Serial Development of Foliar Gemmae in Tortula (Pottiales, Musci), an Ultrastructural Study

The development and liberation mechanism of foliar gemmae havebeen studied by electron microscopy in two mosses, Tortula latifoliaBruch and Tortula papillosa Wils. The gemmae develop on theadaxial surface of mature leaves from single initial cells onboth the lamina and costa in T. latifolia but only on the costain T. papillosa . Elongation of the initial cell is associatedwith the deposition of a highly extensible new wall whilst theold wall and cuticle in the apical dome rupture. The first divisionis transverse and separates a short basal cell embedded in thefoliar tissue and a distal cell, or gemma primordium, protrudingfrom the leaf surface. Subsequent divisions of the gemma primordiumgive rise to a six-to-eight-celled globose gemma with mucilaginousouter walls. During gemma development the basal cell producesa new wall and elongates again whilst the common wall with thegemma splits apart centripetally along the boundary betweenthe old and new wall in the basal cell; plasmodesmal connectionsare gradually severed and eventually the young gemma remainsconnected to the basal cell only by mucilage. After separationof the first-formed gemma, the basal cell may expand and producea second gemma by the same mechanism. The whole process maybe repeated several times resulting in the formation of a chainof gemmae stuck together by mucilage and which are liberatedonly when the leaves are fully hydrated. Accumulation of abundantlipid deposits in the gemmae after symplasmic isolation reflectsconsiderable photosynthetic autonomy. Abscission; bryophytes; cell wall formation; plasmodesmata; vegetative reproduction     

疣壶藓尖叶变种的芽胞形成及其生态意义

该研究以疣壶藓尖叶变种(Gymnostomiella vernicosa var.acuminata)为实验材料,在人工培养条件下观察并记录其芽胞产生及脱落过程,以揭示该分类群及相关类群的芽胞形成过程和机制,为疣壶藓尖叶变种的分类提供参考性特征指标,明确苔藓植物芽胞的进化和生态学意义。结果显示,疣壶藓尖叶变种的芽胞形成过程划分为3个阶段:(1)茎表皮细胞的外切向壁局部向外隆起,外突的疣壶藓尖叶变种表皮细胞富含叶绿体,可进行光合作用。(2)芽胞起始细胞横向分裂,形成一个基细胞和一个顶细胞;基细胞经数次横向分裂,下部形成柄状结构,上部的细胞常呈喇叭形的“柄托”;顶细胞经多方向细胞分裂,形成椭球状至圆球状芽胞体,形成初期为富含叶绿体的厚壁细胞,成熟后叶绿体数目逐渐减少,最终变成深棕色至红棕色。(3)由薄壁的柄细胞随机破损使芽胞从母株上脱落。研究表明,在无法预测的多变栖息环境中,相对于有较高的能量、遗传和生态消耗的有性生殖,疣壶藓尖叶变种产生芽胞是一种风险分担策略,可以提高生存几率和有效利用资源,可视为一种扩大和维持种群的有效途径。     

The Formation of Catenate Foliar Gemmae and the Origin of Oil Bodies in the Liverwort Odontoschisma denudatum (Mart.) Dum. (Jungermanniales): a Light and Electron Microscope Study

