Silica Deposition in Some C3 and C4 Species of Grasses, Sedges and Composites in the USA

We report new information on silica deposition in 15 plant species,including nine grasses, two sedges and four composites. Thesilica depositional patterns found in seven of the grass speciesindicate that they are C₄ plants. However the festucoid grassCortaderia selloana is a C₃ plant with long leaf trichomes andoval silica structures in the leaves. In contrast the panicoidC₄ grasses Chasmathium latifolium, Chasmathium sessiflorum,Imperata cylindrica, Panicum repens, Panicum commutatum andSetaria magna, all produce dumb-bell-shaped silica structuresin the leaves. The chloridoid grasses Spartina patens and Spartinacynosuroides have saddle-shaped structures and no dumb-bellor oval shaped ones. The sedges Rhynchospora plumosa and Scirpuscyperinus were found to have oval phytoliths and may be C₃ plants.Our examination of these and other grasses strongly suggeststhat C₄ grasses tend to produce the same type of silica cells.Grasses and sedges with C₃ type photosynthesis tend to produceoval silica structures. The composite Grindelia squarrosa andsunflowers Helianthus angustifolia, Helianthus atrorubens andHelianthus tuberosus absorb relatively small amounts of siliconand larger amounts of calcium, where both elements deposit inleaf trichomes. We found no clear indicator for the C₃ sunflowersor C₄ types in the Asteraceae. Helianthus tuberosus leaves havemany trichomes on the adaxial surface. These trichomes havea higher concentration of silica than the surrounding leaf surface.Helianthus tuberosus leaves had much higher ash and silica contentsthan those of Helianthus angustifolia and Helianthus atrorubens.The composite Grindelia squarrosa has a usual deposition ofsilica in the basal cells around the guard cells. Silica depositionoften reflects the surface features of a leaf. An exceptionis Scripus cyperinus where the silica structures are deep inthe tissue and do not reflect the surface configurations. Theinforescence of Setaria magna had a 14.64 silica content. Thetufts of white, silky hairs characteristic of Imperata cylindricainflorescence have no silica. C₃ and C₄ plants, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, spectra of elements in plants, trichomes, silica fibres, phytoliths     

Silica and Ash in Seeds of Cultivated Grains and Native Plants

Silica and ash contents and the depositional patterns of opalinesilica have been determined in the seeds of 31 plant species.Included were 13 monocotyledons, eight dicotyledons and theseeds of eight common cereal grains. The cereal grains, exceptfor Oryza sativa L. (3.2%) and Avena sativa L. (1.4%), werequite low in silica. The silica in these cereals was in thelemma. In seeds with high silica content it often makes up morethan 50% of the ash. Silica in seeds occurs largely in the outercoating of the seed. Dicotyledon seeds tend to have less silicathan those of monocotyledons. Energy-dispersive X-ray analysisshows that the distribution of the element silicon is clearlyrelated to certain epidermal structures such as ridges, raisedareas, trichomes and hairs. It also occurs in cell walls. Membersof a specific plant family tend to have very similar silicadepositional patterns in their seeds. Small amounts of K, S,Cl and Ca are also found in seeds. Light-microscopy studiesshow that the silica in the lemma of seeds such as Oryza sativaL. is deposited in cellular sheet-like structures with crenateedges. Silica in seeds also occurs in fibres and in other cellularstructures (silica cells) that become phytoliths. Seeds, epidermis, seed coat, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silica depositional patterns, trichomes     

Silica and Ash in Native Plants of the Central and Southeastern Regions of the United States

