檀香吸器维管组织的个体发育

为了解檀香吸器维管组织的发育过程,采用激光共聚焦显微镜、光学显微镜和透射电镜观察檀香吸器维管组织的个体发育。结果表明,檀香维管组织的分化分为两个时期:入侵前和入侵后。吸器维管组织发育始于盘状吸器时期,起源于吸器基部具有分生能力的细胞,后分为两束。侵入前无向顶的分化,处于吸器基部。侵入后随吸管深入寄主根与寄主根维管束连通,形成具有吸收功能的维管组织。成熟吸器维管组织呈倒烧瓶结构,仅处于吸器烧瓶核心两边,由木质部组成而无韧皮部。檀香的吸器维管组织发育有两个因素诱导,一个是遗传因素,另一个为寄主。这些为檀香半寄生性特性研究提供了形态解剖学基础。     

银州柴胡根的发育解剖学研究

本研究采用常规石蜡切片结合荧光显微镜技术对银州柴胡根的发育解剖学进行了研究。结果表明:（1）银州柴胡根顶端分生组织由原分生组织及其衍生的初生分生组织组成。原生分生组织细胞体积小、排列紧密、细胞质浓厚、细胞核大而明显,具有典型的分生组织的特点;（2）初生分生组织由根冠原、表皮原、皮层原和中柱原组成。在根发育过程中,表皮、皮层和维管柱共同组成其初生结构。银州柴胡根初生木质部为二原型或三原型,外始式;同时在根表皮细胞的径向壁观察到径向壁的细胞壁加厚;（3）在根次生生长过程中,位于初生木质部和初生韧皮部之间的原形成层恢复分裂能力产生维管形成层,维管形成层不断地向外产生次生韧皮部,向内产生次生木质部;同时位于根内皮层内方的中柱鞘细胞恢复分裂能力产生木栓形成层,木栓形成层向外形成木栓层,向内形成栓内层。在维管形成层和木栓形成层分裂的过程中,在次生韧皮部和中柱鞘组织中产生形态大小不同的分泌道,均为次生的裂生型分泌道。研究认为,银州柴胡根的结构类似于药典收录的北柴胡和红柴胡根的结构特点,但其根表皮细胞径向壁加厚、木纤维的分布、分泌道的大小和数量等有别于柴胡属其它植物,可作为柴胡属植物重要的分类鉴定依据。     

Developmental Morphology of Roots and root-borne shoots ofPodostemum subulatum as compared withZeylanidium olivaceum (Podostemaceae—Podostemoideae) Part VII of the series ‘Morphology of Podostemaceae’

The root structure ofPodostemum subulatum is investigated and compared with that ofZeylanidium olivaceum. Podostemum has thread- or ribbon-like roots. The root tip consists of an inner apical meristem and a single-layered root cap. From roots arise numerous shoots of endogeneous origin. Their vascular bundle isab initio connected with the root bundle.By the simple (reduced) apical zonation, the roots ofPodostemum subulatum appear more advanced than the crustose roots ofZeylanidium olivaceum, which bear an ordinary (though asymmetrical) root cap. With regard to the endogeneous root-borne shoots, however,Zeylanidium appears more advanced because of the shoot dimorphism. The floriferous shoots have a short axis that grows plagiotropously above the crust surface, whereas the axes of the vegetative shoots are extremely short and remain, together with the apical meristem, within the crust. Only the leaves protrude from the crust surface.     

Studies on induced changes in growth patterns and polarity in roots of Vicia faba L.

