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     

Development and Liberation of Cauline Gemmae in the Moss Aulacomnium androgynum (Hedw.) Schwaegr. (Bryales): An Ultrastructural Study

The leafy shoots of the mossAulacomnium androgynum form clustersof gemmae borne terminally on long pseudopodial axes. The gemmaearise from single initial cells produced by the activity ofa superficial meristem. Mature gemmae comprise an apical anda basal cell with four to seven cells forming two, sometimesthree, tiers in between. The basal cell is connected to thetip of the pseudopodium by a uniseriate filament consistingof an abscission (tmema) cell and a stalk cell. The first divisionof the initial cell produces a proximal cell and a distal cell.The proximal cell elongates without further division formingthe stalk of the gemma; the distal cell gives rise to a lowerand an upper cell by transverse division. The upper cell dividesrepeatedly by oblique septa forming the apical and middle cellsof the gemma; the basal cell and tmema cell arise from a transversedivision of the lower cell. The first two divisions in gemmadevelopment are highly asymmetrical and exogenous, i.e. preceededby cell expansion. A broad interphase cortical band of microtubulesis associated with intercalary cellular growth during this stage.Subsequent gemma development follows an endogenous pattern withcellular expansion following the completion of proliferativedivisions and involving a conventional system of cortical microtubules.While elongating to about four times its original length withoutdeposition of a distinct new wall the tmema cell undergoes cytoplasmicdegeneration and eventually breaks, causing gemma liberation.The stalk cell elongates about eight-fold and its contents alsodegenerate after gemma liberation. Plasmodesmata in the basaland stalk cells are obliterated by the deposition of additionalwall materials. The highly electron-opaque outer walls of themature gemmae and tips of the stalk cells are water-repellant.The gemmae are dispersed either in water films or by air currents. Abscission; asexual reproduction; bryophytes; morphogenesis; microtubules; ultrastructure     

Protonemal Morphogenesis of the Moss Tetraphis pellucida Hedw. in Culture and in the Wild

In axenic culture, the protonemal filaments of Tetraphis pellucidaderived from either spores and gemmae, or excised stems andleaves, share a mixture of attributes of chloronemata and caulonemata.Indirect immunofluorescence reveals that the tip cells containcortical and endoplasmic arrays of microtubules at interphase,and phragmoplasts associated with cell plate formation, butpre-prophase bands are absent. Protonemal plates originate fromthe same sites as filamentous protonemal side branches or directlyfrom young gemmae or excised stem fragments. These plates havea cylindrical base, the latter producing a single gametophorebud, and a unistratose lamina. The gametophores produce gemmacups in culture with the vegetative life cycle taking approximately28 d. Gibberellic acid (GA₃) has no visible effect on protonemal morphogenesiswhereas the auxin naphthalene acetic acid (NAA) suppresses plateand side branch formation. In the presence of kinetin the platesare callus-like and produce supernumerary buds. Abscisic acid(ABA) induces malformed plates and filaments with swollen cells,similar to those found in ageing cultures. Rhizoids are produced in abundance from gametophores and protonemalplates in nature but were never seen in culture. In the wild,rhizoids produce numerous protonemal plates and occasional gametophorebuds. The former are the main source of new shoots. The filamentousprotonemal phase in nature mainly comprises upright filamentscontaining one or more abscission cells. The protonemal plates in Tetraphis are homologous with thosein the allied genus Tetrodontium but are very different fromthose in Diphyscium and Sphagnum. Differences between cultureand nature are attributed to lower nutrient levels and irradiancesin the wild. Tetraphis pellucida, protonema, moss, morphogenesis, immunocytochemistry, gemmae, tip growth, vegetative reproduction     

Thallus Differentiation in the Marchantialean Liverwort Asterella wilmsii (Steph.) with Particular Reference to Longitudinal Arrays of Endoplasmic Microtubules in the Inner Cells

