,长 /宽比值有差异 ;根据穿孔板的结构可分为 2种形式 :( 1 )单穿孔板 ,这类穿孔板较普遍存在 ;( 2 )梯形穿孔板 ,发现 2属中的红花五味子 (Schisandrarubriflora)、五味子 (S .chinensis)、翼梗五味子 (S .henryi)、铁箍散 (S .propinquavar.sinensis)和黑老虎 (Kadsuracoccinea) 5种植物的导管分子具有此类穿孔板。其中五味子属中的五味子导管分子只具有梯形穿孔板 ,无单穿孔板 ,但横条较少 ,多为 2～ 3条。     

中国木兰属和含笑属导管分子的比较解剖

本文对我国木兰科的39种木兰属和含笑属植物次生木质部的导管分子进行了初步分析。两属导管分子的长度和宽度略有差异。木兰属中多数种的导管分于有单穿孔板,但有的可见到梯状穿孔板。含笑属植物的导管分子大多具有梯状穿孔板,仅有一种可看到单穿孔板。在具有梯状穿孔板的木兰属植物中,穿孔板的横隔数目较含笑属的多。木兰属的导管壁上一般无螺纹加厚;含笑属则相反。此外,在两属之间,导管尚存在一些其它差异。     

东北刺人参茎次生木质部结构植物学研究

东北刺人参为五加科植物,本文对其茎次生木质部进行结构植物学研究,发现其为环孔材,射线为异形Ⅱ型。在离析材料中,发现其导管分子穿孔板有单穿孔板和梯状穿孔板2种。具不同穿孔板的导管分子可分为8种。还有维管管胞、纤维管胞和韧型木纤维。8种导管分子是只有中央具1个单穿孔的导管;中央有2个穿孔的导管,其中一个孔是单穿孔,另一个是梯状穿孔;一端是梯状穿孔板,另一端为孔纹增厚的导管;一端是单穿孔。另一端是孔纹增厚的导管;一端为单穿孔板。另一端为梯状穿孔板的导管;两端都是梯状穿孔板的导管;侧壁具有3个穿孔的导管;两端都具单穿孔的导管。在东北刺人参的个体发育中重演了系统发育过程中导管分子穿孔板的演化过程。     

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。     

Comparative Anatomical Studies on the Vessel Elements of Eight Genera in Magnoliaceae

This paper deals mainly with comparative studies on vessel elements of secondary xylem in Chinese Manglietia, Manglietiastrum, Alcimandra, Parakmeria, Paramichelia, Tsoongiodendron, Aromadendron and Liriodendron. There are some differences between the genera in vessel element length and width. Manglietia species all have scalariform perforation plates, only a few also have simple perforations. The other genera have scalariform perforation plates. only one (Tsoongiodendron odorum Chun) has simple perforations in addition. There are no spiral thickenings in Manglietia except for one species [Manglietia chingii Dandy (M. temuipes Dandy)] . Paramichelia and Liriodendron have no spiral thichenings, but they are present in the other genera. In addition, some other differences in vessel elements between these 8 genera are also discussed.     

Vessel-bearing stems of ASOVINEA TIANIIgen. et sp. nov. (Gigantopteridales) from theUpper Permian of Guizhou Province, China

Permineralized gigantopterid stems of Vasovinea tianii Li et Taylor gen. et sp. nov. were collected from the Upper Permian of Guizhou Province, China. They are slender and bear prickles, trichomes, and compound hooks. Internally, the stems have a sparganum cortex, eustele, and secondary xylem. The mesarch protoxylem tracheids have annular to helical thickenings, and metaxylem tracheary elements have scalariform and/or transversely elongated, bordered pits, while those of the secondary xylem have scalariform to circular bordered pits. Importantly, the inner part of the secondary xylem has large vessel elements with foraminate-like perforation plates. The hooks and other morphological and anatomical characteristics are similar to those found in gigantopterids, suggesting that Vasovinea is a member of the Gigantopteridales. The vegetative plant is reconstructed from permineralized stems and Gigantopteris-type leaves based on the anatomical similarities and intimate association. The eustele, secondary xylem, and other features support the placement of the order among the seed plants. Ecologically, Vasovinea is suggested to have been a vine or liana that used compound hooks to climb among the trees in a Permian tropical rain forest. The occurrence of vessels could have been an efficient adaptation to allow the slender stems to conduct sufficient water to the large Gigantopteris-type leaves.     

