Function of transfer cells in the nodal regions of stems,particularly in relation to the nutrition of young seedlings

Summary The distribution and time course of development of transfer cells in the hypocotyl region of lettuce (Lactuca sativa L.) and groundsel (Senecio vulgaris L.) are examined by light microscopy of serial sections through a sequence of ages of hypocotyls. Investments of xylem transfer cells occur in departing traces to the cotyledons and, later, in the traces to foliage leaves; phloem transfer cells are widely distributed but particularly prominent in those bands of protophloem in the plumule vasculature which lie alongside xylem of the cotyledonary traces. Both classes of transfer cell are well endowed with wall ingrowths before differentiation of xylem and perforation of stomata occurs in the plumule. Autoradiographic evidence is obtained of a transport pathway from cotyledonary trace xylem elements to xylem transfer cell to plumule, and analyses of xylem sap collected from above or below the zones of transfer cells in the hypocotyl show that certain materials can be removed from the xylem sap by transfer cells as it moves towards the cotyledons. From these findings it is concluded that the seedling transfer cells play an important role in nutrition of the young plumule, particularly before the latter has become adequately connected with the vascular systems of cotyledons and root.Experiments on the experimental modification of transfer cell development in the hypocotyl suggest that both photosynthetic fixation of carbon dioxide and a transpirational loss of water by a cotyledon must take place before the presumptive xylem transfer cells in its traces can develop normal sets of wall ingrowths.Discussion is extended to the general role of transfer cells in the nodal regions of stems. Possible functions envisaged are, the general nutrition of young tissues of the apical region, the abstraction of assimilates for local storage, the transfer of assimilates to axillary buds released from apical dominance, and the interchange of assimilates between adjacent vascular traces running through the node.     

DEVELOPMENTAL MORPHOLOGY OF THE EMBRYO AND SEEDLING OF RHIZOPHORA MANGLE L. (RHIZOPHORACEAE)

The embryo of Rhizophora mangle L. is initially attached to the integument by a long multiseriate suspensor. Its basal cells lyse, and intrusive growth of the endosperm envelops the embryo, forces the micropyle open, and often carries the embryo out of the integument. Thus, “germination” is effected by growth of the endosperm rather than of the embryo. The surface of the endosperm differentiates into a layer of peculiar transfer cells. The cotyledonary body initiates as a toroidal primordium, which later becomes lobed; most of the free portions ultimately fuse. After “germination,” the axis of the viviparous seedling grows by a diffuse intercalary meristem below the cotyledonary node. Before seedling abscission, the shoot apex produces three pairs of leaves, the first of which aborts, leaving the rest of the plumule protected by their stipules. The (immersed) radicle apex is nearly inactive, but lateral roots arise early in seedling development; these are usually the first or only roots to grow during establishment. Ten provascular strands “differentiate” in the cotyledons; a hollow provascular cylinder develops in the hypocotyl. Initial vascular differentiation in the latter is of many alternate poles of xylem and phloem; later, de novo differentiation of metaxylem opposite the protophloem poles, and vice versa, produces collateral bundles. Xylem maturation is endarch over most of the length of the hypocotyl, but tangential and random series of metaxylem vessels occur in the radicle end.     

THE DEVELOPMENTAL ANATOMY OF OPUNTIA BASILARIS. I. EMBRYO,ROOT, AND TRANSITION ZONE

The large seeds of Opuntia basilaris Engelm. & Bigel. show an unusually high percentage of germination, followed by a rapid development of the seedling during the first 30 days of growth. The primary root has six xylem arms alternating with six phloem poles around a large central pith. Development of metaxylem opposite each of the primary phloem poles results in the formation of eight collateral bundles. Secondary and tertiary roots have four xylem and phloem poles with xylem developing to the center of the stele. The transition zone is characterized by a gradual disappearance of all but two of the primary xylem arms of the root. Metaxylem development in the central portion of the transition zone interconnects the protoxylem poles forming a primary xylem cylinder around the central pith. The collateral bundles pass through the transition zone essentially without change.     