Summary Roots of Vicia faba were treated with colchicine (0.025%), or IAA (4.7×10^-6 M), or both, for 3 hours and fixed at various intervals over the following 11 days. The axis of spindle orientation and the distribution of mitotic figures, lateral root primordia and xylem vessel elements was examined in the apical 10 mm of median longitudinal sections of these roots.No effect of IAA was found on the orientation of the spindle. However, evidence was obtained indicating that the systems controlling the polarity of cell division and cell expansion differ in some way.The number of lateral root primordia formed was greater in roots treated with IAA or colchicine than in control roots. These primordia were always initiated adjacent to a xylem vessel. Thus, no primordium was closer to the apex than the most apical xylem vessel, suggesting that an endogenous factor involved in primordia initiation is transported in the xylem. The primordia which develop after colchicine treatment grow out as lateral roots; this is in contrast with those which form after IAA treatment and which do not undergo elongation. These results, which it must be emphasized apply only to the apical 1 cm of treated roots, indicate that lateral root primordia become sensitive to IAA at a certain stage in their development. Exogenous IAA acts as an inhibitor.The new meristem, which forms in the primary root apex after colchicine treatment, contains both diploid and polyploid cells, i.e. it was formed from cells that were unaffected and from cells that were affected by colchicine. Following colchicine treatment the size of the meristem shrinks and this can be prevented by treatment with IAA. This and other evidence presented here, suggests that IAA is a factor involved in the control of the size of the apical meristem in normal roots.     

Changes in cell length and mitotic index in vascular pattern-related pericycle cell types along the apical meristem and elongation zone of the onion root

Summary Three pericycle cell types (opposite xylem, opposite phloem and intervening) distinguished by their location in relation to different elements of the vascular system were studied in the adventitious root ofAllium cepa L. Changes in cell length and mitotic index were analysed in these cells along the apical meristem and elongation zone of the root. The opposite phloem and intervening pericycle cells are significantly shorter than the opposite xylem pericycle cells in the apical half of the meristem. Between 1,200 and 1,400 m behind the tip, length became similar in all three pericycle cell types, while in more proximal zones the opposite phloem cells were significantly longer. These results suggest that the number of transverse divisions is different in the three types of pericycle cells. In the apical half of the meristem, mitotic index increased in intervening and opposite xylem cells but remained unchanged in opposite phloem cells, a fact likely to account for the relative lengthening of the latter. In the proximal half of the meristem, mitotic index fell in all three cell types until cell division had ceased. However, mitotic index in opposite xylem cells remained high for longer than in the other two cell types, implying that increase of the mean cell length in the former was slower. These results suggest that differences in mean cell length between the three pericycle cell types are due to different rates of proliferation.     

THE RELATIONSHIP BETWEEN THE PRIMARY THICKENING MERISTEM AND THE SECONDARY THICKENING MERISTEM IN YUCCA WHIPPLEI TORR. I. HISTOLOGY OF THE MATURE VEGETATIVE STEM

Anatomical observations were made on 1-, 2-, and 3-yr-old plants of Yucca whipplei Torr, ssp. percursa Haines grown from seed collected from a single parent in Refugio Canyon, Santa Barbara, California. The primary body of the vegetative stem consists of cortex and central cylinder with a central pith. Parenchyma cells in the ground tissue are arranged in anticlinal cell files continuous from beneath the leaf bases, through the cortex and central cylinder to the pith. Individual vascular bundles in the primary body have a collateral arrangement of xylem and phloem. The parenchyma cells of the ground tissue of the secondary body are also arranged in files continuous with those of the primary parenchyma. Secondary vascular bundles have an amphivasal arrangement and an undulating path with frequent anastomoses. Primary and secondary vascular bundles are longitudinally continuous. The primary thickening meristem (PTM) is longitudinally continuous with the secondary thickening meristem (STM). Axillary buds initiated during primary growth were observed in the leaf axils. The STM becomes more active prior to and during root initiation. Layers of secondary vascular bundles are associated with root formation.     

远志根的形态发生及组织化学研究

应用植物解剖学方法对远志(Potygda tenuiflia Willd.)根的发生和发育过程,以及1 a生与2 a生根的结构进行了比较观察,还应用组织化学方法对远志根储藏物质及主要药用成分积累部位进行了研究.结果表明:远志的药用部位为其主根,发育过程包括原分生组织、初生分生组织、初生结构和次生结构4个发育阶段.原分生组织来源于胚根,由3群原始细胞组成,具有典型分生组织的细胞学特征;初生分生组织包括根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和维管柱组成,初生木质部为二原型;次生生长主要是依靠维管形成层和木栓形成层的活动来完成.木栓形成层由中柱鞘细胞恢复分裂能力而形成,并且产生多层栓内层薄壁细胞.2 a生远志根的基本结构与1 a生的基本相同,只是栓内层增加至10层以上.远志根的储藏物质主要是脂类物质及少量的多糖.远志皂苷积累在远志根的薄壁细胞中,而山酮类化合物主要分布在根的木栓形成层、栓内层薄壁细胞和次生韧皮部中.     