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。     

Origin and nature of vessels in monocotyledons. 5. Araceae subfamily Colocasioideae

Tracheary elements from macerations of roots and stems of one species each of five genera of Araceae subfamily Colocasioideae were studied by means of SEM (scanning electron microscopy). All of the genera have vessel elements not merely in roots, as previously reported for the family as a whole, but also in stems. The vessel elements of stems in all genera other than Syngonium are less specialized than those of roots; stem vessel elements are tracheid-like and have porose pit membrane remnants in perforations. The perforations with pit membrane remnants demonstrate probable early stages in evolution of vessels from tracheids in primary xylem of monocotyledons. The vessel elements with such incipient perforation plates lack differentiation in secondary wall thickenings between perforation plate and lateral wall, and such vessel elements cannot be identified with any reliability by means of light microscopy. The discrepancy in specialization between root and stem vessel elements in genera other than Syngonium is ascribed to probable high conductive rates in roots where soil moisture fluctuates markedly, in contrast with the storage nature of stems, in which selective value for rapid conduction is less. Syngonium stem vessels are considered adapted for rapid conduction because the stems in that genus are scandent. Correlation between vessel element morphology and ecology and habit are supported. Although large porosities in vessel elements facilitate conduction, smaller porosities may merely represent rudimentary pit membrane lysis.     

Apostasiads, systematic anatomy, and the origins of Orchidaceae

Anatomical study of several species of the putatively primitive orchids Apostasia and Neuiviedia was based on specimens of leaves, stems and rootS. Research was carried out in the hope of providing objective information from anatomy towards unravelling the relationships of these plants, and especially towards answering the question of whether extant orchids evolved from these two genera, or from plants like them. The morphology of Apostasia and Neuwiedia has evoked the dogma of primitiveness because of the two or three anthers borne on separate filaments and the free style and stigma which characterize the flowerS. In most other orchids there is only one anther and filaments and styles are fused to form the column (gynostemium). The general anatomical structure of apostasiads shows features present in other orchids; none is restricted to Apostasia and Neuwiedia. Tracheary anatomy, however, shows vessels in roots, but not in any part of the shoot system. Vessel members are characterized by chiefly simple perforation plates in contrast with other orchids where scalariform perforation plates predominate in vessel members of the root. These phenomena, considered from the phylogenetic standpoint, would cast serious doubt on the possibility that plants with scalariform perforation plates, the ancestral, or primitive condition, could have arisen from plants with simple perforation plates, the derived, or advanced condition. On this basis the apostasiads could not have given rise to the di- and monandrous orchidS. Of the several suggested origins for orchids, Hypoxidaceae, or plants similar to them, whether in Asparagales, Liliales or Haemadorales of different authors, could have been the progenitors of orchids as a group, including the apostasiadS. Because of the unique combination of floral features in the apostasiads, their predominantly simple perforation plates, and their overall anatomical similarity to orchids in general, it would appear appropriate to consider them as a subfamily, Apostasioideae, of Orchidaceae sensu lato.     

Leaf and twig anatomy of the Pterostemonaceae (Engl.) Small: ecological and systematic features

An anatomical investigation of the leaves and twigs of Pterostemonaceae (Engl.), a monogeneric family of two species, has been made. The following anatomical characters in the leaf are of particular interest: glandular hairs, hydathodes on the marginal dentations and secretory substances in the glands and palisade cells. Characters of interest in the twig xylem include: vessel lumina of very small tangential diameter and with simple perforation plates; fibriform vessels with scalariform plates having one to six bars and also plates with perforations in irregular patterns.     

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.      

Comparative wood anatomy of epacrids (Styphelioideae,Ericaceae s.L.)

The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel-ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze-thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.     

VESSELS IN JUNCALES: II. CENTROLEPIDACEAE AND RESTIONACEAE

The occurrence and specialization of vessels in 6 genera and 14 species of Centrolepidaceae and 25 and 72, respectively, of Restionaceae are described. Those in Anarthriaceae (1 and 3) and Ecdeiocoleaceae (1 and 1), newly separated from Restionaceae, were also examined. Vessels in Ecdeiocolea are highly specialized; those in Centrolepidaceae are relatively primitive. Vessels in Restionaceae and Anarthriaceae are intermediate. Vessels in Cyperaceae and Gramineae are compared with those in Juncales in a review of vessels in Glumiflorae. Thurniaceae has most primitive vessels on the average, Juncales has more primitive vessels in leaves and inflorescence axes than does Cyperales; the opposite is true for roots, rhizomes and stems. Vessels in Graminales on the average are much more specialized than those in Cyperales and Juncales.     