This article describes the ultrastructural events associatedwith the differentiation and liberation of the exogenous gemmaeproduced in branched acropetal chains along the margins of theleaves in the liverwort Odontoschisma denudatum. Formation ofa dorsal protrusion from young leaf cells containing a largecentral nucleus, small vacuoles, starch-free chloroplasts, scatteredcytoplasmic lipid droplets but no oil bodies, signals the onsetof the formation of the initial cell of a gemmiferous filament.The protrusion enlarges and the nucleus migrates into its base,therein dividing with the equator of the spindle virtually fillingthe central isthmus between the leaf surface and the now swollentip of the initial cell. Subsequent divisions of the initialcell produce a chain of cells in atropetal succession. Transverselyorientated microtubules line the cortical cytoplasm along thelateral walls of the terminal cells of the gemmiferous filaments,but are absent from the tips, thus suggesting that these cellselongate by intercalary, rather than by tip, growth. Duringmitosis microtubules are closely associated with the envelopesof spindle-shaped prophase nuclei, radiate from ill-definedspindle poles surrounded by plastids at metaphase and anaphaseand form a dense phragmoplast array during telophase. Pre-prophasebands are absent and it may be that the nuclear equator determinesthe plane of division in gemmiferous filaments. Chloroplastdivision, associated with extremely transient plastid-dividingrings, takes place during interphase. Lateral branches of thegrowing filaments arise from subapical cells by reiterationof the first division mechanism. Immediately following the proliferative divisions, which takeplace in cells measuring only 5-6 µm in diameter, oilbodies suddenly appear as flat pleomorphic cisternae associatedwith endoplasmic reticulum and occasional microtubules. Theircontents are electron-transparent apart from scattered osmiophilicdroplets. Throughout their ontogeny the oil bodies are closelyassociated with cytoplasmic lipid bodies but there is no evidenceof fusion. The nascent oil bodies swell rapidly to their finaldiameter, become ovoid to spherical in outline and are eventuallysuspended by fine cytoplasmic bridges within the vacuoles. Thelatter rapidly increase in size together with an expansion ofthe cells themselves until these reach their final diameterand length. The final event in gemma maturation is an endogenousdivision with the formation of a new internal wall along a phragmosome.Separation of the bicellular gemmae proceeds basipetally andinvolves the appearance of an electron-transparent line alongthe middle lamella in the cross walls, which often develop convexthickenings, and severing of the plasmodesmata. After theirliberation shallow scars are visible on the leaf surface underthe SEM. Gemma maturation sees a marked increase in the electron-opacityof the walls and dense staining of these together with Golgivesicles with the periodic acid/thiocarbohydrazide/silver proteinatetest for non-cellulosic carbohydrates. This change in wall chemistryand ultrastructure may be related to the fact that the maturinggemmae become extremely water repellant and are probably dispersedeither on the surface of water films or in the air.Copyright1995, 1999 Academic Press Bryophyta, gemmae, liverwort, microtubules, morphogenesis, oil bodies, polar growth, vegetative reproduction     

Presence and absence of the preprophase band of microtubules in moss protonemata: a clue to understanding its function?

Summary InFunaria protonemata, preprophase bands (PPBs) of microtubules do not develop when the tip cell divides, when side branches are initiated or in intercalary regeneration divisions. We report here that PPBs do, however, develop when a tmema cell is formed. In the former cases, cell division is not coupled with an expansion of the mother cell wall at the site where the cell plate will attach. In the latter case, the mother cell wall ruptures at that site and the tmema cell elongates. This observation and the findings on presence and absence of the PPB in other cell types indicate a connection between PPB occurrence and mother cell wall expansion. They support the idea that the PPB might be involved in the local secretion of cell wall material. We extend this notion, suggesting that the microtubules of the PPB control the oriented deposition of a thin layer of cellulose microfibrils at the mother cell wall which supports the firm attachment of the cell plate when the mother cell wall expands.Abbreviations FITC fluorescein isothiocyanate - IgG immunoglobulin G - MT microtubule - PPB preprophase band of microtubules - TC tmema cell     

Embryology ofCymbidium sinense:the Microtubule Organization of Early Embryos

InCymbidium sinense, the pattern of embryo development is unusualin that oblique cell divisions result in the formation of severalsuspensor cells prior to the development of the embryo proper.Characteristic changes in microtubular distribution can be foundwithin the zygote and the proembryo during their development.After fertilization, the ellipsoid-shaped zygote has randomlydistributed microtubules within its cytoplasm. As the zygotetakes on a more rounded appearance, microtubules organize intoa dense meshwork. Furthermore, microtubule bundles appear atthe chalazal region of the cell prior to the first mitotic divisionof the zygote. At the preprophase stage of mitosis, a preprophaseband of microtubules appears in the cytoplasm of the zygote.The zygote divides obliquely and unequally and gives rise toan apical cell and a slightly larger basal cell. Many randomly-alignedmicrotubules can be found in the cortex of the basal cell. Theincrease in the abundance of microtubules coincides with theisotropic expansion of the basal cell. The early division ofthe basal cell and subsequent division of the apical cell resultsin the formation of a four-celled embryo, of which three cellsnear the micropylar pole develop as suspensor cells. In thesuspensor cells, the microtubules tend to orient in the samedirection as the long axis of the cell. In addition, prominentmicrotubules can also be found near the adjoining cell wallsof the four-celled embryo. The terminal cell is highly cytoplasmicwith abundant microtubules within the cell. Subsequent divisionsof the terminal cell give rise to additional suspensor cellsand the embryo proper. In the mature embryo, five suspensorcells are usually present; one eventually grows through themicropyle of the inner integument and four grow towards thechalazal pole. The cortical microtubules of suspensor cellsredistribute from a longitudinal to a transverse direction asthey grow towards their respective poles.Copyright 1998 Annalsof Botany Company Embryogenesis, endosperm, microtubules, preprophase band, suspensor cells,Cymbidium sinense(Andr.) Willd.     