The structure and development of the haustorium in parasitic Scrophulariaceae*

The structure and development of roots and haustoria in 37 species of parasitic Scrophulariaceae was studied using light microscopy. The mature haustorium consists of two regions: the swollen “body” and the parent root, which resembles non-haustorial roots in structure. The body arises from the parent root and is composed of an epidermis, cortex, central region of xylem (the vascular core), a region of parenchyma (the central parenchymatous core), and the portion of the haustorium contained in the host tissue (the endophyte). The xylem of the vascular core is composed predominately of vessel elements. The central parenchymatous core is composed of parenchyma and col-lenchyma. Vessels extend from the vascular core through the central parenchymatous core to the endophyte. The endophyte is composed of parenchyma cells and vessel elements. No phloem is present in the body of the haustorium. Early stages in the development of the haustorium are exogenous. Initial periclinal divisions in the epidermis or outer cortex are followed by hypertrophy of cortical parenchyma. These events are followed by development of the vascular core from the pericycle, attachment of haustorium to the host by a specialized layer of cementing cells or root hairs, and penetration of the host by dissolution of host cells.     

DISTRIBUTION AND RELATIONSHIP OF CELL DIVISION AND MATURATION EVENTS IN PISUM SATIVUM (FABACEAE) SEEDLING ROOTS

Pea roots have open apical organization, where discrete initial cells do not exist. Differentiation of all tissues occurs in cylinders and vascular sectors that blend gradually with each other. This study reports the distribution of dividing cells and their relationship to maturation events in the 2 mm root tip, and in the 8–10 and 18–20 mm segments. Up to 200 μm from the root body/cap junction, cell division is uniformly distributed throughout all meristem regions. By 350 to 500 μ, xylem tracheary elements and cells of the pith parenchyma and middle cortex have stopped dividing. At this level cell division is almost entirely restricted to two cylinders, one composed of the inner root cap, the epidermis, and the outer cortex (outer cortex cylinder) and another composed of cells of the inner cortex, the pericycle and vascular tissue (inner cortex cylinder). When the protophloem matures, all cells in the phloem sector of the inner cortex cylinder, including the 1 layered pericycle, the endodermis and the phloem parenchyma, stop dividing. The 3–4 layered pericycle in the xylem sectors continues dividing until about 10 mm from the body/cap junction following the maturation of the protoxylem tracheary elements.     

ANATOMY OF MACROZAMIA COMMUNIS LATERAL ROOTS AND ROOT NODULES FORMED IN VITRO STUDIED WITH LIGHT AND SCANNING ELECTRON MICROSCOPY

The anatomy of Macrozamia communis L. Johnson lateral roots and nodules was studied following axenic culture in light and darkness. Pointed lateral roots from dark cultures had an open apical organization similar to that of other cycads and gymnosperms. A distinct protoderm-derived epidermis was not observed. At the apex, the dermis was formed by the outer root capcortical cell layer. Subapically, the outer cortex formed the dermis. No evidence of an algal zone was observed in these roots. The stele was bounded by a distinct endodermis and contained an exarch, diarch xylem. Apogeotropic nodules which developed at the root-shoot junction in darkness, branched dichotomously and had rounded tips covered by tangentially-enlarged root cap cells. The root cap was reduced to a few cell layers and was confined to the extreme nodule apex. The central region of the apical meristem was enlarged, and meristematic cells contained differentiated amyloplasts. A presumptive algal zone was present in some but not all nodules and divided the cortex into inner and outer regions. Stelar anatomy was similar to that observed in pointed, dark-grown lateral roots, except that there was greater xylem differentiation. Nodules which developed in the light were similar to dark-formed nodules, except that root cap cells were radially enlarged and extended over the flanks of the nodule forming a persistent root cap. The heteromorphic lateral roots of M. communis formed a developmental continuum not a heterorhizic root system.     