This light and electron microscope study of the liverwort Asterellareveals that, as in other Marchantiales, the cells lining thedorsal air chambers are highly vacuolate with numerous amylochloroplastsin the peripheral cytoplasm. The ventral parenchyma in the midribof the thallus contains aseptate fungal hyphae surrounded byan interfacial matrix and host cytoplasm forming transvacuolarstrands. These are lined with microtubules, rarely seen in otherfungal-hepatic associations or in mycorrhizas. Numerous lipidbodies found in all the thallus cells are thought to be associatedwith perennation during the winter dry season. Elongated, thick walled inner thallus cells, between the dorsalair chambers and the fungus-containing tissue, have a cytologicalorganization not previously recorded in land plants. Initiallyhighly vacuolate, with numerous microtubules of random orientationlining the tonoplast, these cells subsequently show interdigitationof vacuoles and cytoplasm producing a labyrinth of sphericaland elongate tonoplast profiles lined by longitudinal arraysof microtubules. At the same time the cytoplasm becomes increasinglyelectron-lucent and the ribosomes, progressively lost from theER, clump together. At maturity the inner thallus cells arehighly polarized with most of the vacuoles lying nearer thethallus apex. In pits in the end walls, numerous plasmodesmata,with expanded cytoplasmic annuli recall the plamodesmatal fieldsin the mesophyll and phloem of the leaves in vascular plants. Far from being supporting parenchyma or sclerenchyma as assumedhitherto, the inner thallus cells of Asterella are clearly highlydifferentiated. Their vacuole microtubule associations are highlysuggestive of a microtubule-based translocation system akinto that seen in many animal cells and perhaps fungal hyphae,but very different from bulk flow in sieve elements and actin-basedcytoplasmic streaming.Copyright 1994, 1999 Academic Press Cytoskeleton, liverwort, microtubules, plasmodesmata, solute transport     

Scapania praetervisa Meyl., not S. mucronata Buch in Britain

Abstract

The protonema of Orthotrichum obtusifolium, regenerated from leafy shoots, comprises a dense mass of largely unbranched exclusively upright filaments. All the cells have transverse cross-walls and contain numerous chloroplasts which become larger and discoidal in older filaments. Differentiation into chloro- and caulonema is absent. When upright filaments are transplanted onto an agar surface, they act as the main axes for a further set of upright branches. Buds and side branches are produced symmetrically at right angles to the mother cells. Protonemal morphogenesis hardly changes on nutrientfree medium and spherical brood cells are not produced after prolonged culture or in the presence of abscisic acid. Rhizoids with oblique septa and pigmented walls arise from the base of gametophores. These revert to protonemata when in contact with nutrient agar.

The upright filaments produce terminal and intercalary filamentous gemmae with well defined abscission cells. The latter swell and lose their contents prior to liberation. Cells immediately below the abscission cells produce side branches in acropetal succession. Secondary gemmae arise by percurrent proliferation and gemmae frequently germinate in situ.     

Targeted destruction of fungal structures of Erysiphe trifoliorum on flat leaf surfaces of Marchantia polymorpha

In this study, we observed the germination behaviour of airborne conidia from powdery mildews that settle on thalloid surfaces. We inoculated thalli (flat, sheet‐like leaf tissues) and gemmae (small, flat, sheet‐like leaf tissues that propagate asexually via bud‐like structures) of the common liverwort (Marchantia polymorpha) with conidia from tomato powdery mildew (Oidium neolycopersici; KTP‐02) and red clover powdery mildew (Erysiphe trifoliorum; KRCP‐4N) and examined their germination and subsequent appressorium formation under a high‐fidelity digital microscope. Conidial bodies and germ tubes of the inoculated KRCP‐4N conidia were destroyed on both the thalli and gemmae. The destruction of these fungal structures was observed only for KRCP‐4N conidia inoculated onto M. polymorpha on both leaf surfaces. No differences in destruction of the KRCP‐4N fungal structures between thalli and gemmae were observed. At 4 h post‐inoculation, destruction of the germ tube tip was observed when it reached the gemmae leaf surface. At 6 h post‐inoculation, the conidial bodies and germ tubes were destroyed. In contrast, KTP‐02 conidia were not destroyed and formed normal, well‐lobed appressoria on the surface of M. polymorpha gemmae.     

An Outline of the Bryophytes of County Laois (Queen's County)

Abstract

Scapania lingulata is reported from five vice-counties in Scotland and from one in Ireland, two gatherings bearing ♂ shoots and one with perianths. It differs from S. scandica in its larger gemmae, from S. praetervisa in its broadly rounded postical leaf lobes, and from both species in its larger leaf cells, more numerous oil bodies and polyploid chromosome complement. Usually it grows on base-deficient earthy banks but also on base-rich rock ledges.     

The Distribution of Silicon Deposits in the Fronds of Pteridium aquilinum L.