Ash and silica contents and depositional patterns were determinedfor different tissues of 11 plants growing in the southeasternand central parts of the USA. Silica content was high in theleaves, sheaths and inflorescences of the grasses studied, especiallyso in the inflorescence of the C₃ grass, Stipa comata Trise.and Rupr. The ash content was especially high in leaves of Polymniauvedalia L., which are also high in calcium. Calcium depositionwas largely in trichomes and in veins of the leaf. Energy-dispersiveX-ray analysis showed that the distribution of the element siliconis closely related to certain epidermal structures such as ridges,cell walls, rows of irregularly-shaped structures lying lenghthwisealong the leaf, dumb-bell shaped structures and trichomes. Thesestructures also correspond to the phytoliths left behind afterdecay of the plant. The C₃ grasses differed from the C₄ in thatthey showed oval structures and produced correspondingly ovalphytoliths. Silicified trichomes (particularly in the C₃ grasses)and long, narrow, silica fibres were common in the inflorescencesof the grasses studied. These sharp particles could be irritatingto oesophageal and other tissues. Similar fibres in other grasseshave been implicated in certain cancers. High silicificationof the inflorescence structures might afford protection forthe seed, as reported for other grasses. C₃ and C₄ grasses, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, spectra of elements in plants, trichomes, silica fibres, phytoliths     

Silica and Ash in Tissues of Some Plants Growing in the Coastal Area of Mississippi, USA

Ash and silica contents and their depositional patterns in differenttissues of 27 plants growing in the Ocean Springs area of Mississippi(many grow elsewhere), were determined. Silica content of driedplant parts varied from no more than 0 per cent in Borrichiafrutescens (L.) D.C. stems to 18.76 per cent in Arundinariagigantea (Walt.) Muhl. leaves. Ash content varied from 0·73per cent in Cliftonia monophylla (Lam.) Britt. ex Sarg. stemsto 44·02 per cent in Batis maritima L. leaves. Plantssuch as Batis maritima L., Borrichia frutescens (L.) D.C., Salicorniabigelovii Torr. and Salicornia virginica L. which grew in salinemarshes had high ash contents due NaCl in their tissues. Morusrubra L. leaves had a high silica content for a dicotyledonousplant (3·12 per cent). Energy-dispersive X-ray analysisshows that the distribution of the element silicon is clearlyrelated to certain epidermal structures such as ridges, cellwalls, rows of irregular shaped structures lengthwise of theleaf, dumb-bell shaped ones and especially in trichomes. Therewas a high concentration of silica containing trichomes alongthe veins on the underside of Morus rubra L. leaves and thiswould protect them from insects. The outer parts of the inflorescencesof Ctenium aromaticum (Walt.) Wood, Elymus virginicus L., Juncuspolycephalus Michx. and phragmites communis Trin. were highlysilicified. This should give the seed some protection from insects.The sharp particles would be irritating to oesophageal tissuesand might be implicated in such a cancer.     

TINY BRANCHED HAIR functions in multicellular trichome development through an ethylene pathway in Cucumis sativus L.

The fruit trichomes of Cucurbitaceae are widely desired in many Asian countries and have been a key determinant of cucumber (Cucumis sativus L.) cultivar selection for commercial production and breeding. However, our understanding of the initiation and development of cucumber trichomes is still limited. Here, we found that the cucumber TINY BRANCHED HAIR (TBH) gene is preferentially expressed in multicellular trichomes. Overexpression of CsTBH in tbh mutants restored the trichome phenotype and increased the percentage of female flowers, whereas silencing of CsTBH in wild-type plants resulted in stunted trichomes with a lower rate of female flowers. Furthermore, we provide evidence that CsTBH can directly bind to the promoters of cucumber 1-Aminocyclopropane-1-Carboxylate Synthase (CsACS) genes and regulate their expression, which affects multicellular trichome development, ethylene accumulation, and sex expression. Two cucumber acs mutants with different trichome morphology and sex morphs compared with their near-isogenic line further support our findings. Collectively, our study provides new information on the molecular mechanism of CsTBH in regulating multicellular trichome development and sex expression through an ethylene pathway.     