STRUCTURALLY PRESERVED FOSSIL PLANTS FROM ANTARCTICA. I. ANTARCTICYCAS,GEN. NOV., A TRIASSIC CYCAD STEM FROM THE BEARDMORE GLACIER AREA

Silicified stems with typical cycadalean anatomy are described from specimens collected from the Fremouw Formation (Triassic) in the Transantarctic Mountains of Antarctica. Axes are slender with a large parenchymatous pith and cortex separated by a narrow ring of vascular tissue. Mucilage canals are present in both pith and cortex. Vascular tissue consists of endarch primary xylem, a narrow band of secondary xylem tracheids, cambial zone, and region of secondary phloem. Vascular bundles contain uni- to triseriate rays with larger rays up to 2 mm wide separating the individual bundles. Pitting on primary xylem elements ranges from helical to scalariform; secondary xylem tracheids exhibit alternate circular bordered pits. Traces, often accompanied by a mucilage canal, extend out through the large rays into the cortex where some assume a girdling configuration. A zone of periderm is present at the periphery of the stem. Large and small roots are attached to the stem and are conspicuous in the surrounding matrix. The anatomy of the Antarctic cycad is compared with that of other fossil and extant cycadalean stems.     

夏侧金盏花幼苗初生维管系统的解剖学研究

夏侧金盏花(Adonis estivalis L.)为毛莨科(Ranuncula-ceae)侧金盏花属(Adonis)植物。其幼苗可明显地分为上胚轴苗区、子叶节区和下胚轴根区。幼苗以子叶节区为中心,往上其子叶节区中部和上部为子叶节—茎过渡区;向下其下胚轴为子叶节—根过渡区。目前对毛莨科某些属幼苗初生维管系统的个体发育研究已有一些报道。但夏侧金盏花幼苗的子叶节—茎过渡区的转变与已报道的其他属均不同。主要表现为其子叶节区下部的中始式二原型双肩状单中柱,到达子叶节区中部,其后生木质部弦向发育成二唇形外韧维管束雏型,在中柱中央出现薄壁组织,进一步发育则形成髓;再往上,即子叶节区上部,便一分为多个内始式的外韧维管束雏型,直接形成上胚轴(茎)的真中柱。此研究为再一次验证子叶节区理论的正确性与进一步揭示被子植物初生维管系统的演化规律积累一份新资料。     

Anatomical Studies on Hypocotyl of Circaeaster

The stem of Circaeaster agrestis Maxim. is very short but the length of hypocotyl is comparatively long, almost occupying the whole length of the plant. This tender hypocotyl is mainly supported by the thickening of cuticle on the outer wall of the epidermal cell and the primary xylem in the center. Between primary xylem and primary phloem there are 2–3 layers of parenchymatous cells, regularly or irregularly arranged, but no cambial zone can be recognized. The transition region where root and stem meet showed no evidence of twisting, splitting or inversion of the strands in the primary vascular tissues which are common in most of the dicots. The extending cotyledon traces differentiate directly from the parenchymatous cells which locate on the outside of the poles of primary xylem. The first and the second leaf traces are organized in the middle of the primary phloem.     

PROCAMBIUM VS. CAMBIUM AND PROTOXYLEM VS. METAXYLEM IN POPULUS DELTOIDES SEEDLINGS

The concept of a procambium-cambium continuum was examined in Populus deltoides by following its development in serially sectioned bud and stem tissues. As in other species, the term cambium is used to refer to that part of the continuum associated with the formation of secondary vascular tissues; i.e., with secondary growth. However, that part of the continuum associated with the formation of primary vascular tissues is subdivided to facilitate interpretation of the consecutive stages of primary xylem differentiation. Thus, the procambium as envisioned by other authors is subdivided into procambium, initiating layer, and metacambium, all of which develop acropetally and in complete continuity. The procambium is derived from the residual meristem in the form of acropetally developing strands and traces. The initiating layer is represented by the first, tangentially separated, periclinal divisions that delineate the position of the prospective cambium. The metacambium is a later stage during which additional periclinally dividing cells unite the initiating layer into a tangentially continuous meristem within a trace bundle. After establishment of the initiating layer, the procambial trace is completely phloem dominated. Protoxylem differentiation begins in an originating center at the base of the leaf primordium and it progresses basipetally to form the protoxylem pole. Cells of the initiating layer do not contribute to the formation of either protoxylem or protophloem. However, those cells of the initiating layer directly opposite the protoxylem pole divide precociously and later differentiate to metaxylem, thus forming a radial file of protoxylem-metaxylem elements. Protoxylem elements of lateral traces are longitudinally continuous with the protoxylem of their parent traces, whereas those of a central trace are longitudinally continuous with the metaxylem of its parent trace. Metaxylem is formed later than protoxylem and it is derived from the metacambium. Metaxylem does not form a continuous system with protoxylem of the same trace because of the different temporal and spatial origins of the two kinds of xylem. Rather, metaxylem is longitudinally continuous with secondary xylem of older traces below. An attempt was made to determine the functional significance of the pattern of protoxylem and metaxylem differentiation in relation to primary and secondary plant development.     