Development and germination of rhizoidal gemmae of Bryum violaceum

Approximately four to six weeks after transferring gametophores of Bryum violaceum Crundwell & Nyholm to fresh soil, abundant new rhizoids with stalked gemmae were present at the bases of the gametophores. The development of gemmae on rhizoids was followed from single cell initials to multicellular, three-dimensional forms. Mature gemmae, the vegetative diaspores, were pale orange, purple or reddish brown and each had a uniseriate stalk of two to four cells. While remaining on soil, the rhizoidal gemmae showed no in situ germination. However, following the removal of gametophores with rhizoidal gemmae, or rhizoids alone with gemmae, and their placement upon wet filter paper in Petri dishes in light, the gemmae germinated. During germination, the protonemata emerged consistently from mature gemmae at their distal ends, revealing the existence of gemma polarity. In the case of immature gemmae, on the other hand, the protonemata emerged from any surface cell indicating that gemma polarity had not yet been established.     

Some noteworthy bryophytes from Tenerife

Abstract

Mosses exhibit a greater variety of cellular separation mechanisms than any other group of land plants. Diaspore liberation mechanisms range from (1) the random breakage of thin-walled stalk cells to (2) the formation of new internal walls that separate from the old walls, (3) severance along the middle lamella of the basal cell with or without the rounding off of the cells, (4) the formation of highly specialized abscission or tmema cells and (5) breakage along an intercalary region of thinwalled living cells. Rhizoidal gemmae are the only propagules lacking a separation mechanism other than by the decay of the filament system that produces them. In some species, two and sometimes three different kinds of diaspore are formed simultaneously in culture. Diaspore germination patterns in mosses are even more diverse than the liberation mechanisms. With a few exceptions new growth from diaspores is filamentous. Most diaspares are highly polarized and the germination pattern is fixed during development. Protonemal and rhizoidal gemmae are defined on the basis of the filament systems that produce them and from the presence or absence of specialized abscission mechanisms.     

Growth and Dormancy in Lunularia cruciata (L.) Dum.: I. RESUMPTION OF GROWTH AND ITS CONTINUATION

Lunularia cruciata may become dormant at three stages in itslife history: mature thallus, gemma, and spore. The resumptionof growth and its continuation in various conditions have beenstudied in thalli and gemmae. Air-dry, mature thalli of theIsrael strain, planted on a suitable medium, produce adventitiousbranches ventrally from the region immediately posterior tothe existing meristem, which itself fails to resume growth.When dormant gemmae are taken from the gemma-cup, however, theexisting apical cells and meristems simply recommence growing,so that the new growth is continuous with the old. Except inthe case of mature thalli aroused from dormancy, apical dominanceis pronounced, and branching occurs only by bifurcation. Thisdominance can be broken by applying sucrose solution to thegrowing tips—possibly a plasmolytic effect. The growth in area of freshly planted gemmae accelerates forabout 25 days before its relative rate slows to any great extent.During the first half of this period, growth is due exclusivelyto the expansion of existing cells, but subsequently cell numbersincrease rapidly from the 8–10,000 present in the dormantgemma. Cell numbers were estimated by using a modified macerationtechnique, in which chelation followed prolonged fixation. Lunularia grows successfully at quite low light intensities.Of the mineral nutrient deficienccs investigated, lack of Plimits growth most severely, although N-deficiency also restrictsit to a very low level. Gemma-cup production appears to be unaffected by light intensity,at least within levels permitting growth. There is, however,a large temperature effect, cup production decreasing markedlyabove 12° C.     

The Floral Nectaries of Hibiscus rosa-sinensis: 1. Development of the Secretory Hairs

