杨柳科(Salicaceae)7种植物次生韧皮部的比较研究

本文研究和比较了杨柳科2属7种植物次生韧皮部解剖结构。结果表明:(1)杨属和柳属植物在次生初皮部解剖上有某些共同特征:次生韧皮部具有明显分层现象;韧皮纤维和含晶细胞与筛管分子、伴胞和韧皮薄壁组织细胞是切向带相间排列;筛管分子均为复筛板,端壁倾斜平均含有7-8个筛域。(2)两属植物在射线和晶体类型上有明显区别:柳属植物次生韧皮部无石细胞;杨属植物不具功能韧皮部中含有石细胞。(3)两属植物均有一些较为原始的韧皮部解剖特征。     

水松的次生韧皮部解剖及其系统位置的讨论

在光学显微镜和扫描电子显微镜下观察,水松茎次生韧皮部的主要特征为：韧皮部由轴向系统和径向系统组成。轴向系统由筛胞、韧皮薄壁组织细胞、蛋白细胞和韧皮纤维组成,径向系统由韧皮射线组成。在横切面上,轴向系统的各组成分子以单层切向带交替有规律的排列,其排列顺序为：筛胞－韧皮薄壁组织细胞－韧皮纤维-筛胞。筛胞的径向壁上嵌埋有草酸钙结晶,韧皮纤维仅一种类型,韧皮射线同型、单列。根据水松茎次生韧皮部的解剖研究,并与杉科其它各属的有关资料进行比较,我们认为：水松属与水杉属和落羽杉属有较近的亲缘关系。     

Anatomy of Secondary Phloem of Glyptostrobus in Relation to Its Systematic Position

A anatomical characters of secondary phloem in Glyptostrobus pensilis (Staunt.)Koch were observed by means of both light and scanning electron microscopy(SEM). The secondary phloem is composed of axial and radial systems. In the axial systems, the phloem consists of sieve cells, phloem parenchyma cells, albuminous cell and phloem fibers. In the radial systems, it consists of phloem rays. The alternate arrangement of different cells in cross section results in tangential bands. The sequence of radial arrangement follows the pattern of sieve cells, phloem parenchyma cells, sieve cells and phloem fibers, sieve cells. Many crystals of calbium oxalate are embedded in the radial walls of seive cells. The phloem fibers are of only one type. The phloem rays are homogeneous, uniseriate. According to the anatomical characters of secondary phloem of Glyptostrobus pensilis (Staunt.)Koch and comparison with the other genera of Taxodiaceae, Glyptostrobus, Metasequoia and Taxodium have close relationships.     

PHLOEM STRUCTURE IN THE CARBONIFEROUS FERN PSARONIUS (MARATTIALES)

Phloem anatomy in stems of Psaronius is described from coal ball specimens collected at the Berryville, IL and Lewis Creek, KY localities. Phloem completely surrounds the C-shaped xylem segments, but is more extensively developed on the abaxial side of the trace. The phloem zone consists of a central band of large diameter (approximately 90–120 μm) sieve elements surrounded by a mixed zone of smaller sieve elements and phloem parenchyma. Phloem is separated from the xylem by a parenchymatous xylem sheath. On the abaxial side of the trace, a discontinuous arc of very small diameter cells (7.8 μm) is present between the xylem sheath and the metaxylem. These cells corrrespond in position and size to protophloem cells in living marattialeans. Metaphloem sieve elements exhibit discrete, circular-oval sieve areas on their side and end walls, some of which show evidence of sieve pores. Phloem structure in Psaronius is compared with that known for living members of the Marattiales.     