Ultrastructural Localization of a Bean Glycine-Rich Protein in Unlignified Primary Walls of Protoxylem Cells

A polyclonal antibody was used to localize a glycine-rich cell wall protein (GRP 1.8) in French bean hypocotyls with the indirect immunogold method. GRP 1.8 could be localized mainly in the unlignified primary cell walls of the oldest protoxylem elements and also in cell corners of both proto- and metaxylem elements. In addition, GRP 1.8 was detected in phloem using tissue printing. The labeled primary walls of dead protoxylem cells showed a characteristically dispersed ultrastructure, resulting from the action of hydrolases during the final steps of cell maturation and from mechanical stress due to hypocotyl growth. Primary walls of living protoxylem and adjacent parenchyma cells were only weakly labeled. This was true also for the secondary walls of proto- and metaxylem cells, which in addition showed high background labeling. Inhibition of lignification with a specific and potent inhibitor of phenylalanine ammonia-lyase did not lead to enhanced labeling of secondary walls, showing that lignin does not mask the presence of GRP 1.8 in these walls. Dictyosomes of living proto- and metaxylem cells were not labeled, but dictyosomes of xylem parenchyma cells without secondary walls, adjacent to strongly labeled protoxylem elements, were clearly labeled. These observations suggest that GRP 1.8 is not produced by xylem vessels but by xylem parenchyma cells that export the protein to the wall of protoxylem vessels.     

扁圆封印木（相似种）茎干的解剖特征

贵州省水城矿区晚二叠世煤核中扁圆封印木（相似种Ｓｉｇｉｌｌａｒｉａ ｃｆ．ｂｒａｒｄｉｉＢｒｏｎｇｎ．）茎干的主要解剖特征如下：管状中柱,具多边形薄壁细胞组成的髓。初生木质部成环带状,外缘呈规则的齿槽状,向心式发育。次生木质部显束状特征,横切面管胞为方圆至长方形,纵切面为梯状壁增厚,并具流苏纹。射线１—２列细胞宽,数个至十余个细胞高。叶迹起源于初生木质部外缘的槽中,中始式,但以向心发育为主。     

SECRETORY RESERVOIRS (DUCTS) OF TWO KINDS IN GIANT RAGWEED (AMBROSIA TRIFIDA; ASTERACEAE)

Two types of tubular secretory reservoirs occur in Ambrosia trifida, the first such example known in plants. Paraffin and resin sections, and clearings showed that, although each type consists of many separate unbranched tubes, they differ in anatomy, secretory contents, distribution, and length. Reservoirs (PAR) containing a red substance (presumably a polyacetylene) and lined with a biseriate epithelium parallel the largest leaf and stem vascular bundles. One PAR arises near the base of each leaf lobe midrib and extends through the petiole to the node or continues in the stem cortex to the node below. Other PARs start at the cotyledonary node or in cotyledons and extend down into the primary root, where they have only a single layer of unspecialized epithelium. PARs realign themselves, and more form de novo, until the primary root has two to four separate arrays of PARs abutting the endodermis, each with three to six parallel PARs. Branch roots have similar PAR arrays but unconnected to PARs of the parent root. Inflorescence PARs occur only in bracts, and in petals of male flowers. The second type of reservoir (OR) has a uniseriate epithelium and contains an unidentified oil. ORs occur in phloem, and in pith next to xylem, of stem and large leaf bundles. They dwindle in successively smaller veins until the two smallest orders lack them. ORs occur only in phloem in the hypocotyl; none occur in cotyledons, roots, or floral parts.     

