Surface-viewed shoot apex ofAngiopteris lygodiifolia Ros. (Marattiaceae)

Marattian ferns are thought to be an exception to the rule that a single apical cell is always present in the shoot apex of ferns; the occurrence of plural apical initials has been generally accepted for these ferns. However, a contradicting conclusion was reached in this study which examined the apical organization of the shoot ofAngiopteris lygodiifolia Ros., using fresh materials which had not been fixed. Shoot apices were hand-sectioned transversely into thin sections, including the surface layer of the shoot apex, which were observed by differential interference contrast microscopy without staining. In contrast with the generally accepted view, the shoot apex ofA. lygodiifolia was found to usually possess a single apical cell with three cutting faces. The segments cut off from the apical cell are regularly arranged in a helical sequence. The apical cell seems to actually function as an initial cell of the whole shoot apex. The shoot apices, particularly those of plants cultivated in a greenhouse, sometimes show somewhat irregular organization. In extreme cases, no apical cell is recognizable. However, even in these exceptional cases of such apparently irregular shoot apices, plural apical initials are not found.     

SHOOT APICAL MERISTEMS AND MUTATION: FIXATION OF SELECTIVELY NEUTRAL CELL GENOTYPES

Shoot apical meristems are interpreted as either structured, that is having a permanent set of apical initials, or stochastic, having apical initials which represent “... momentary representatives of the continuous meristematic residue at the apex of the relevant layer or zone” (Newman, 1965). The two main parameters of stochastic growth are the average number of apical initials (α) and the number of mitotic cycles (r) of the initials and their daughter cells prior to the random selection of subsequent initials. Mathematical analysis and computer simulation studies of stochastic growth have shown that if one starts with 1 mutant initial and α-1 nonmutant initials, eventually a mosaic plant results. The frequency of shoot apices composed of mutant cells is 1/α and the frequency of shoot apices composed of only nonmutant cells is (1 – α)/α. These asymptotics are only attained after considerable growth, thus mericlinal chimeras can persist for many nodes and give the appearance that a permanent set of initials is present.     

TRANSLOCATION OF 14C IN MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1,2

Translocation patterns in the giant kelp, Macrocystis integrifolia Bory, were investigated in situ using ¹⁴C tracer; sources and sinks were identified. Export was first detected after 4 h of labeling; experiments were routinely 24 h continuous ¹⁴C application. Mature blades exported ¹⁴C to young blades on the same frond and on younger fronds, as well as to sporophylls and frond initials at the bases of the fronds. Blades <0.3 m from the apex imported and did not export; this distance did not change seasonally. In spring export from blades 0.3–1.25 m from the apex was exclusively upwards; older blades also exported downwards. In fall downward export began 0.5 m from the apex, and blades >2 m from the apex exported exclusively downwards. Carbon imported by frond initials, young fronds, and sporophylls in fall may partly be stored for growth in early spring. No translocation was seen in very young plants until one blade (secondary frond initial) bad been freed from the apical blade; this blade exported to the apical blade for a time, but imported when it began to develop into a frond. The second and third formed blades on the primary fronds (sporophylls also exported when <0.3 m from the apex, and later stopped. Frond initials and sporophylls on later-formed fronds did not export at all. The translocation pattern in M. integrifolia differs from that previously reported in M. pyrifera in seasonal change and in distances from the apex at which the changes take place.     

THE CYTOHISTOLOGICAL AND CYTOHISTOCHEMICAL ZONATION OF THE SHOOT APEX OF BOTRYCHIUM MULTIFIDUM

Cytohistologically, the shoot apex of Botrychium multifidum is composed of three zones—a zone of surface initials in which there is usually a centrally located apical cell, a zone of subsurface initials, and a cup-shaped zone that is subdivided into a pheripheral zone and a rib meristem. The results of cytohistochemical tests for total protein, RNA, total carbohydrate, histones, and DNA localization support this concept. Thus, the cytohistological zonation of the apical meristem of the Ophioglossales is essentially identical to that of the Filicales, and furthermore, is comparable to that of the seed plants.     