HECATE2 acts with GLABROUS3 and Tu to boost cytokinin biosynthesis and regulate cucumber fruit wart formation

Warty fruit in cucumber (Cucumis sativus L.) is an important quality trait that greatly affects fruit appearance and market value. The cucumber wart consists of fruit trichomes (spines) and underlying tubercules, in which the existence of spines is prerequisite for tubercule formation. Although several regulators have been reported to mediate spine or tubercule formation, the direct link between spine and tubercule development remains unknown. Here, we found that the basic Helix-Loop-Helix (bHLH) gene HECATE2 (CsHEC2) was highly expressed in cucumber fruit peels including spines and tubercules. Knockout of CsHEC2 by the CRISPR/Cas9 system resulted in reduced wart density and decreased cytokinin (CTK) accumulation in the fruit peel, whereas overexpression of CsHEC2 led to elevated wart density and CTK level. CsHEC2 is directly bound to the promoter of the CTK hydroxylase-like1 gene (CsCHL1) that catalyzes CTK biosynthesis, and activated CsCHL1 expression. Moreover, CsHEC2 physically interacted with GLABROUS3 (CsGL3, a key spine regulator) and Tuberculate fruit (CsTu, a core tubercule formation factor), and such interactions further enhanced CsHEC2-mediated CsCHL1 expression. These data suggested that CsHEC2 promotes wart formation by acting as an important cofactor for CsGL3 and CsTu to directly stimulate CTK biosynthesis in cucumber. Thus, CsHEC2 can serve as a valuable target for molecular breeding of cucumber varieties with different wart density requirements.

Fruit wart is a key external quality trait, and CsHECATE2 promotes wart formation by interacting with a spine regulator and a tubercule factor to directly stimulate cytokinin biosynthesis in cucumber     

Structure and Function of Silica Bodies in the Epidermal System of Grass Shoots

Silica (SiO₂.nH₂O) is deposited in large quantities in the shootsystems of grasses. In the leaf epidermal system, it is incorporatedinto the cell wall matrix, primarily of outer epidermal walls,and within the lumena of some types of epidermal cells. This biogenic silica can be stained specifically with methylred, crystal violet lactone, and silver amine chromate. At theultrastructural level, the silica in lumens of silica cells,bulliform cells and long epidermal cells is made up of rodsabout 2.5 µm in length and 0.4µm in width. Ultimateparticles in the rods range from 1 to 2 nm in diameter. In contrast,silica in the cell wall matrix of trichomes and outer wallsof long epidermal cells is not rod-shaped, but rather, formsroughly spherical masses. Detailed analyses are presented on the frequencies of occurrenceof the different types of epidermal cells that contain silicain the leaves of representative C₃ and C₄ grasses. The C₄ grasseshave higher frequencies of bulliform cell clusters, silica cells,and long epidermal cells, whereas the C₃ grasses have higherfrequencies of trichomes. No correlation was found in the frequencyof occurrence of silica bodies in bulliform cells for C₃ grassesas compared with C₄ grasses. Of all the grasses examined, Coix,Oryza, and Eleusine had the highest densities of such bodies,and some taxa had no silica bodies apparent in their bulliformcells. The idea that silica bodies in bulliform cells and silica cellsmight act as "windows’ and trichomes might function as‘light pipes’ to facilitate light transmission throughthe epidermal system to photosynthetic mesophyll tissue belowwas tested. The experimental data presented do not support eitherof these hypotheses. C₂ and C₄ grasses, biogenic silica, light pipes, window hypothesis, silica staining, silica ultrastructure     

Epidermal Structure in the Ligule of Rice (Oryza sativa L.)