Floral nectaries of Hibiscus rosa-sinensis occur on the lowerinner side of the fused sepals and each one consists of numerous(50000–55000) secretory hairs, occupying a cylinder-likezone completely lining the inner side of the sepals. Each hairoriginates from a single protodermal mother cell and, at maturity,it is built up of a basal cell, a stalk, 35–40 intermediatecells and a tip secretory cell. Development of protodermal cellsinto secretory hairs is asynchronous, the first cells to initiatedevelopment being those situated in the lowermost part of thecylindrical zone, and development progressing upwards. Volume increase of protodermal mother cells initiating developmentis accompanied by cell polarization manifested by organelledisplacement towards the apical region. Secretory hairs areformed through a sequence of transverse and, later on, anticlinaldivisions. Divisions of apical cells are preceded by well definedpre-prophase microtubule bands, which foreshadow the plane ofthe forthcoming division and predict with accuracy the sitesof parental walls where the new cell plate fuses at cytokinesis. Stalks consist of either one or two cells. Two-celled stalksoccur in 40 per cent of secretory hairs and derive from a transversedivision of one stalk cell; the wall formed is always depositedparallel to the proximal and distal walls, but never to thelateral ones. The significance of this mode of division is discussedin relation to the fact that lateral walls are entirely impregnatedwith a cutin-like material that blocks apoplastic movement ofsolutes. Hibiscus rosa-sinensis, nectaries, development, preprophase microtuble bands, stalk cells     

Microscopic analysis and seasonality of gemma production in the freshwater red alga Hildenbrandia angolensis (Hildenbrandiales, Rhodophyta)

The development and release of the unique vegetative propagules of the freshwater encrusting alga Hildenbrandia angolensis Welwitsch ex West et West, gemmae, were studied using several different microscopic and histochemical techniques. In addition, the seasonality of gemma production was monitored bimonthly over a 12‐month period in two spring‐fed streams in Texas, USA. Gemmae differentiate within the thallus and are subsequently released from the surface of the crust. Release of the gemmae most likely occurs by digestion of surrounding cells, as suggested by the presence of starch granules and lipid globules in the region between the released gemma and the thallus. The initial separation of the gemmae from the thallus occurs from the sides of the gemma or the bottom, or possibly simultaneously. Contrary to previous studies, we have observed that gemma production occurs endogenously within the thallus of freshwater Hildenbrandia, rather than on the surface of the crust in raised structures. Histochemical tests and electron microscopic examination indicate that the cells of the gemmae contain a large amount of floridean starch. The starch granules frequently form rings surrounding the nuclei of both gemma and thallus cells; a feature infrequently reported for florideophyte red algae. Our seasonality investigations indicate that large fluctuations in gemma production occur over 1 year, but at least some gemma production continues year‐round in the streams examined.     

ROLE OF ASYMMETRIC CELL DIVISION IN PTERIDOPHYTE DIFFERENTIATION. II. EFFECT OF CA2+ ON ASYMMETRIC CELL DIVISION,RHIZOID ELONGATION,AND ANTHERIDIUM DIFFERENTIATION IN VITTARIA GEMMAE

Gametophytes of Vittaria graminifolia reproduce vegetatively by means of gemmae. Each gemma consists of a linear array of six cells: four body cells and a knob-shaped terminal cell at each end. When gemmae are shed from the gametophyte onto Knop's mineral medium, the two terminal cells do not divide, but elongate to form primary rhizoids. The body cells undergo asymmetric cell division, and the smaller daughter cells differentiate into either secondary rhizoids or prothalli. When gibberellic acid is included in the medium, antheridia are formed as a result of asymmetric cell division instead of vegetative structures. We studied the effect of Ca²⁺ on asymmetric cell division, rhizoid elongation, and antheridium formation in gemmae cultured on Knop's mineral medium and variations of Knop's medium. Ca²⁺ inhibited the onset of cell division and rhizoid elongation, but was required for differentiation of antheridia. Treatments which lowered the Ca²⁺ content of gemmae (EGTA and dilute HCl extraction, culture on verapamil-containing and Ca²⁺-deficient medium) caused an early onset of cell division and rhizoid elongation. The stimulation of growth was most pronounced when gemmae were deprived of Ca²⁺ during the first 24 hr of culture. The proportion of cell divisions which differentiated into antheridia in response to GA was greatly reduced when the Ca²⁺ status of gemmae was lowered with verapamil and Ca²⁺-EGTA buffers.     

ABSORBING TRICHOMES IN THE PLEUROTHALLIDINAE (ORCHIDACEAE)

Glandular trichomes occur on both surfaces of leaves of all examined genera and species of the subtribe Pleurothallidinae (Orchidaceae). Trichome initiation is effected by one periclinal division of a protodermal cell, producing a thin-walled, globose apical cell with a relatively large nucleus and a subapical stalk cell with heavily cutinized lateral walls. In some species a second periclinal division produces a third small basal cell also having thick lateral walls but thin transverse walls. As leaf development proceeds, the trichome apparatus assumes a sunken position due to continued anticlinal divisions of protoderm. Prior to laminar expansion and guard-mother-cell division on the abaxial surface, the wall of the apical cell ruptures and is replaced by a brown opaque residue. Finally, after vascular tissue differentiation and the cessation of meristematic activity, two or more pitted foot cells develop at the base of the trichome and adjacent to the water-storing hypodermal layers. Preliminary investigations indicate that the trichome apparatus is absorptive throughout its development and similar in function to tillandsioid scales in Bromeliaceae.     