Ontogeny of the vascular system ofBotrychium multifidum (S. G. Gmelin) Rupr.(Ophioglossaceae) and its bearing on stellar theories

Basipetal to the shoot apex, a procambial ring with parenchymatous gaps is present. The protoxylem poles are endarrh in both the ectophloic siphonostele and the collateral vascular bundle which comprises the leaf trace. Each leaf trace has an anastomosing system of protoxylem poles that decreases in number basipetally from five to three to two. Differentiation of the leaf trace procambium and protoxylem is bidirectional, that is the differentiation first occurs near the base of the leaf and acropetally in the leaf and basipetally in the stem. Then a fascicular cambium differentiates betweem the primary xylem and phloem in the leaf. This vascular cambium which is also present in the stem is unidirectional and only produces secondary xylem centripetally. Limited secondary growth also occurs in roots. Medullary tracheids when present are longitudinally continuous with the vascular system. The stele of the stem is interpretated as a sympodium of leaf traces and the pith is considered to be fundamental tissue enclosed by the anastomosing of leaf traces.     

Empirical Models for Xylogenesis in Populus deltoides

Empirical quantitative models were constructed for Populus deltoidesdescribing temporal and spatial changes in vessel characteristicsof metaxylem, both within individual central leaf traces andwithin all central leaf traces considered as a morphologicalunit at a given transverse level in the stem (the central tracesympodia). Similar models were constructed for secondary vesselcharacteristics. The growth processes of the stem segment throughwhich the vasculature extended were incorporated in these modelsto illustrate how a functional vascular system is maintainedin the stem as a whole. The central trace sympodia representedthe integrals of the temporal and spatial functions for individualcentral leaf traces. Metaxylem vessel production ceased in individualleaf traces two plastochrons before the cessation was reflectedin the central trace sympodia because of the integrative natureof the sympodial complex. A functional continuum of developmentwas apparent between metaxylem vessels of the central tracesympodia and secondary vessels of the stem. The transition betweenmetaxylem and secondary xylem production in the central tracesympodia corresponded with cessation of leaf and internode elongation. Populus deltoides Bartr. ex Marsh., cottonwood, primary xylem, secondary xylem, primary-secondary vascular transition, leaf growth, xylogenesis     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND SECONDARY ROOT OF GLYCINE MAX (L.) MERR.

The primary vascular connection between primary and secondary root of Glycine max (L.) Merr. was derived from stelar parenchyma and pericycle. Inner stelar parenchyma, associated with the parent metaxylem and outer stelar parenchyma adjacent to the pericycle, were resonsible for the histogenesis of the primary xylem connection. Acropetal maturation of the diarch xylem connection occurred after the lateral root emerged from the parent root. Development of tetrarchy occurred distal to the diarch xylem connection. The concentric primary phloem connection was derived from the pericycle and outer stelar parenchyma. Acropetal maturation of the primary phloem connection occurred prior to lateral root emergence from the parent root. Secondary growth quickly augmented the primary vascular connection. A substantial amount of mature secondary xylem formed prior to maturation of the secondary phloem. The structure of the primary and secondary vascular connections is described.     

Mechanisms of Reduction of the Stelar Pattern along Barley Roots

The stelar pattern along the seminal and nodal roots of barley (Hordeum vulgare L.) is gradually simplified due to a decreasing frequency of longitudinal cell division in the apical meristem. The decrease involves the proportion of stelar parenchyma, the number of vascular strands on the periphery of the stele and, in nodal roots with a more complex structure, the number of central metaxylem files. In spite of the fact that the stelar parenchyma is reduced in distal parts of the roots to approximately one half, the discontinuity of central and peripheral metaxylem is preserved. Reduction of the number of central metaxylem files is due to fusion. In the reduction of peripheral xylem and phloem strands, the development of certain xylem strands is discontinued and they are terminated blindly. Two phloem strands that had alternated radially with them, approach each other, coalesce and a single phloem strand continues to develop. In this way the regular alternation of phloem and xylem is re-established. The importance of fusions ensuring reduction of the functional continuity in vascular tissue by formation of a network structure must be stressed. This reduction mechanism is involved not only in files of the wide central metaxylem but also in phloem strands which are thus preferred over blindly terminating peripheral xylem strands.     

Gymnosperm trees from the Permian of Antarctica: An anatomically preserved trunk of Kaokoxylon sp.