The structure of the abaxial epidermis of the ligule of rice(Oryza sativa L.) as seen in the light and scanning electronmicroscope is described. Long cells, silica cells, cork cells,prickle hairs, microhairs and stomata-like structures were found.Reasons for considering these latter structures to be true stomataare given. Results of an X-ray analysis for silicon are presentedand the distribution of silica discussed in relation to itsfunctional role in the ligule. Oryza sativa L, rice, Gramineae, ligule, epidermis, scanning electron microscopy, X-ray analysis, silica, stomata     

Silica and Ash in Tissues of Some Coastal Plants

Ash and silica contents and their depositional patterns in differenttissues of 44 Mississippi coastal plants were determined. Silicacontent of dried plants varied from no more than a trace inChenopodium album L. leaves to 7.37 per cent in Zizanopsis miliacea(Michx) Doell & Aschers leaves. Ash content varied from2.50 per cent in Lythrum lineare L. stems to 28.24 per centin Borrichia frutescens (L.) DC leaves. Plants in the same familytend to be alike in their ability to absorb or not absorb silica.Poaceae and Cyperaceae had consistently high concentrationsof silica. In contrast, the Asteraceae studied had very lowsilica contents but often had high contents of other minerals.Dicotyledonous plants studied had consistently lower silicacontents than the monocotyledons. Plants growing in salt watercontained considerable sodium chloride. Spectra were obtainedfor major elements in four different plants. Energy-dispersiveX-ray analysis shows that distribution of the element siliconis clearly related to certain epidermal structures such as guardcells, ridges, dumb-bells and balls that appear in electronmicrographs. Silica was deposited differently in each type ofplant studied. In many of the plants silica was deposited inrows of irregular-shaped particles running lengthwise of theleaf and in guard cells. In others, like Zizanopsis miliacea(Michx) Doell & Aschers, the deposit was sheet-like. Zizaniaaquatica L. not only had a sheet-like deposit, but the depositwas ridged and there were rows of dumb-bell-shaped silica cells.Related plants had similar structures. Euchlaena mexicana Schrad.,Tripsacum dactyloides (L.) L and Manisuris rugosa (Nutt.) Kuntzeall had irregular phytoliths similar to those in Zea mays L. coastal plants, marsh plants, ash content, silica deposition, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, X-ray diffraction patterns, spectra of elements in plants     

Silica and Ash Content and Depositional Patterns in Tissues of Mature Zea mays L. Plants

Ash and silica content and their depositional patterns in differenttissues of the mature corn plant (Zea mays L.) were determined.Ash and silica were highest in the leaf blades (up to 16.6 and10.9 per cent, respectively) followed by the leaf sheath, tassel,roots, stem epidermis and pith, and ear husk. The percentageof ash as silica was also highest in the leaves. Silica wasextremely low in the kernels. The upper stem epidermis and pithcontained nearly twice the silica content as did the lower portion.The patterns of ash and silica distribution were similar inplants grown in two different areas of Kansas, but were in lowerconcentration in the leaves and leaf sheaths from the area withlower soluble silica in the soil. Silica was deposited in theepidermis in a continuous matrix with cell walls showing serratedinterlocking margins in both leaves and stem. Rows of lobedphytoliths of denser silica were found in the epidermis as wellas highly silicified guard cells and trichomes. The silica matrixof the epidermis appears smooth on the outer surface and porousor spongy on the inner surface. Zea mays L. Corn, maize, ash content, silica deposition, scanning electron microscopy     

A New Histochemical Method for Localization of the Site of Monoterpene Phenol Accumulation in Plant Secretory Structures

A new method is reported for the histochemical localizationof monoterpene phenols in essential oil secretory structures.The method was adapted from a spot test originally devised forin vitro detection of phenolic compounds in organic analyses.Plant subjects were the Lamiaceae species Thymus vulgaris L.,Oreganum vulgare L. and Mentha x piperita L., which accumulateessential oil in glandular trichomes. A reagent consisting of4-nitrosophenol in conc. H₂SO₄was applied to sample leaves ofeach species. A positive test for phenol was indicated by theproduction of coloured indophenols. Using this method, monoterpenephenols were identified in the trichomes of T. vulgaris(thymol)and O. vulgare(carvacrol), indicated by colour changes to redand green respectively. No phenol was detected in trichomesof M. x piperita. Results were confirmed by GC-MS analysis ofleaf volatile extracts from each species, and in vitro testswith thymol and carvacrol. The method could be used in fieldsurveys for rapid identification of potential medicinal plantsand bioactive compounds. Copyright 2001 Annals of Botany Company Histochemistry, secretory structures, glandular trichomes, Lamiaceae, Thymus vulgaris, Oreganum vulgare, Mentha x piperita, essential oil, aromatic monoterpenes     