Anatomical Studies on Secondary Phloem of Euonymus bungeanus

The anatomical structures, especially the type, distribution and arrangement of the constituent elements in the secondary phloem of Euonymus bungeanus Maxim. have been studied. The results showed that the secondary phloem was thicker, consisted of sieve-tube elements, companion ceils ,phloem parenchyma cells ,secretory ceils and rays. Sieve-tube elements, phloem parenchyma cells and secretory cells were alternately arranged in tangential bands, forming a conspicuous zone-like constitution. There was no obvious boundary between the functional phloem and the non-functional phloem. Sieve-tube elements were long, slender cells with very oblique end walls and compound sieve plates. Sieve areas on lateral wall were highly differentiated. Companion cells were triangular in transection and slender in radial section. Mostly,two or three companion cells stayed along with one sieve-tube element. In the functional phloem, phloem parenchyma cells were also slender, containing a few starch grains;but in the nonfunctional phloem they enlarged and contained abundant starch grains. Secretory cells were longer than sieve-tube elements, consisting of rubber-like material. Rays were uniseriate. Finally, the authors also discussed the phylogenetic position of E. bungeanus, which may provide some references for further study of the classification of different genera of Celastraceae.     

Seasonal Changes in the Structure of the Secondary Phloem of Grewia tiliaefolia, a Deciduous Tree from India

Structure of the secondary phloem of Grewia tillaefolia Roxb.was studied in samples of bark collected at monthly intervalsfrom forest populations of Gujarat in western India. The secondaryphloem in this species is vertically storied and the axial elementsoccur as alternate tangential bands of fibres and sieve elementsproduced in succession. On average, two to four bands of fibresand corresponding bands of sieve elements are produced in ayear. The sieve elements function for more than one season anddifferent phloem increments are separated by terminal zonesmade up of very narrow sieve elements which mature just beforeand immediately after the period of dormancy. The tree becomesleafless about eight to ten weeks preceding the spring equinox.Cambial activity, phloem differentiation and phloem functionare suspended during this period. Differentiation of phloembegins after bud break which occurs in April, and continuesuntil January, but most of the phloem is produced between Julyand September when the rainy season is well advanced. The widthof the conducting zone is maximal at the end of the period ofgrowth when the tree is in full leaf. Inactivation of sieveelements, apparently by callose plugging the sieve plates, beginswith leaf abscission. The sieve elements produced in the precedingseason, just before dormancy is imposed resume function in thefollowing growing season and the older elements die. Companioncells and axial parenchyma cells surrounding sieve elementsappear to have s significant role during senescence of the conductingelements. The development and activity of the secondary phloemseem to be related to other developmental phenomena occurringwithin the tree.     

THE PHLOEM OF ETAPTERIS LECLERCQII AND BOTRYOPTERIS TRIDENTATA

The phloem of Etapteris leclercqii and Botryopteris tridentata petioles is described from Lower Pennsylvanian coal balls. Petioles of B. tridentata are characterized in transverse section by an omega-shaped xylem trace, a phloem zone which extends from 2-10 cells in width, and 2-parted cortex. Etapteris leclercqii petioles exhibit a 4–9 cell-wide phloem zone surrounding the central clepsydroid xylem mass, and a 3-parted cortex. In both taxa a 1–2 cell layer parenchyma sheath separates the xylem from the extra-xylary tissues. The phloem of both species consists of sieve elements that average about 20 μm in diam by 200 μm in length in Botryopteris, and 100 μm in length in Etapteris, with horizontal-slightly oblique end walls. In transmitted light, the radial walls of the sieve elements form an irregular reticulate pattern enclosing elliptical lighter areas. With the scanning electron microscope, these areas appear as horizontal-slightly oblique furrows on the cell wall, with many small indentations lining the furrows. These indentations, because of their regular occurrence and size (from a few fractions of a micron up to 1.0 μm in diam), are interpreted as sieve pores, and the elliptical areas that enclose them as sieve areas. The phloem of E. leclercqii and B. tridentata is compared with that described for other fossil genera and with that of extant ferns.     