Roles for Class III HD-Zip and KANADI genes in Arabidopsis root development

Meristems within the plant body differ in their structure and the patterns and identities of organs they produce. Despite these differences, it is becoming apparent that shoot and root apical and vascular meristems share significant gene expression patterns. Class III HD-Zip genes are required for the formation of a functional shoot apical meristem. In addition, Class III HD-Zip and KANADI genes function in patterning lateral organs and vascular bundles produced from the shoot apical and vascular meristems, respectively. We utilize both gain- and loss-of-function mutants and gene expression patterns to analyze the function of Class III HD-Zip and KANADI genes in Arabidopsis roots. Here we show that both Class III HD-Zip and KANADI genes play roles in the ontogeny of lateral roots and suggest that Class III HD-Zip gene activity is required for meristematic activity in the pericycle analogous to its requirement in the shoot apical meristem.     

Developmental anatomy of the root cortex of the basal monocotyledon, Acorus calamus (Acorales, Acoraceae)

BACKGROUND AND AIMS: The anatomical structure and development of adventitious roots were analysed in the basal monocotyledon, Acorus calamus, to determine to what extent those features are related to phylogenetic position. METHODS: Root specimens were harvested and sectioned, either with a hand microtome or freehand, at varying distances from the root tip and examined under the microscope using a variety of staining techniques. KEY RESULTS: Roots of Acorus calamus possess a unique set of developmental characteristics that produce some traits similar to those of another basal angiosperm group, Nymphaeales. The root apical meristem organization seems to be intermediate between that of a closed and an open monocotyledonous root apical meristem organization. The open-type root apical meristem consists of a curved zone of cortical initials and epidermal initials overlying the vascular cylinder initials; the epidermal part of the meristem varies in its association with the cortical initials and columellar initials of the promeristem. The cortex develops an endodermis with only Casparian bands, a dimorphic exodermis with Casparian bands and suberin lamellae, and a polygonal aerenchyma by differential expansion, as also observed in the Nymphaeales and some dicotyledonous species. The stele has characteristics like those of members of the Nymphaeaceae. CONCLUSIONS: Specific anatomical and developmental attributes of Acorus roots seem to be related to the phylogenetic position of this genus.     

Seedling developmental anatomy of an undescribed Malaccotristicha species (Podostemaceae, subfamily Tristichoideae) with implications for body plan evolution

The seedling development of an undescribed Malaccotristicha species was observed by using seedling culture and microtomy to infer the evolution of body plan with a focus on the root, which is a developmentally leading organ of most Podostemaceae. The young seedling has a small primary shoot apical meristem and a primary root apical meristem. The shoot meristem develops into a plumular ramulus, and the root meristem, into a cylindrical radicle with no root cap. The radicle transforms to a dorsiventral, flattened, capped primary root. An adventitious root develops endogenously on the lateral side of the hypocotyl and is similar to the primary root. This is a new pattern in Podostemaceae. Comparison of this and described patterns of Podostemaceae (and the sister-group Hypericaceae) suggests that the radicle was lost in the early evolution of Podostemaceae and instead adventitious roots replaced it as a leading organ.     

The extent and differences in mitotic activity of the root tip ofVicia faba L.