STUDIES ON THE ONTOGENY OF THE DWARF MISTLETOES,ARCEUTHOBIUM. II. HOMOLOGY OF THE ENDOPHYTIC SYSTEM

Cohen , L. I. (Washington State U., Pullman.) Studies on the ontogeny of the dwarf mistletoes, Arceuthobium. H. Homology of the endophytic system. Amor. Jour. Bot. 50(5): 409–417. Illus. 1963.—Development of the seedling in Arceuthobium, as well as the mode of penetration and infection, is described. The promeristem of the root apex of the seedling, as in the embryo, is composed of 4 zones: (1) a uniseriate layer of apical surface initials; (2) a subapical zone of central initials; (3) a peripheral zone; and (4) a zone of procambial initials. After germination, the seedling grows on the host surface until the root apex is confronted by a spur shoot or by a fragment of raised bark. The root apex becomes attached to the host by means of a massive holdfast. It is the procambial initials which actually invade the host tissue. After penetrating the host, the individual procambium initials proliferate in the cortex, each giving rise to a cortical strand. It was concluded that the endophytic system of Arceuthobium cannot be interpreted in terms of classical concepts of plant homology; it is, in fact, an organ sui generis.     

PLASTOCHRONIC CHANGES AND THE CONCEPT OF APICAL INITIALS IN EPHEDRA ALTISSIMA

Paolillo , Dominick J., Jr ., and Ernest M. Gifford , Jr . (U. California, Davis.) Plastochronic changes and the concept of apical initials in Ephedra altissima. Amer. Jour. Bot. 48(1): 8–16. Illus. 1961.—The occurrence of distal mitoses and the differentiation of derivatives of the distal cells of the apex of E. altissima are correlated with stages of the plastochronic cycle. The relative rates of growth of elements within the various regions of the apex are characterized in terms of elongation gradients. The numbers of mitoses found in median sections of individual apices are recorded. It is concluded that the distal zone is the source of all other cells of the apex and should be regarded as a zone of initials. The concept of “méristème d'attente” and “anneau initial” is discussed in relation to the observations made on E. altissima.     

AN ELECTRON MICROSCOPIC EVALUATION OF CYTOHISTOLOGICAL ZONATION IN THE SHOOT APICAL MERISTEM OF PINUS BANKSIANA

Shoot apical meristems of jack pine (Pinus banksiana) were examined by light and electron microscopy. Cytohistological zonation was evident when meristems were fixed in Craf IV, embedded in paraffin, and stained with Chlorazol Black E. When meristems were fixed for electron microscopy the cytoplasm of the apical initials and central mother cells each contained numerous lipid bodies and their nuclei contained little, if any, heterochromatin. The cytoplasm of the peripheral zone was rich in ribosomes. The nuclei of the peripheral zone and rib meristem were heterochromatic. Thus, the lack of heterochromatin in the nuclei and the dissolution of lipids in the cytoplasm of the apical initials and central mother cells appeared to contribute most to the organization and appearance (cytohistological zonation) of the shoot apex when standard histological techniques are used.     

CYTOLOGY AND MORPHOGENESIS IN THE RHIZOMORPH OF ARMILLARIA MELLEA

Aided by the techniques of thin sectioning and electron microscopy, the apical region of the rhizomorph of Armillaria mellea has been examined. This region is composed of concentric zones of morphologically distinct tissues derived from a subapical meristematic zone designated the apical center. Meristematic activity is of two types: (1) primary, localized in the apical center, in which new hyphal elements are formed from apical initials, and (2) secondary, localized in the lateral regions of the apex, in which elaboration of the hyphal elements by means of elongation and secondary crosswall formation takes place. From these meristematic zones the tissues of the mature rhizomorph are derived and include: (a) peripheral hyphae, (b) cortex, (c) subcortex, and (d) primary and secondary medulla. The manner of differentiation of an apical initial appears unique and involves synchronous nuclear divisions accompanied by segmentation in many planes. The result of this activity is the formation of multinucleate hyphae. Apical initials are usually highly cytoplasmic and possess peculiar non-membrane-bound fibrous bundles, but in all other respects they resemble the hyphae of most Basidiomycetes thus far examined with the electron microscope.     

THE ROOT APEX OF LINUM

Several developmental stages of primary roots of 13 species of Linum were studied. The basic pattern of root apex organization in three species consisted of a single tier of cortical initials. Ten species had a basic pattern of two tiers of cortical initials. Variations, manifested by either an increase or a decrease in the tiers of cortical initials, were observed in some roots in those species that had a basic pattern of two tiers of cortical initials. Although these variations were interpreted as being ontogenetic, there was no total reorganization of the root apex. There was anatomical and cytological evidence that a quiescent center is established in Linum roots.     