白蜡虫七种寄主植物枝条树皮比较解剖研究

采用常规石蜡切片法解剖观察了白蜡虫7种寄主植物一年生枝条树皮横切面结构特征,结果表明:白蜡虫7种寄主植物一年生枝条树皮从内到外由次生韧皮部、初生韧皮部纤维束、皮层和周皮组成;次生韧皮部横向系统均由筛管、伴胞和薄壁细胞组成;轴向系统由射线组成。木栓层以美国白蜡和流苏细胞层数最多,达10～12层;华南小蜡、紫药女贞和白枪杆次之,为5～8层;女贞和白蜡树最少,分别为2～3和3～4层。初生韧皮部纤维束排列整齐连接为带状或分散,女贞属纤维连接成带状,白蜡属和流苏属纤维分散。带状纤维层厚薄不均,厚度在26.93±13～59.15±7μm之间,以白枪杆纤维层最厚,为59.15±7μm;美洲白蜡次之,为50.05±7μm;白蜡树最薄,为26.93±13μm。分散型纤维束直径在25.12±13～76.15±36μm之间,纤维束直径大小顺序为:流苏(76.15±36μm)>紫药女贞(43.44±10μm)>女贞(25.12±13μm)。女贞、紫药女贞和流苏纤维束间距分别为78.53±39μm、149.78±27μm和212.02±95μm。次生韧皮部厚度在48.52±12～377.44±24μm之间,以女贞的次生韧皮部最厚,达377.44±24μm,华南小蜡最薄,为48.52±12μm。树皮次生韧皮部厚、木栓层数少和纤维束直径小为白蜡虫优良寄主植物的显著特征。     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

大关杨树皮的形成与季节变化的关系

大关杨的输导韧皮部限于当年新生的次生韧皮部。４月中旬至５月中旬大量出现,７月底停止发育。其成熟筛分子中具一细胞核,以后才消失。前一个生长季产生的筛管变形,故二个生长季的韧皮部蛤限明显。韧皮纤维出现迟,呈切向束状分布。含晶细胞分布于纤维束两侧。木栓形成层５月中旬开始活动,７月进入发育高峰。     

Wood Anatomy and the Development of Interxylary Phloem of Ipomoea hederifolia Linn. (Convolvulaceae)

In Ipomoea hederifolia Linn., stems increase in thickness by forming successive rings of cambia. With the increase in stem diameter, the first ring of cambium also gives rise to thin-walled parenchymatous islands along with thick-walled xylem derivatives to its inner side. The size of these islands increases (both radially and tangentially) gradually with the increase in stem diameter. In pencil-thick stems, that is, before the differentiation of a second ring of cambium, some of the parenchyma cells within these islands differentiate into interxylary phloem. Although all successive cambia forms secondary phloem continuously, simultaneous development of interxylary phloem was observed in the innermost successive ring of xylem. In the mature stems, thick-walled parenchyma cells formed at the beginning of secondary growth underwent dedifferentiation and led to the formation of phloem derivatives. Structurally, sieve tube elements showed both simple sieve plates on transverse to slightly oblique end walls and compound sieve plates on the oblique end walls with poorly developed lateral sieve areas. Isolated or groups of two to three sieve elements were noticed in the rays of secondary phloem. They possessed simple sieve plates with distinct companion cells at their corners. The length of these elements was more or less similar to that of ray parenchyma cells but their diameter was slightly less. Similarly, in the secondary xylem, perforated ray cells were noticed in the innermost xylem ring. They were larger than the adjacent ray cells and possessed oval to circular simple perforation plates. The structures of interxylary phloem, perforated ray cells, and ray sieve elements are described in detail.     

Sieve-element characters of Myristicaceae: Nuclear crystals, S-and P-type plastids, nacreous walls

The phloem of the Myristicaceae is composed of sieve elements, parenchymatous cells, and fibers. Within the metaphloem and secondary phloem parenchymatic layers including prominent secretory elements alternate with tangential bands of fibers and layers composed of sieve elements, companion cells and phloem-parenchyma cells. among the latter the sieve elements are most abundant and easily identified by the presence of thick (nacreous) walls. The most characteristic feature of the sieve elements of Myristicaceae (and found nowhere else among the Magnoliiflorae) are nuclear crystals, which are released into the lumen during nuclear degeneration and persist in the mature cell. P-and S-type sieve-element plastids were recorded for the 18 species investigated. Both types of the plastid are characterized by large diameters and many medium-sized starch grains. The sizes and contents (small protein crystals only) of the P-type plastids of the Myristicaceae do not conform to the tiny P-type plastids (with large protein crystals) of the Annonaceae, a family to which the Myristicaceae is traditionally allied.     