The occurrence of silicon in mature fronds of Pteridium aquilinumwas investigated using light and scanning electron microscopyand X-ray micro analysis. The heaviest deposits were locatedin the outer tangential walls of the epidermis of the petiole,but the upper epidermal cells of the costal regions of the laminaealso accumulated small quantities. No discrete bodies were associatedwith these deposits. However, the mid-ribs of the laminae exhibitedsome long siliceous fibres associated with the luminae of thestrengthening tissues. The significance of these accumulationsin relation to deposition mechanisms and possible carcinogeniceffects are discussed. Pteridium aquilinum L., bracken, frond, microanalysis, silica     

Sieve-Element Ontogeny in the Aerial Shoot of Equisetum hyemale L.

The aerial shoots of Equisetum hyemale L. var. affine (Engelm.)A. A. Eat. were examined with the electron microscope as partof a continuing study of sieveelement development in the lowervascular plants. Young E. hyemale sieve elements are distinguishablefrom all other cell types within the vascular system by thepresence of refractive spherules, proteinaceous bodies whichdevelop within dilated portions of the endoplasmic reticulum(ER). Details of cell wall thickening differ between protophloemand metaphloem sieve elements. Following cell wall thickeningthe ER increases in quantity and aggregates into stacks. Shortlythereafter, nuclear degeneration is initiated. During the periodof nuclear degeneration some cytoplasmic components-dictyosomes,microtubules and ribosomes-degenerate and disappear, while organellessuch as mitochondria and plastids persist. The latter undergostructural modifications and become parietal in distribution.Eventually the massive quantities of ER are reduced, leavingthe lumen of the cell clear in appearance. At maturity the plasmalemma-linedsieve element contains a parietal network of tubular ER, aswell as mitochondria, plastids, and refractive sphemh At thistime many of the spherules are discharged into the region ofthe wall. Sieveelement pores occur in both lateral and end walls.At maturity many pores are traversed by large numbers of ERmembranes. The metaphloem sieve elements of the mid-internodalregions apparently are sieve-tube members. The connections betweenmature protophloem sieve elements and pericycle cells are associatedwith massive wall thickenings on the pericyclecell side.     

Fine Structure and Cytochemistry of the Abscission Zone Cells of Phaseolus Leaves: II. Localization of Peroxidase and Acid Phosphatase in the Separation Zone Cells

The sub-cellular localization of acid phosphatase and peroxidaseactivities has been studied by cytochemical procedures in cellsat the surface of the separation layer during the abscissionof leaves of Phaseolus vulgaris. Intense staining for both enzymeswas found in the cell walls, Golgi bodies and endoplasmic reticulum.The wall staining for acid phosphatase was chiefly associatedwith the middle lamellar region while staining for peroxidasewas found throughout the wall. These observations are discussedin relation to the possible involvement of these enzymes inthe changes occurring in the wall during abscission and to therole of the Golgi bodies in the separation process.     

Histochemical Localization of Citral Accumulation in Lemongrass Leaves (Cymbopogon citratus(DC.) Stapf., Poaceae)

Lemongrasses (Cymbopogonspp., Poaceae) are a group of commerciallyimportant C₄tropical grasses. Their leaves contain up to 1.5%(d.wt) essential oils with a typical lemon-like aroma, consistingmainly of citral (a mixture of the isomeric acyclic monoterpenealdehydes geranial and neral). To specifically locate the sitesof citral accumulation in lemongrass we employed Schiff's reagent,which reacts with aldehydes and gives a purple-red colorationwith citral. Using this technique, single oil-accumulating cellswere detected in the adaxial side of leaf mesophyll, commonlyadjacent to non-photosynthetic tissue, and between vascularbundles. Cell walls of these oil cells are lignified. Our resultssuggest that citral accumulation takes place in individual oilcells within the leaf tissues.Copyright 1998 Annals of BotanyCompany Lemongrass;Cymbopogon citratus; Poaceae; oil cells; histochemistry; citral; aldehydes; Schiff's-reagent.     