Structure and Cytogenesis of the Shoot Apex of Syringa oblata var. affinis Lingelsh

The structure, growth and mitotic activity of 211 shoot apices of developing sprouts of Syringa oblata var. affinis Lingelsh. in longitudinal sections and 67 in transverse sections have been studied with the view to understanding the nature of zonation patterns and cytogenesis of the apical meristems during a double plastochron. The external morphology and the anatomical structure of the apices in 4 plastochronic stage-early, middle, late Ⅰ and late Ⅱ stages are described. In the shoot apices examined, especially those at late plastochronic stage, the following zones may be delimited: Zone of tunica initials, zone of corpus initials, peripheral zone and zone of rib meristem. The location and orientation of mitotic figures observed in longisections of the apices in 4 plastochronic stages are plotted in diagrams and the mitotic frequency has been calculated. Information obtained from these investigations reveals that the tunica and corpus inititals constitute an active region of the apex, but their mitotic activity changes periodically within the double plastochron. In the middle plastochronic stage when the apex is at its minimal area and the cells of peripheral zone and rib meristem zone have been completely transformed into constituent parts of foliar primordia and the subjacent tissues of the stem and the pith mother cells respectively, the mitotic frequency of the initials is at its maximium and its intensity of mitotic activity is not much lower than that of any other meristematic zone at any stage. When the apical dome is reformed by the activity of these initials in late plastochronic stages, the mitotic frequency of the initials gradually drops and the region of high mitotic frequency shifts to the flank of the apex, the peripheral zone. Anticlinal divisions are predominant in this zone. On the other hand, those cells directly left behind by the corpus initials, which constitute the rib meristem, are vacuolated and marked by the pre- dominance of transverse divisions. Thus the entire zonation pattern reappears. In the next early plastochronic stage, the mitotic frequency of the tunica and corpus initials drops to its mimimium, but other regions of the apex still maintain a high mitotic frequency. It may be concluded that the tunica and corpus initials form a cytogenerative center of the shoot, and the cytohistological zonation is actually a result of the fact that different regions of apical meristems are different in mitotic activety, different in state of cell differentiation and different in their function in morphogenesis.     

SHOOT APEX ORGANIZATION AND ORIGIN OF THE RHIZOME-BORNE ROOTS AND THEIR ASSOCIATED GAPS IN DENNSTAEDTIA CICUTARIA

The shoot apex of Dennstaedtia cicutaria consists of three zones—a zone of surface initials, a zone of subsurface initials, and a cup-shaped zone that is subdivided into a peripheral region and central region. A diffuse primary thickening meristem, which is continuous with the peripheral region of the cup-shaped zone, gives rise to a broad cortex. The roots occurring on the rhizomes are initiated very near the shoot apex in the outer derivatives of the primary thickening meristem. The roots that occur on the leaf bases also differentiate from cortical cells. Eventually, those cortical cells situated between the newly formed root apical cell and the rhizome procambium (or leaf trace) differentiate into the procambium of the root trace, thus establishing procambial continuity with that of the rhizome or leaf trace. Parenchymatous root gaps are formed in the rhizome stele and leaf traces when a few of their procambial cells located directly above the juncture of the root trace procambium differentiate into parenchyma. As the rhizome procambium or leaf trace continues to elongate, the parenchyma cells of the gap randomly divide and enlarge, thus extending the gap.     

ORIGIN AND DEVELOPMENT OF THE TERMINAL CARPEL IN PSEUDOWINTERA TRAVERSII

Flowers of Pseudowintera traversii (Buchan.) Dandy possess a terminal unicarpellate gynoecium. The present study of carpel morphogenesis was initiated for the purposes of (1) providing additional developmental documentation of the occurrence of terminal carpels in the Winteraceae and (2) comparing the mode of initiation and development of the ascidiate terminal carpel of P. traversii with the essentially conduplicate terminal carpel of Drimys lanceolata. Following its axillary origin, the floral apex of P. traversii initiates 2–3 connate sepals, 5–6 petals, 4–15 stamens, and usually a single terminal carpel, in acropetal succession. Bicarpellate gynoecia may occur with a frequency of up to 15 % on a given plant. The floral apex is zonate and shows increased expression of its zonation during later stages of floral development. The terminal carpel is ascidiate from inception and originates as a cylindrical growth around the entire circumference of the floral apex; transformation of the floral meristem into a carpel primordium terminates apical growth of the floral axis. Carpel growth continues to be cylindrical and is mediated by a ring of marginal and submarginal initials at its summit. Earlier and more extensive division of initials and their derivatives on the dorsal rim causes the primordium to become canted adaxially, shifting the apical cleft to a subterminal adaxial position. Continued marginal meristematic activity results in closure of the cleft as well as elevation and elaboration of the stigmatic crests. Five to seven bitegmic ovules are initiated at the same time as crest elaboration and arise in two rows from the adaxial (laminar) position. Carpel maturation is signified by tannin deposition and oil cell differentiation, beginning at the base and proceeding acropetally; carpel margins bordering the cleft are the last to differentiate. Carpel procambialization is continuous and acropetal from inception, with the dorsal median bundle differentiating before the ventral strands. The significance of occasional bicarpellate flowers is discussed.     

