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.     

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.     

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。     

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。     

慈姑不定根中导管分子的观察

慈姑导管仅在根中出现。根的后生木质部中央导管由顶端平截、单穿孔的网纹导管分子连接组成;周围较小的导管分子和管胞有从梯纹管胞向导管分子演化的各种过渡类型;有一至多个梯形穿孔或单穿孔发生在导管分子的端壁或侧壁,并有分枝型导管分子存在,特别在根与主茎连接处尤为明显。管胞亦有分枝与不分枝的类型。     

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。     

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。     

Functional anatomy and xylem cavitation resistance of three species of monocotyledons grown on flooded substrates

There are few investigations that analyze the xylem functional anatomy of monocotyledons, as the methods have been developed for woody plants. This study describes the root, rhizome and aerial stem xylem anatomy and functional anatomy of Canna indica, Cyperus papyrus and Phragmites communis grown on flooded substrates; and it aims to evaluate the relationship between the xylem anatomy and its cavitation resistance. To calculate the indexes of vulnerability, mesomorphy, collapse and relative hydraulic conductivity in the three organs mentioned, the diameter, number of vessels per mm², thickness of the walls and the length of the tracheal elements were recorded. In addition, the xylem specific conductivity of the aerial stem was measured with the pipette method, and its resistance to cavitation was determined experimentally by the air injection technique. The protoxylem is xeromorphic, it has longer vessel elements, smaller diameters, thin walls and scalariform perforation plates, whereas the metaxylem is mesomorphic, with shorter vessel elements, larger diameters, thicker walls and simple perforation plates. Both present low collapse resistance but have a high relative hydraulic conductivity. P. communis recorded the highest cavitation resistance, and the number of vessels per mm² was related to xylem cavitation resistance in Canna indica and Cyperus papyrus. The experimental results of this investigation match partially the anatomical indexes and showed that the xylem of these species has a low specific conductivity and is more vulnerable to cavitation than that of other monocots.     

细枝木麻黄导管分子及穿孔板解剖学研究

运用细胞图象分析系统和显微照相的方法对细枝木麻黄(CasuarinacunninghamianaMiq.)次生木质部导管分子进行了观察研究。细枝木麻黄次生木质部导管分子有3种类型,即两端具尾导管、一端具尾导管和无尾导管。导管分子穿孔板有4种类型:两端均为梯状穿孔板、一端为单穿孔板,另一端为梯状穿孔板、两端均为单穿孔板、一梯状穿孔板与一单穿孔板位于同一端壁两侧相互对应以及一些过渡类型穿孔板。根据观察结果,分析了各类型穿孔板之间的演化关系。     

余甘子次生木质部导管分子观察研究

运用细胞图象分析系统及显微照相的方法,对余甘子次生木质部导管分子进行观察研究.结果发现,余甘子次生木质部导管分子中存在着许多不同的样式,导管分子大多数具尾;其穿孔板存在着两种类型:(1)两端均为1个单穿孔板;(2)一端为1个单穿孔板另一端为2个单穿孔板;(3)极少数的导管分子具有特殊的内含物;(4)管间纹孔式为互列纹孔式;(5)导管射线间纹孔式为混合型纹孔与横列刻痕状纹孔以及梯状穿孔.     

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。     

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.     

基于不同方法的毛茛族(毛茛科)导管穿孔板比较研究

导管分子的演化与植物的进化相关联.毛茛科是联系基部被子植物和核心真双子叶植物的关键类群,对其导管穿孔板的研究具有重要的系统学意义.为研究毛茛科毛茛族不同属植物导管穿孔板类型及其与环境的相关性,该文利用新的扫描电子显微镜和半薄切片技术,对该族7属8种植物次生木质部导管状分子进行了比较观察.结果表明:木质部管状分子以单穿孔...     

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.     

不同纬度对鹅掌楸次生木质部结构的影响

对生长在北纬23至32度的鹅掌楸〖WTBX〗（Liriodendron chinense〖WTBZ〗 Sarg.）的次生木质部进行了比较解剖学研究。观察的特征有：导管分子的长度、直径、壁厚、梯状穿孔板横闩的数目;导管密度、纤维长度、直径、壁厚;射线的高度、宽度,单列射线的比例,射线的密度等。结果表明,随着纬度的增加,导管分子变短变细,壁变薄,梯状穿孔板的横闩数目减少;纤维也变短变细,壁变薄;木射线变矮变窄,单列射线的比例和射线密度增加。     

不同纬度对鹅掌楸次生木质部结构的影响

对生长在北纬23至32度的鹅掌楸（Liriodendron chinense Sarg.）的次生木质部进行了比较解剖学研究。观察的特征有：导管分子的长度、直径、壁厚、梯状穿孔板横闩的数目;导管密度、纤维长度、直径、壁厚;射线的高度、宽度,单列射线的比例,射线的密度等。结果表明,随着纬度的增加,管分子变短变细,壁变薄,梯状穿孔板的横闩数目减少;纤维也变短变线,壁变薄;木射线变敌变窄,单列射线的比例和射线密度增加。     

黄三七( 毛茛科) 茎次生木质部导管穿孔板的研究*

利用扫描电子显微镜对东亚特有植物黄三七( Souliea vaginata (Maxim. ) Franch. ) 茎的次生木质部
离析材料进行了观察, 结果表明, 黄三七茎次生木质部中的导管分子端壁上具网状穿孔板( 麻黄式穿孔
板) 、梯状穿孔板、网状- 梯状混合穿孔板、网状- 梯状- 单穿孔混合型穿孔板、梯状- 单穿孔混合型穿
孔板及单穿孔板, 同时也观察到了端壁多穿孔板和侧壁穿孔板, 并对不同类型穿孔板中纹孔膜的残留也进
行了观察。其中, 网状穿孔板、各种过渡类型的穿孔板均为毛茛科植物中首次报道。根据观察结果, 对导
管分子穿孔板的演化及黄三七属植物的系统位置进行了分析。     

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。     

Water Flow Through Vessel Perforation Plates--The Effects of Plate Angle and Thickness for Liriodendron tulipifera

The effects of scalariform perforation plate thickness and anglewithin vessel elements of Liriodendron tulipifera were studiedwith a computational fluid dynamical model. The pressure gradientand hence resistance to flow through the plate increased asthe perforation plate increased in thickness. Increasing theangle of the plate relative to the axis of the vessel (samenumber of pores) also increased the pressure gradient alongthe modelled cell. For the model matching the actual vesselelement, the plate contributed 8% to the flow resistance ofthe vessel element. This contribution increased only to 11%for doubled plate thickness, and to 14% for a plate at an angleof 60° to the vessel axis. The perforation plate alteredthe velocity profile across the vessel element, but to a differentextent depending on the angle of the plate. A plate at an acuteangle to the vessel axis has little effect on the paraboloidprofile as found upstream from the plate, while obtuse angleplates change the flow profile such that fluid through poresnear the wall is accelerated to a greater velocity than foundin the centre of the cell. Key words: Conductance, perforation plate, vessel, water flow