Aurora B Kinase Activity is Required to Prevent Polar Cortical Ingression during Cytokinesis

Cytokinesis requires proper regulation of microtubule dynamics. It has been suggested that dynamic astral microtubules prevent cortical ingression. However, it remains unknown how astral microtubules maintain their dynamic state. Here we show that aurora B kinase, a component of the chromosome passenger complex, is required to sustain the dynamic state of astral microtubules during cytokinesis. Treatment of HeLa cells with Hesperadin, an inhibitor of aurora B kinase, caused abnormal cortical protrusion, leading to cortical ingression in the protruding region and cytokinesis failure. Actin filaments, myosin II, and RhoA failed to localize at the equator but instead distributed along the lateral and/or polar cortex in cells treated with Hesperadin. Time-lapse analyses of microtubule dynamics showed that, in cells treated with Hesperadin, abnormally bundled astral microtubules targeted the protruding region. Mitotic kinesin-like protein 1 (MKLP1), a component of the spindle midzone required for bundling of microtubules, was not detected along bundled astral microtubules in cells treated with Hesperadin, suggesting that factors other than MKLP1 may be involved in this process. Our results suggest that aurora B kinase activity is required for proper regulation of microtubule dynamics to ensure that cytokinesis occurs precisely at the cell equator.     

Intracellular Silica Deposition in Mature and Senescent Leaves of Sieglingia decumbens (L.) Bernh

Following 1, 2, 4, or 8 weeks of growth in a silica-minimalsolution, tillers of Sieglingia decumbens (Heath Grass) weretransferred for 8 days to a nutrient solution which contained50 or 100 ppm silica. The resultant formation of intracellularopal phytoliths (silica bodies) was compared for the four developmentalstages of leaf 3. Characteristically different phytolith typesoccurred in leaves of different ages. Senescent, in contrastto younger, mature leaves, typically exhibited extensive extracellularsilicification of the mesophyll, in addition to deposits instomatal, long, and bulliform cells of adaxial epidermis; depositionin abaxial silica cells and long cells, characteristic of youngleaves, was much reduced, or absent. Physico-chemical factors and cytoplasmic changes associatedwith senescence are discussed in relation to intracellular opalphytolith formation in mature leaves of S. decumbens. In thisrespect, the tendency of silicic acid sols to polymerize inthe presence of an organic matrix and mineral cations is consideredto be significant.     

The Position of Bud Differentiation on Protonema of the Moss, Physcomitrium sphaericum

The position of the gametophytic bud was examined in relationto the development of protonema in the moss, Physcomitrium sphaericum. Positions of protrusion formation, of the development of protrusionsinto lateral filaments, and of the differentiation of protrusionsinto buds are restricted within the narrow regions of the filaments.The number of cells from the apical cell of the filament tothese positions are constant in any size filament. The growth pattern of the protonema is shown as follow. As afilament grows one-dimensionally through divisions of the apicalcell, new protrusions are produced successively on the 5th cellfrom the apical cell or on its vicinity. The cells which intervenebetween the apical cell and this protrusion increase in numberas the apical cell divides. When this protrusion is positionedat the 8th or 9th cell from the apex, it differentiates intoa bud or a lateral filament. This growth pattern is common toboth the main and lateral filaments. Buds are differentiated not only on caulonema cells in the mainand lateral filament, but also on chloronema cells at the baseof the lateral filaments. (Received December 14, 1981; Accepted April 24, 1982)     

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.     

中国藓类植物无性繁殖体的初步观察

无性生殖在苔藓植物的生活史中起着重要的作用,并且常通过各种无性繁殖体来完成。无性繁殖体的形态常被用来辅助鉴定一些不育的藓类植物。本文通过对38种藓类植物的无性繁殖体进行显微观察,结果显示：无性繁殖体在不同的藓类植物之间已经过多次演化;无性繁殖体的形态在种内是相对稳定的,且与其着生位置、配子体的分枝方式密切相关,而与植物的系统位置以及生境并无直接的关系;无性繁殖体的颜色与其胞壁的厚度以及表面纹饰密切相关。此外,无性繁殖体的产生常常与假根、原丝体有共存关系。在研究中发现,藓类植物的无性繁殖体主要包括原丝体芽胞、无性芽胞（叶生芽胞、中肋芽胞、枝生芽胞）、芽体、假根芽胞和假根状块茎;其中原丝体芽胞和无性芽胞最为常见。     

The oral apparatus of Tetrahymena pyriformis, strain WH-6. IV. Observations on the organization of microtubules and filaments in the isolated oral apparatus and the differential effect of potassium chloride on the stability of oral apparatus microtubules.