Ultrastructure of Mitosis in Entosiphon sulcatum (Euglenida)1

ABSTRACT The ultrastructural features of cell division in the biflagellate, phagotrophic euglenoid, Entosiphon sulcatum, have been examined. Prophase is marked by the appearance of daughter feeding apparatuses and the emergence of two additional flagella. Pairs of flagella begin to migrate laterally along the surface of the elongating nucleus and remain lateral to the developing spindle poles. As the nucleolus elongates, it becomes dumbbell-shaped and the chromosomes move to the center of the nucleus, forming a loosely organized metaphase plate. Microtubules from opposing spindle poles attach to one of the pair of kinetochores found on each chromosome. The initial chromosome separation occurs during anaphase as the nucleus elongates. The length of the chromosomal microtubules does not decrease until late anaphase/early telophase. As the nucleus elongates, it forms a dumbbell-shaped structure. Most of the remaining microtubules are positioned in the interzone between the forming daughter nuclei. The interzonal spindle becomes somewhat constricted but remains intact until it is broken by the impinging cleavage furrow. Replication of the pellicular strips is not completed until late in cytokinesis.     

A Sequential Study of Cell Divisions and Expansion Patterns on a Single Developing Shoot Apex of Vinca major

During the growth of a single developing vegetative apex ofVinca major, both the orientation and frequency of cell divisions,and the pattern of cell expansion, were observed using a non-destructivereplica technique. Micrographs taken at daily intervals illustratethat the central region of the apical dome remains relativelyinactive, except for a phase of cell division which occurs after2 d of growth. The majority of growth takes place at the proximalregions of the dome from which develop the successive pairsof leaves. The developing leaf primordia are initiated by aseries of divisions which occur at the periphery of the centraldome and are oriented parallel to the axis of the subsequentleaves. The cells which develop into the outer leaf surfaceof the new leaves undergo expansion and these cells divide allowingfor the formation of the new leaf. This paper describes thefirst high-resolution sequential study of cell patterns in asingle developing plant apex. Sequential development, cell division, expansion patterns, SEM, Vinca major, apical dome, leaf primordium, leaf initiation     

In Vitro Culture of a Root Parasite, Aeginetia indica L. II. The Plane of Cell Division in the Tendril

Factors affecting septation (cell division) of the tendril whichfacilitates the organic connection with the host were studiedin a root parasite Aeginetia indica L. Transverse cell division,which occurs perpendicular to the long axis of the tendril,was promoted by additions of sucrose, glucose and cytokininsto the basal medium. Longitudinal cell division of the tendril,which takes place parallel or obliquely to the long axis, wasstimulated by cytokinins, but not by sucrose. The latter typeof cell division was frequent in basal and sub-basal cells ofthe tendril but was extremely rare in apical cells. The orientationof the planes of these cell divisions was closely related tocell shape. Abnormal growth of the tendril was seen in germinatingseeds grown for six weeks or more in media containing both Miscanthus(a host) root extract and cytokinin. (Received February 23, 1984; Accepted June 12, 1984)     

Microtubules in the epidermal cells of Carausius morosus (Br.). Their pattern and relation to pigment migration

The existence of one or several systems of microtubules, consisting of a central bundle that branches off towards the basal and distal ends of epidermal cells of Carausius morosus has been shown by indirect immuno-fluorescence microscopy using monospecific anti-tubulin. The pattern of microtubules coincides with the position of the ommochrome granules and their migration path during physiological colour change. The ommochrome granules stick to isolated bundles of microtubules. Small bundles of microtubules extend almost perpendicularly towards the apical cell membrane where they are attached. Distally they are covered by pore channels within the cuticle. Parallel to the basal cell membrane and in close contact with it, there are very small bundles of microtubules. These findings strongly support the idea that during physiological colour change the ommochrome granules migrate along a firmly fixed system of microtubules.     