Seasonal Variations of Secondary Phloem Development in Pterocarya Stenoptera and Its Relation to Feeding of Kerria yunnanensis

Early in April of 1987, cells in an undifferentiated state which overwintered on the phloem side of the cambial zone in the branch of Pterocarya stenoptera began to differentiate into merebets of phloem. Cambium divided actively in mid-April and ceased to decide by early-Novembet. Five to eleven bands of fibers alternating with the bands of sieve tubes, companion cells and phloem parenchyma cells produced every year. By mid to late April, new xylem differentiation began. Phloem and xylem differentiation ceased almost simultaneously. Functional sieve tube elements were present all the year round in the phloem. During winter, most sieve tubes produced in the current year ceased functioning, leaving only the zone of functional sieve tube of several rows of cells in width with open pores in the sieve plates. These sieve tubes did not collapse until mid-May. In October, several rows of partially differentiated sieve elements appeared near the cambial zone. They still possessed nuclei. The companion cells had produced but no P-protein. They matured during April of the following year and collapsed by July to September. The life span of sieve elements extended for 8 months at the most. In winter, there were less functional sieve tubes in the branch. This may be one of the reasons that only few Kerria yunnanensis survive on the branch of Pterocarya stenoptera.     

CO2浓度加倍对辽东栎维管组织结构的影响

 以辽东栎(Quercus liaotungensis)的13年生幼树为材料,分别培养在大气CO2浓度加倍(700μl·L-1)与对照(350μl·L-1)的开顶式熏气室中,研究CO2浓度升高对其茎次生木质部和次生韧皮部结构的影响。结果表明：经CO2浓度加倍处理的两个生长季内,辽东栎的年轮宽度明显增加,为对照的300％～370％,其中晚材宽度的增加更为显著,为对照的750％～830％。另外,晚材中导管的密度和径向直径分别比对照增加50％和20％;木纤维细胞的比例约为对照的170％。但早材的导管分子和木纤维细胞与对照相比均无显著变化。在CO2浓度加倍条件下,辽东栎的次生韧皮部中含晶韧皮薄壁细胞的数目,每条韧皮纤维切向带中韧皮纤维细胞的数目,以及韧皮纤维长度均有显著增加(p≤0.05)。相反地,韧皮纤维细胞的直径和筛管分子长度却无明显变化。值得提出的是,在CO2浓度加倍的条件下,次生韧皮部的宽度、筛管分子的直径、以及每年形成的韧皮部细胞总数分别为对照的82％、87％和80％。综上所述,大气CO2浓度加倍对辽东栎次生木质部的生长发育具明显的正效应,而对次生韧皮部的细胞总数与筛管分子的影响则呈负效应。     

HYPERPLASTIC PHLOEM IN SUGARBEET LEAVES INFECTED WITH THE BEET CURLY TOP VIRUS

Electron microscopy of sugarbeet leaves infected with the beet curly top virus confirmed earlier findings by light microscopy that the hyperplastic phloem consists mainly of sieve elements that are more or less abnormal in structure. Some parenchyma cells and occasional companion cells may be present. The hyperplastic phloem develops in the place of normal phloem and sometimes in the adjacent ground tissue and the xylem. The sieve elements vary in shape and may be haphazardly arranged. The protoplasts of the sieve elements have the usual characteristics of this type of cell. The sieve element plastids develop from chloroplasts if the hyperplasia occurs in chloroplast-containing parenchyma cells. The cell walls have sieve areas that often are less well differentiated than those of normal sieve elements. The hyperplastic growth in the phloem of curly top diseased plants is discussed with reference to plant tumors induced by certain other plant viruses.     