Mitotic activity does not stop for different meristematic cells of the root apex at the same distance from the initials. The differences are connected with the functional heterogeneity of the apical meristem of the root. The arrangement of vascular bundles,i.e. the alternation of independent xylem and phloem groups, is of major importance. In broad bean roots, the protophloem sieve elements stop dividing first. The centre of the stelei. e. late metaxylem elements stop dividing next. Division in the stele gradually ceases centrifugally, while it ceases centripetally in the peripheral part of the root. The cylindrical region with prolonged cell division includes internal layers of the cortex including endodermis, pericycle and adjoining cells of the stele. Proximally apical meristem is reduced to isolated strands of cells adjacent to the protoxylem poles. Pericycle cells stop dividing last at a distance of approx. 9–10 mm from the initials. The number of the division cycles is limited and is specific for individual cell types. Epidermal and cortical cells divide in broad bean roots transversely approximately seven times, cells of late metaxylem approximately five times. Root apical meristem is an asynchronous cell population with a different duration of the mitotic cycle. We determined local variations in the duration of the mitotic cycle in the apical meristem of broad bean root by means of colchicine-induced polyploidy. The cells of the quiescent centre had the longest mitotic cycle after colchicine treatment. The region of the proper root adjacent to the quiescent centre was mixoploid (2n and 4n). Isolated cells with a long cycle occurred also in the cortex and in the central cylinder. Cells with a division cycle of 18h were found in the root cap, in the epidermis, in the cortex and in the central cylinder. Relatively numerous cells with the shortest division cycle, approx. 12 h, occurred farther of the quiescent centre in the epidermis, in the cortex, in the pericycle, and in adjacent layers of the stele through-out the entire meristematic region. The results derived from the analysis of the apical meristem are discussed in connection with the ontogenesis of different types of cells taking part in the primary structure of the root.     

ADVENTITIOUS ROOT DEVELOPMENT IN TYPHA GLAUCA,WITH EMPHASIS ON THE CORTEX

Adventitious root development was investigated in Typha glauca plants grown under experimental conditions with the previous year's dead, sterile stalk either emerged above or submerged below the surface of Hoagland's solution. Adventitious roots emerged from buds in which most primordia had been earlier formed. Most roots elongated to 14–19 cm in 3–4 weeks and produced abundant lateral roots to their tips. Root apical meristem organization was typically monocotyledonous with a single tier of ground meristem/protoderm over the procambium. The ground meristem had zones of periclinal divisions in its innermost and outermost layers; the innermost layer initiated the endodermis and midcortex, and the outermost layers initiated the hypodermis. Crystalliferous cells with raphides were produced in the midcortex, and aerenchyma resulted from the radial expansion of schizogenous air spaces and some lysigeny in the midcortex. The procambium produced a vascular cylinder with 10–13 phloem and xylem poles, 6–9 large metaxylem elements, and central sclerenchyma. As roots stopped elongating, they narrowed, the vascular cylinder diminished in size, typical aerenchyma was lost from the cortex, crystal production ceased, and the rootcap diminished in size with its storage starch used up. Growth was determinate in these adventitious roots. The results suggested that a periclinally derived outer ground meristem was a prerequisite for a hypodermis, which, in turn, was necessary as a structural framework for aerenchyma. Without a hypodermis, typical aerenchyma was not present.     

Pericycle cell proliferation and lateral root initiation in Arabidopsis

In contrast with other cells generated by the root apical meristem in Arabidopsis, pericycle cells adjacent to the protoxylem poles of the vascular cylinder continue to cycle without interruption during passage through the elongation and differentiation zones. However, only some of the dividing pericycle cells are committed to the asymmetric, formative divisions that give rise to lateral root primordia (LRPs). This was demonstrated by direct observation and mapping of mitotic figures, cell-length measurements, and the histochemical analysis of a cyclin-GUS fusion protein in pericycle cells. The estimated duration of a pericycle cell cycle in the root apical meristem was similar to the interval between cell displacement from the meristem and the initiation of LRP formation. Developmentally controlled LRP initiation occurs early, 3 to 8 mm from the root tip. Thus the first growth control point in lateral root formation is defined by the initiation of primordia in stochastic patterns by cells passing through the elongation and young differentiation zones, up to where lateral roots begin to emerge from the primary root. Therefore, the first growth control point is not restricted to a narrow developmental window. We propose that late LRP initiation is developmentally unrelated to the root apical meristem and is operated by a second growth control point that can be activated by environmental cues. The observation that pericycle cells divide and lateral root primordia form without intervening mitotic quiescence suggests that lateral organ formation in roots and shoots might not be as fundamentally different as previously thought.     