Cell Division in Tetraspora

Cell division in Tetraspora sp. is described. The cell becomesimmotile some while before mitosis and the basal bodies withdrawfrom the cell surface. The preprophase nucleus migrates to thebasal body complex, around which increasing numbers of microtubulesgather. The spindle is closed with open polar fenestrae; a basalbody complex is always closely associated with at least onepole. No spindles were observed to have basal bodies at bothpoles, and the spindle may possibly be unicentric. During anaphase,spindle microtubules penetrate through the fenestrae. Aftertelophase, the nuclei come together as a phycoplast forms betweenthem; cytokinesis is effected by furrowing. Forming basal bodiesare frequently encountered in late telophase and cleaving cells;no evidence was obtained that the basal bodies replicated beforemitosis. The protoplast rotates inside the cell wall duringcleavage. Cell division is compared with that of other greenalgae, and in particular, Chlamydomonas.     

FINE STRUCTURE OF THE MALE AND FEMALE GAMETES OF ATRACTOMORPHA PORCATA (SPHAEROPLEACFAE,CHLOROPHYTA) WITH EMPHASIS ON THE DEVELOPMENT AND ABSOLUTE CONFIGURATION OF THE FLAGELLAR APPARATUS1

Gametogenesis in Atractomorpha porcata Hoffman was initiated b the synchronous mitotic division of nuclei within a multinucleate gametangium. Uninucleate gametes were subsequently produced following two series of cytokinetic divisions. The first series involved the formation of phycoplast microtubules (phycoplastic cytokinesis), whereas the second series did not (nonphycoplastic cytokinesis). Centrioles were connected by a rudimentary striated distal fiber by the time they migrated to the planes of division preceding the first series of cytokinetic division. These first divisions produced binucleate gametocytes. A well-developed flagellar apparatus lay near the cell surface in close proximity to each nucleus of the gametocyte prior to the second series of cytokinetic divisions that produced the uninucleate gametes. As seen in apical view, the paired basal bodies were directly opposed, with no lateral displacement of their longitudinal axes. In lateral view, the paired basal bodies diverged from one another at an angle of 130–180° (female) or 170–180° (male) and were connected by an arched, distal striated fiber about 670–750 nm long and 600 nm at its widest part. Four electron-opaque, pyramid-shaped lateral bodies flanked the basal bodies in close contact with their undersurfaces. The flagellar roots demonstrated a cruciate arrangement, with s = 6–9 over 1 (female gametes) or 7–10 over 1 (male gametes) microtubules and d= 2 microtubules. In male gametes, one of the multistranded roots was located close to the eyespot, and a second system of cytoskeletal microtubules was detected internally. Based on gamete ultrastructure, Atractomorpha porcata appears to be the most undifferentiated member of the genus.     

Morphology, ontogeny and histochemistry of secretory trichomes of Geranium robertianum (Geraniaceae)

Geranium robertianum bears three types of glandular uniseriate trichomes which originate from a single protodermal cell and develop through periclinal divisions. Type I trichomes are procumbent and have an oval apical cell, two stalk cells and a basal cell. Type II trichomes are erect and have a pear shaped apical cell, two stalk cells and a basal cell. Type III trichomes are much longer than the other two types and have an elongated apical cell, five long stalk cells and a basal cell. Type I and type II trichomes are common on leaves while III trichomes are more abundant on flower structures.
Type I and type II trichomes secrete terpenoids and phenols. Type III trichomes are characterized by the accumulation of anthocyanins in the apical cell and secrete flavonoids.     

Fine structure of protonemal apical cells of the mossPhyscomitrium turbinatum

Summary Elongating caulonemal apical cells of the mossPhyscomitrium turbinatum were cultivatedin vitro and observed during successive stages of cell elongation and division. Actively-growing cells which had completed approximately half of their growth in length were examined by electron microscopy. The distribution of many organelles changes progressively from the cell tip to the distal edge of the large basal vacuole, establishing an apical-basal gradient in organization. Whereas the vacuoles become progressively more extensive in more mature parts of the cell, the dictyosomes, chloroplasts and smooth endoplasmic reticulum are more numerous in younger regions. Some mitochondria in the younger regions of the cell contain localized areas of membrane invagination. Attempts were made to clarify the origin and growth of vacuoles, which become increasingly prominent as the apical cell elongates.Morphological evidence suggests that vacuoles arise in close association with endoplasmic reticulum and dictyosomes as a result of ER dilation and/or cytoplasmic sequestration. The number of vacuolar profiles is highest at the cell tip, decreasing progressively toward the base of the cell, Conversely, the mean area of vacuolar profiles increases progressively toward more basal regions of the cell. These features, along with the increasing number of closely grouped vacuolar profiles along an apical-basal gradient are compatible with the concept of vacuolar growth by coalescence, culminating in their union with the basal vacuole.