PHLOEM ANATOMY OF THE CARBONIFEROUS COENOPTERID FERNS ANACHOROPTERIS AND ANKYROPTERIS

Phloem histology in the petioles of two genera of Pennsylvanian ferns is detailed from coal balls collected at various localities in North America. Both Ankyropteris and Anachoropteris have primary phloem that completely surrounds the central xylem trace and is separated from it by a parenchymatous sheath. Ankyropteris contains very narrow (about 13.5 μm diam) sieve elements and a few strands of phloem parenchyma. End walls are either horizontal or slightly oblique and sieve areas as well as scattered individual pores have been observed. Anachoropteris phloem contains two different sizes of sieve elements. Small sieve elements that surround the C-shaped trace are similar to those seen in Ankyropteris. Larger elements (approximately 50–120 μm in diam) are present only within the C-shaped trace, and are elongate (up to 2.5 mm) with very oblique end walls. Sieve areas on these large cells are conspicuous, 5–8.5 μm in diam and aggregated into groups. The cell wall within each sieve area appears to be composed of criss-crossed fibrillar material. Phloem anatomy in these two ferns is compared to that previously described in other Carboniferous vascular cryptogams, as well as that known from extant plants.     

PHLOEM ANATOMY IN STAUROPTERIS BISERIATA FROM THE PENNSYLVANIAN OF NORTH AMERICA

Phloem anatomy in the coenopterid fern Stauropteris biseriata is detailed from Lower-Middle Pennsylvanian coal ball specimens from eastern Kentucky. Axes exhibit a cruciate-shaped xylem trace in transverse section. Phloem tissue completely surrounds the xylem, but is more extensively developed in the embayments between the xylem arms. Phloem is composed of elongate conducting elements with a few scattered parenchyma cells. Large and small sieve cells are present, with larger ones occurring in the embayments within the primary plane of symmetry of the axes. Large elements are approximately twice the diameter of the smaller sieve elements. Oval sieve areas and pores have been observed on lateral and oblique end walls of both large and small elements. The structure and composition of Stauropteris phloem is discussed in relationship to the available information on phloem anatomy in other fossil cryptogams.     

Anatomy of the Secondary Phloem and the Crystalliferous Phloem Fibers in the Stem of Torreya grandis

The structure of the secondary phloem and the development of the crystaleiferous phloem fibers in the stem of Torrey grandis were observed under the ligth microscope and SEM. The secondary phloem is composed of sieve cells, phloem parenchyma cells, crystalliferous phloem fibers and stone cells in the longitudinal system, and the uniserite homogeneous phloem rays consisting of parenchyma cells only in the radial system. In the cross section, there are 3–9 sieve cells in radial rows forming discontinuous tangential layers, the crystalliferous phloem fibers often in a single discontinuous tangential layer and the stone cells dispersed in rangential layer of phloem parenchyma. The developmental process of crystalliferous phloem fibers is as follows: initial cells appeared in the end of April and were well differentiated in the first week of May. Some crystals were deposited in the primary wall, while others were free in the cell. At the end of May, the secondary wall of most crysalliferous phloem fibers started to be thickened. With the thickening of the secondary wall, all the crystals were embedded in the wall from June to August From the end of September to the early days of October, the crystalliferous phloem fibers reached their full maturation. It is shown by microchemical identification and EDAX analysis that the crystals embedded in the wail of crystalliferous phloem fibers are calcium oxalate crystals.     

OBSERVATIONS ON METAPHLOEM IN THE VEGETATIVE PARTS OF PALMS

Metaphloem was studied in available vegetative parts of 374 species in 164 genera of palms. Sieve elements usually have compound sieve plates except in the subfamilies Lepidocaryoideae and Nypoideae. Sieve elements in roots usually have oblique to very oblique end walls, whereas in stems and leaves they have transverse to oblique walls. Within a phloem strand the degree of compounding of a sieve plate is directly correlated with element diameter. Plastids are normally present in functioning, enucleate sieve elements. Small quantities of “slime” substances have been detected in young sieve elements in stems and petioles of a few species. Many sieve plates in functioning sieve elements lacked callose in materials quick-killed in liquid nitrogen or chilled acetic-alcohol. Definitive callose is confined to sieve elements just before their obliteration. Sieve tubes in leaf and stem are usually ensheathed by contiguous parenchyma cells while those in root have very few contiguous parenchyma cells. Two types of contiguous parenchyma cells can be distinguished by difference in cytoplasmic density, especially with the electron microscope. Cells with denser cytoplasm are interpreted as companion cells. Lignified contiguous parenchyma cells are occasionally present in metaphloem of petioles. The possible diagnostic and taxonomic features of metaphloem are discussed.