PARASITISM IN PHOLISMA (LENNOACEAE). I. EXTERNAL MORPHOLOGY OF SUBTERRANEAN ORGANS

The external features of the subterranean organs of coastal Californian Pholisma depressum Greene are described. A dimorphic root system exists. Long, stout roots (pilot roots) serve for vegetative propagation in reaching other host roots. Short, unbranching, more slender roots originating from the pilot roots are haustorial in function. Shoots seem to arise only on pilot roots. Host attachment is achieved through the direct transformation of the apical meristem of haustorial roots into the haustorial organ. The basal part of the shoot is also able to produce roots when near host roots, and may form lateral branch shoots either directly or after new haustorial connections are formed. Both types of roots arise endogenously. The invaded host root undergoes considerable hypertrophy. The intrusive organ itself forms a massive but irregular body. More than one haustorial root may enter even a small host root. Self parasitism may occur but is not common.     

ORIGIN AND DEVELOPMENT OF LATERAL ROOTS IN TYPHA GLAUCA

Lateral roots of Typha glauca arose from the pericycle of the parent adventitious root. Periclinal divisions of the pericycle gave rise to two layers; the outermost initially produced the ground meristem and protoderm, and the innermost produced the procambium. The immature endodermis of the parent root contributed to the early stages of the root tip as an endodermal covering. Prior to emergence, the ground meristem/protoderm produced cells into the endodermal covering. After emergence, the endodermal covering was replaced by a calyptrogen, which was derived from the ground meristem/protoderm and which, in turn, formed the rootcap. A typical monocotyledonous three-tiered meristem was then produced. An outer ground meristem also arose before emergence to form a hypodermis in many lateral roots; in these, crystalliferous cell production began in midcortex cells before emergence, and a small aerenchyma developed in their cortices. The rootcap columella stored small amounts of starch shortly after emergence. Lateral roots of T. glauca were smaller than their parental adventitious roots; they normally had only two to six poles of xylem and phloem, and the cortex was less than six cells across. During 1–3-cm elongation, the lateral root apical meristem and mature regions narrowed, stored starch disappeared, fewer crystals formed, aerenchyma production ceased, and the roots stopped elongation.     

Effects of Zinc on Meristem Size and Proximity of Root Hairs and Xylem Elements to the Root Tip in a Zinc-tolerant and a Non-tolerant Cultivar of Festuca rubra L.

A 4 d exposure to zinc (0.1 and 02 µg Zn cm^–3) reducedthe length of the root apical meristem in a Zn-sensitive cultivar(S59) of Festuca rubra L. to a much greater extent than in aZn-tolerant cultivar (Merlin). In S59, Zn treatment also inducedroot hair and xylem formation much closer to the root cap boundarythan in control roots, whereas Merlin was only marginally affectedby Zn treatment. The data are discussed in relation to previouslyestablished effects of Zn on the cell cycle and other cellularcharacters of the two cultivars. zinc, meristem size, root hair, xylem, Festuca rubra     

远志根的发育解剖学研究

运用石蜡切片法对远志根的发育过程及1~3年生根的结构进行解剖学研究。结果显示:远志根的原分生组织由3群原始细胞组成,具有典型分生组织的细胞学特征。初生分生组织分化为根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和中柱组成,初生木质部为二原型。次生生长是依靠维管形成层和木栓形成层的活动完成,次生结构从外到内由周皮和次生维管组织组成;远志根次生结构特点为:次生韧皮部在次生维管组织中占主要部分,次生韧皮部中以韧皮薄壁细胞为主且其中储存有丰富的内含物,随着根龄的增加,韧皮薄壁细胞中的内含物也随之增加。3年生的主根中次生韧皮部薄壁细胞中的内含物最丰富;不同年份远志的主根随根龄的增加,周皮、次生韧皮部和次生木质部的面积都呈增加趋势,其中韧皮部和木质部的面积比值随根龄增长呈由小到大的变化,这是远志根的显著特点;根中的周皮发达,具有较厚的木栓层,次生木质部中导管和纤维发达,导管分布频率较高,并具有较大的口径。周皮和次生木质部的结构特征与远志的抗旱特性相适应。