Anatomically preserved gymnosperm axes are relatively abundant in Permian localities of Antarctica, but their anatomy has rarely been studied in detail, which limits comparison with other Gondwanan morphotaxa. Here we describe a silicified trunk collected from the Upper Permian Buckley Formation at Coalsack Bluff, in the central Transantarctic Mountains. The trunk has a small heterogeneous pith approximately 4 mm in diameter containing conspicuous sclerotic nests, endarch primary xylem maturation, paired leaf traces, and secondary xylem of the Araucarioxylon type. Comparison with contemporaneous gymnosperm axes from Antarctica indicates that the Coalsack Bluff trunk represents a new Permian morphotaxon for the region. The anatomical characters of the pith and secondary xylem suggest an affinity with the genus Kaokoxylon Kräusel, previously reported from Permian and Triassic localities of Southern Africa, South America, India, and Australia.     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND LATERAL ROOT OF LYCOPERSICON ESCULENTUM

The primary xylem connection between the diarch parent root and the diarch lateral root was derived from the pericycle and stelar parenchyma. Early in lateral root development stelar parenchyma that was positioned between the parent xylem and the primordium divided transversely. These transverse divisions produced a plate of cells, most of which subsequently differentiated into vessel element connectors. After emergence of the lateral root, xylem maturation began in the stelar vessel element connectors and maturation proceeded acropetally into the lateral root. Protoxylem of the lateral root was connected to the metaxylem of the parent root via stelar vessel element connectors. The circular phloem connection was pericyclic in origin. Axial phloem connections which vascularized the lateral root were established with sieve tube elements of both parent phloem poles. Maturation of the phloem connection occurred prior to lateral root emergence. Transaxial phloem, positioned in arches above and below the lateral root vascular cylinder, was derived from the pericycle; and each arch consisted of three to four sieve tube elements. No transfer cells were found in the transaxial phloem.     

Anatomical Characteristics of the Stem of Sigillaria cf. brardii from Coal Balls in Guizhou Province,China

The stem specimens of Sigillaria cf. brardii were collected from the coal balls of Upper Permian in Shuicheng Coal Mines in Guizhou Province. The main anatomical characteristics of Sigillaria cf. brardii are described as follows: The stem is siphonostelic, with pith composed entirely of polygonal parenchyma cells, there are secondary walls in some pith cell cavities these secondary walls show the characters of cell division. Surrounding the pith is the continuous cylindrical primary xylem which consists entirely of tracheids. The outermost, and part are the protoxylem elements show spiral secondary thickenings. In cross section, the outer edge of exarch primary xylem appears regularly sinuous, with trace of mesarch leaf originating from the furrows. The centripetal metaxylem is characterized by scalariform wall thickenings on the tracheids, and delicated strands of secondary wall materials extending between abjacent bars, these structures are called fimbris, or williamson striations, and are characteristic in lepidodendrids. The secondary xylem consists of tracheids and vascular rays. The tracheids, too, have scalariform wall thickenings and fimbris. The rays are one-to twocell width and several to more than ten cells in height.     