五味子科比较木材解剖及其系统学意义

在光学显微镜和扫描电镜下观察了南五味子属Kadsura 7种21个样品和五昧子属Schisandra 8种14个样品的木材解剖特征,结果表明次生木质部的导管分子类型、导管一射线间纹孔的排列方式、射线类型、射线细胞形状等性状在科的水平上很稳定,这些共同特征都支持五味子科Schisandraceae是比较自然的类群。在五昧子科中发现木材导管单生、具梯状穿孔板、导管壁具梯形排列的纹孔以及木射线异型等原始性状,支持五味子科在被子植物中的原始地位。此外,该科木材还具有单穿孔板导管、导管次生壁具螺纹加厚、具分隔纤维等较为特化的性状状态,这种性状进化水平的异等级现象,使五味子科表现出不同进化水平性状的镶嵌组合。根据木材解剖性状对五味子科进行UPGMA聚类分析,所得结果显示南五昧子属和五味子属在木材解剖特征方面有一定的交叉和重叠,这与分子系统学的结论一致,表明这两个属关系密切,可能起源于共同的祖先。通过比较五昧子科与八角科Illiciaceae的木材解剖特征,进一步证明两个科的亲缘关系很近,不支持将五味子科从八角目Illiciales中独立出来成立五味子目Schisandrales的观点。     

Morphological analysis of vessel elements for systematic study of three Zingiberaceae tribes

Zingiberaceae containing over 1,000 species that are divided into four subfamilies and six tribes. In recent decades, there has been an increase in the number of studies about vessel elements in families of monocotyledon. However, there are still few studies of Zingiberaceae tribes. This study aims to establish systematic significance of studying vessel elements in two subfamilies and three tribes of Zingiberaceae. The vegetative organs of 33 species processed were analysed by light and scanning electron microscopy and Principal Component Analysis was used to elucidate genera boundaries. Characteristics of vessel elements, such as the type of perforation plate, the number of bars and type of parietal thickening, are proved to be important for establishing the relationship among taxa. Scalariform perforation plate and the scalariform parietal thickening are frequent in Zingiberaceae and may be a plesiomorphic condition for this taxon. In the Principal Component Analysis, the most significant characters of the vessel elements were: simple perforation plates and partially pitted parietal thickening, found only in Alpinieae tribe, and 40 or more bars composing the plate in Elettariopsis curtisii, Renealmia chrysotricha, Zingiber spectabile, Z. officinale, Curcuma and Globba species. Vessel elements characters of 18 species of Alpinieae, Zingibereae and Globbeae were first described in this work.     

Structural and systematic study of an unusual tracheid type in cacti

Wide-band tracheids are a specialized tracheid type in which an annular or helical secondary wall projects deeply into the cell lumen. They are short, wide and spindle-shaped, and their bandlike secondary walls cover little of the primary wall, leaving most of it available for water diffusion. Wide-band tracheids appear to store and conduct water while preventing the spread of embolisms. They may be the most abundant tracheary element in the xylem, but they are always accompanied by at least a few vessels. Typically, fibers are absent wherever wide-band tracheids are present. Wide-band tracheids occur in the primary and secondary xylem of succulent stems, leaves and roots in genera of all three subfamilies of Cactaceae but were not found in the relictual genusPereskia, which lacks succulent tissues. In the large subfamily Cactoideae, wide-band tracheids occur only in derived members, and wide-band tracheids of North American Cactoideae are narrower and are aligned in a more orderly radial pattern than those of South American Cactoideae. Wide-band tracheids probably arose at least three times in Cactaceae.     

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。     

Water Flow Through Vessel Perforation Plates--A Fluid Mechanical Approach

The effect of scalariform perforation plates on the flow ofwater through plant vessels remains poorly understood. In thisstudy, a new computational tool based on finite element methodsolutions to the Navier-Stokes equation was applied to modellingfluid flow through these structures in plant vessels. Modelsdeveloped for Liriodendron tulipifera vessel elements were solvedfor cells with and without the perforation plate to study effectsof the plate on the pressure drop along the cell. Results indicatethat the pressure gradient was 5-fold greater through the platethan for regions before and after the plate. However, the perforationplate in this species accounts for only about 8% of the resistanceto flow through typical vessels because the plate influencesflow for only a short distance along the cell relative to itslength. Details of the flow characteristics through pores ofthe perforation plate are also described. Key words: Conductance, finite element method, perforation plate, vessel, water flow