THE DEVELOPMENTAL ANATOMY OF LONG-BRANCH TERMINAL BUDS OF PINUS BANKSIANA

A study of the composition of long-branch terminal buds (LBTB) of Pinus banksiana Lamb. and the yearly periodicity associated with their formation, development, and elongation was undertaken. Each LBTB has lateral bud zones and zones of cataphylls lacking axillary buds. When present, staminate cone primordia differentiate from the lowest lateral buds in the lowest lateral bud zone of the LBTB. Ovulate cone primordia and lateral long-branch buds can differentiate from the upper lateral buds in any lateral bud zone. When both types of buds are present, lateral long-branch buds are uppermost. Dwarf-branch buds occur in all lateral bud zones. During spring LBTB internodes elongate, new cataphylls are initiated, dwarf branches elongate, needles form and elongate, pollen forms and is released, and ovulate cones are pollinated. During summer buds form in the axils of the newly formed cataphylls. By early fall the new LBTB are in overwintering condition and the four types of lateral buds are discernable. The cytohistological zonation of the LBTB shoot apex is similar to that of more than 20 other conifer species. Cells in shoot apices of pine are usually arranged in distinct zones: apical initials, subapical initials, central meristem, and peripheral meristem. Periclinal divisions occur in the surface cells of the apex; therefore no tunica is present. At any given time, shoot apex volume and shape vary among LBTB in various positions on a tree. In any one LBTB on a tree, shoot apex shape changes from a low dome during spring to a high dome during summer to an intermediate shape through fall and winter.     

A model of frequency and distribution of mutations among genetically effective initials in the plant shoot apex

In an embryo shoot apex more different mutations may be induced after the mutagenic treatment of seeds. These mutations may be distributed among the apical initial cells in different ways. According to the proposed mathematical model it is possible to estimate the proportion of initial cells with 0, 1, 2, …,x mutations in the apioes with a given number of initials,k, which is characteristic for each plant object, and under a certain number of induced mutations per apex,n, determined by the conditions of the mutagenio treatment. Owing to the disadvantage of multimutations in plant breeding it is possible according to the formula P(X > 1) = 1 −e^−n/k(l + n/k). to use, at a knownk, suchn’s which secure minimum multimutations.     

SHOOT ORGANIZATION IN THE FILICALES: THE PROMERISTEM

The classical pteridophyte apical cell theory of meristem organization is not flexible or realistic enough to encompass the variation encountered in a comprehensive anatomical survey of fern meristems. The meristems of 28 taxa of ferns (both eu- and leptosporangiate) were studied. This analysis has led to the formulation of the concept of a promeristem composed of two zones: the surface and subsurface initials. This concept is flexible and sufficient to describe the variation encountered in the ferns as a group. No differences in promeristem organization were observed in plants with various rhizome morphology (e.g., upright and radial; prostrate and dorsiventral). Marking experiments, performed on living surface cells of fern promeristems, correlate with anatomical observations of division planes in the promeristem and indicate that the central, surface initials are not quiescent. Feulgen determinations indicate that Nephrolepis stolons have no endomitotically polyploid cells in the promeristem. Additional work is needed before a generalization can be made concerning ploidy levels in a more typical fern apex. Ferns, in general, have a zoned promeristem which is parallel to the zonation described for higher vascular plants.     

Die Polarisierung der Scheitelzellen von Stypocaulon scoparium

Summary Young isolated apical initials of Stypocaulon scoparium usually regenerate in a bipolar way. Older initials after having produced the basal cell of a branch are reduced in their regenerating capacity to unipolarity, i. e. after each division the polarity of the apical cell must be newly induced.

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Herrn Prof. Dr. R. Harder zum 80. Geburtstag gewidmet.     