Palaeosmunda Emended and Two New Species

The genus Palaeosmunda was established by R. E. Gould in 1970 based upon some Late Permian Osmundaceous trunks with well-developed leaf gaps and rhomboidal sclerotic ring within petiolar base seen in cross section. As he thinks that the latter character is more important than the former, this genus could not be assigned to any subfamily of Osmundaceae. However, the leaf gap is one of the most important characters in the structure of the fern stem, so the author suggests that this genus should be assigned to subfamily Osmundoideae and its diagnosis must be emended as follows: The genus Palaeosmunda is represented by some rhizomes (or trunks), roots and leaf bases of ferns which structurally are preserved, resembling Osmundacaulis but which can’t be assigned to any group of this genus. Stem containing an ectophloic dictyoxylic siphonostele; if tracheids present in the pith, they being multiseriate scalariform pitted; pith or cortex sometimes contain ing groups of secretory cells or sclerenchyma; number of leaf traces seen in a tran sverse section of cortex more than 30; leaf traces adaxially curvature, rarely oblong shaped; petiolar bases with or without stipular expansion, containing a C-shaped vascular strand; root diarch. Type species——Palaeosmunda williamsii. According to this diagnosis some primitive osmundaceous species with the leaf gaps, which have already found in Upper Permian and Lower Triassic, could be assigned to this genus. Two of them are P. williamsii Gould and P. playfordii Gould, and Osmundacaulis beardmorensis, which was from Lower Triassic of Antarctica in 1978, should be assigned to the genus Palaeosmunda. In this paper two osmundaceous new species: P. primitiva and P. plenasioides were found in the coal balls of Upper Permian age from Wangjiazhai of Shuicheng of Guizhou Province, China. P. primitiva is represented by two trunks; stem about 4 cm in diameter; stele actophloic dictyoxylic siphonostele; pith cavity about 3—4 mm in diameter, contianing parenchyma and tracheids; xylem cylinder thin, less than 10 tracheids in radial thickness, dissected by leaf gaps. Inner cortex about 1.5 cm thick, mainly parenchymatous, but sometimes containing a few sclerenchymatous; number of leaf traces seen in a transverse section about 50—60; leaf traces departing at 35—45º,open C-shaped at point of departure, gradually becoming shallow C-shaped or V-shaped in different parts; protoxylem in base of leaf traces single, endarch; when leaf traces pass through inner cortex, protoxylem biturcating. Petiole bases without stipular expansion, probablyloosely embracing the stem; xylem strand of potiole trace shallow C-shaped, surrounded by selerenchyma; sclerotic ring round, connected with single sclerenchyma mass in the concavity of the petiole trace. Root arising singly from leaf trace, diarch, with inner and outer cortex. P. plenasioides is represented by a rhizome; stem more than 4 cm in diameter; stele actophloic dictyoxylic siphonostele; xylem cylider with about 20 tracheids in radial thickness, dissected by leaf gaps; xylem bundle U-, O-, or crosier- (i.e. query-) shaped; pith and inner cortex parenchymatous, with many groups of secretory cells; leaf trace C-shaped, its base containing two endarch protoxylem groups; root diareh,with inner and outer cortex, arising singly from leaf trace or its base.     

双子叶植物出土幼苗根茎转变区维管组织发育动态

关于根茎初生维管系统之间的连接以及与子叶的关系,在文献中已有广泛的论述,有过各种不同的解释。大部分早期关于根茎转变区的文献研究的是初生组织已完成发育的幼苗。这些研究者认为转变区域是根和茎这两种轴器官之间维管组织发生转变、相互连接的区域。但由于茎中的初生维管组织可以认为是叶迹及叶迹的延伸的综合,转变区域应被看作是轴维管系统与叶迹维管系统之间的连接。因此,转变区的研究必须说明根维管系统与最早的真叶叶迹之间的关系。通过对北乌头和大豆胚胎及幼苗维管组织的解剖学研究,本工作显示在出土萌发的双子叶植物中,初生维管组织在根-下胚轴-子叶中形成一连续系统,并完成根与子叶叶迹之间的维管组织过渡转变。而上胚轴中的维管组织是位于根-下胚轴-子叶上方独立形成的第二维管系统。上胚轴中维管组织的分化起始于第一真叶叶迹基部,向上分化进入叶片,向下进入胚轴并在子叶节下方与根-下胚轴-子叶维管系统相连接。真叶叶迹的木质部与下胚轴中靠近韧皮部的后生木质部或次生木质部连接。根与上胚轴之间不存在维管组织的过渡、转变,而只是在同样发育方向的组织中有一种直接的简单的连接．     

Secondary xylem development in Arabidopsis: a model for wood formation

Our understanding of the molecular controls regulating the identity of the vascular cambium and the development of secondary xylem and phloem have not yet benefited much from the use of Arabidopsis as a genetic system. Under appropriate growth conditions Arabidopsis undergoes extensive secondary growth in the hypocotyl, with the development of both a vascular and a cork cambium. The secondary xylem of the hypocotyl develops in two phases, an early phase in which only vessel elements mature and a later stage in which both vessel elements and fibres are found. During this second phase the secondary xylem of Arabidopsis closely resembles the anatomy of the wood of an angiosperm tree, and can be used to address basic questions about wood formation. The development of the vascular cambium and secondary growth in Arabidopsis hypocotyl is described and its utility as a model for wood formation in trees is considered.