This report is an ultrastructural analysis of the organization of the isolated oral apparatus of Tetrahymena pyriformis, strain WH-6, syngen 1. Attention has been focused on the organization of microtubules and filaments in oral apparatus membranelles. Oral apparatus membranellar basal bodies were characterized with respect to structural differentiations at the distal and proximal ends. The distal region of membranellar basal bodies contains the basal plate, accessory microtubules and filaments. The proximal end contains a dense material from which emanate accessory microtubules and filaments. There are at least two possibly three different arrangements of accessory structures at the proximal end of membranellar basal bodies. All membranellar basal bodies appear to have a dense material at the proximal end from which filaments emanate. Some of these basal bodies have accessory microtubules and filaments emanating from this dense material. A possible third arrangement is represented by basal bodies which have lateral projections, from the proximal end, of accessory microtubules and filaments which constitute cross or peripheral connectives. There are at least three examples of direct associations between oral apparatus microtubules and filaments: (1) filaments which form links between basal body triplet microtubules, (2) filaments which link the material of the basal plate to internal basal body microtubules, (3) filaments which link together microtubule bundles from membranellar connectives. KCl extraction of the isolated oral apparatus resulted in the selective solubilization of oral apparatus basal bodies, remnants of ciliary axonemes and fused basal plates. Based on their response to KCl extraction two distinct sets of morphologically similar micro tubules can be identified: (a) microtubules which constitute the internal structure of basal bodies and ciliary axonemes, (b) microtubules which constitute the fiber connectives between basal bodies.     

Changes in Microtubules in Onion Leaf Sheath Cells during Bulb Development

Cortical microtubules are oriented transversely to the cellaxis in leaf sheath cells of onion plants (Allium cepa L. cv.Osaka-Okute) that have not started bulb formation. As the bulbdevelops and the leaf sheath cells swell, the microtubules becomedisoriented and scattered and finally disappear. The microtubuleinhibitors colchicine and cremart [O-ethyl O-(3-methyl-6-nitrophenyl)N-sec-butylphosphorothioamidate]cause swelling of leaf sheath cells and make the basal partof the plant bulbous. The cortical microtubules may have animportant role in regulating bulb development in onion plants. (Received August 21, 1982; Accepted December 6, 1982)     

Comparative Leaf Epidermal Anatomy of Mutants of Barley (Hordeum vulgare L. 'Himalaya') which Differ in Leaf Length

We wished to determine the nature of differences in epidermalcell numbers and dimensions between leaves of different lengthin mutants of barley (Hordeum vulgare L. ‘Himalaya’).Three comparisons were made: leaf one (L1)vs. leaf four (L4);wild typevs. nine dwarf mutants and wild typevs. a slender mutant.L1 was shorter than L4, and for most lines this was associatedwith a change in epidermal cell number for the blade, and inboth cell number and length for the sheath. Compared to wildtype, the smaller leaves of dwarf plants generally had shorterand fewer cells in both blade and sheath. The blade of slenderplants was the same length (L1) or longer (L4) than wild type,while the sheath was longer than that of wild type for bothL1 and L4. Slender plants had longer but fewer cells than thewild type along the blade of L1, and shorter but more cellsfor the blade of L4. In the sheath, slender plants had longerand more (L1) or fewer (L4) cells than did the wild type. ForL1, variation in blade width amongst the barley lines was associatedwith a change in file width and file number. For L4, blade widthvaried only with file number, except for slender plants wherenarrow blades were associated with reduced file width. Hencethere was no consistent correlation between changes in cellsize or cell (or file) number with changes in leaf length orwidth. Differences depended on the leaf (L1vs. L4), leaf part(bladevs. sheath), and the nature of the mutation (dwarfvs.slender). Barley (Hordeum vulgare L. ‘Himalaya’); leaf epidermis; dwarf mutant; slender mutant     

An Ultrastructural Study on the Development of Silicified Tissues in the Leaf Tip of Barley (Hordeum sativum Jess)

The development of silicified tissues in the tip of the fourthand fifth leaf of 7-week-old barley plants grown in nutrientsolution was investigated by transmission electron microscopy.The pattern of silicification was similar in the epidermal cells,sclerenchyma cells and intercellular spaces. Silica granulesfirst appeared around the periphery and eventually filled theentire cell lumen or space. The granules then polymerized toform a solid mass. The walls of the sclerenchyma cells alsobecame silicified but to a less extent than those of the epidermalcells, in which silicification sometimes occurred prior to thatin the lumen. The results are discussed in relation to the chemistry of silicaand possible mechanisms of silicification. Hordeum sativum Jess, barley, silica bodies, opalines, epidermal cells, sclerenchyma, ultrastructure