Development of mantle leaves inPlatycerium bifurcatum (Polypodiaceae)

InPlatycerium bifurcatum the leaf primordia emerge alternately right and left below the shoot apical cell on the dorsal surface of the rhizome. They arise from groups of small cells, a single large cell becoming the initial of the leaf apical cell. The longitudinal axis of the leaf apical cell is at a right angle to the rhizome axis and the leaf primordia are arranged longitudinally in two rows. The leaf apical cell gives rise to marginal initials which are responsible for leaf growth in one plain. Early marginal cells are crescent-shaped while the later ones are wedge-shaped. Hairy marginal cells appear in the very early stages of development. The interpretation of these cells as a promeristem and as initials are discussed.     

DEVELOPMENTAL ANATOMY IN ROOTS OF IPOMOEA PURPUREA. I. RADICLE AND PRIMARY ROOT

The developmental anatomy of the primary root of Ipomoea purpurea was studied at several growth stages, beginning with the radicle. The radicle is generally composed of three superimposed tiers of initials, which produce the vascular cylinder, cortex, and columella; and a peripheral band of lateral rootcap-epidermal initials. The radicular cortex contains 16–19 immature laticifers; none of the tissue regions in the radicle contains mature cells. Following germination and during the first 2–3 cm growth of the primary root the apical meristem and its derivative tissues undergo a series of modifications. Root apical diameter decreases as cells in lateral portions of the rootcap elongate; meanwhile, the columella enlarges vertically. The relationship between cortical and columellar initials changes as fewer mitoses occur in the former while the latter remain active. In longer roots the columellar initials are directly in contact with the vascular initials. Cortical size diminishes during early root growth as cortical laticifers and their associated cells cease to be produced by the outer cortical initials and ground meristem. Early procambium, at the level of vascular pattern initiation, decreases in diameter by cellular reorientation, and the vascular cylinder decreases in overall diameter although the tetrarch pattern remains unchanged.     

RNA Synthesis in the Shoot Apex of Polytrichum formosum Hedw.

RNA metabolism in the cells of the gametophyte shoot apex ofPolytrichum formosum was investigated using both microspectrophotometricand autoradiographic methods along with an accurate measurementof surfaces and volumes of nuclei, nucleoli, free cytoplasmand vacuolar systems. On a per cell basis, the amount of RNAand the rate of RNA synthesis were shown to be highest in theapical cell. On the other hand, both RNA concentration and rateof synthesis for a unit quantity of cytoplasm were found tobe higher in leaf initials and in the cells of young leavesthan in theapical cell, the segments and the segmental derivatives.For the various types of cells in the shoot apex it was establishedthat the more voluminous the nucleolus, the higher the RNA syntheticrate per cell. These results were correlated with the data previouslyobtained on the mitotic cycles in Polytrichum. It is concludedthat in the apical cell, notwithstanding its differentiatedfeatures, RNA metabolism must be considered on the whole tobe very active. These various data are compared with those obtainedon angiosperm shoot apices.     

EMBRYOGENY AND HISTOGENESIS IN NERIUM OLEANDER II. ORIGIN AND DEVELOPMENT OF THE NON-ARTICULATED LATICIFER

Mahlberg , P. G. (U. Pittsburgh, Pittsburgh, Pa.) Embryogeny and histogenesis in Nerium oleander. II. Origin and development of the non-articulated Iaticifer. Amer. Jour. Bot. 48(1): 90–99. Illus. 1961.—Laticifer initials, collectively considered as a laticifer system, are differentiated in the globular embryo from meristematic cells which occupy a position within the potential procambial tissue. A total of usually 28 initials, in Nerium oleander, arise as an irregular ring of cells directly below the embryonic shoot apex, during initiation of the cotyledonary primordia. No anastomoses occur between laticifer initials. During subsequent development of the embryo, the laticifer initials grow in a bi-directional manner and penetrate into the root, cotyledons and toward the shoot apex. Upon enlargement the initials bifurcate repeatedly, many branches penetrate into the cotyledons, others grow into the cortex of the hypocotyl or penetrate between cells of the procambium. Repeated nuclear divisions within each initial result in the formation of a multinucleated protoplast in this cell type. The tips of laticifers occupy intercellular spaces during their growth; they do not penetrate into or through adjacent cells. A plexus of laticifer branches is formed within the cotyledonary node of the mature embryo. No new initials are formed during subsequent growth of the plant, rather certain branches from the cotyledonary nodal plexus penetrate into the enlarging shoot system. The nature of their growth habit and branching suggests that the tips of laticifer initials exhibit an intrusive form of growth.