SILICATE UPTAKE AND SILICA SHELL FORMATION BY SYNCHRONOUSLY DIVIDING CELLS OF THE DIATOM NAVICULA PELLICULOSA (BRÉB.) HILSE1,2,3

Cells of the freshwater diatom Navicula pelliculosa were induced to divide synchronously by the addition of excess silicate (20 mg Si/liter) to a culture which had been silicate-starved for 14 hr. After a 3- to 4-hr lag period, characterized by rapid silicate uptake (6.5 mg Si/liter/4 hr) and silica shell formation, the cell number then doubled in the succeeding 2-3 hr. This system promises to have significance in the study of silicon metabolism, transport phenomena, and silica deposition.     

Silicon Distribution and Anatomy of the Grass Rhizome, with Special Reference to Miscanthus sacchariflorus (Maxim.) Hackel.

Rhizome anatomy is described for Miscanthus sacchariflorus (Maxim.)Hackel. Solid silica deposits, detected as elemental siliconby electron-probe microanalysis and energy-dispersive X-rayanalysis, are confined to cell walls of three concentric zonesconsisting of the uniseriate epidermis, and parenchyma layersaround the cortical air lacunae, and the central cavity, respectively.Si is localized in outer tangential walls of the epidermis,while occurring in all walls of nucleated, parenchyma cellsforming the two internal zones. In comparison, the root exhibitsonly one Si zone. Rhizome Si distribution more closely resemblesthat for Phragmites australis, than for related members of theAndropogoneae. P. australis similarly exhibits aerenchyma anda central cavity. Thus, internal anatomy may strongly influencesilicon distribution. A comparison of taxa of four tribes indicatesthat epidermal wall deposition is common, followed by specificinternal localization in up to three zones of perivascular tissues. Silicon accumulation occurs early in the epidermis of the youngapex of M. sacchariflorus, decreasing sharply across an internodetransection. In comparison, the oldest, basal internodes exhibitvery high Si X-ray counts in each of the three zones, the highestoccurring in the most internal zone around the central cavity.Early Si mobilization in the rhizome apex may resist shearingand abrasion during horizontal growth extension, while depositsbordering aerenchyma of older internodes may resist compression. Miscanthus sacchariflorus (Maxim.) Hackel, plume grass, rhizome, silicification, anatomy, aerenchyma     

Biogenic Opal in Three Species of Gramineae

Particulate biogenic opaline silica is concentrated in cellwalls, intercellular deposits and cell lumina of all portionsof the above-ground plant body of three species of PanicoidGramineae,Andropogon gerardi, Sorgastrum nutans and Panicumvirgatum. Morphologically distinct opal phytoliths form notonly in long cells, short cells, trichomes, stomatal elementsand bulliform cells of the epidermis but also within the cellularstructure of mesophyll, vascular, and sclerenchyma tissues.Roots and rhizomes contain measurable quantities of opalinesilica, and phytoliths develop in epidermal long cells, saddle-shapedshort cells, vascular cells, and intercellular deposits. A morphologicallyunique plate-phytolith, formed by silicification of the innertangential wall of the endodermis, is present in the roots ofall three species. Differences in the quantity of opaline silicaoccur between species and between parts of the same species.The amount of opal deposited in the soil annually by root systemsand above-ground parts is approximately equal in magnitud Andropogon gerardi, Sorgastrum nutans, Panicum virgatum, opaline silica deposition     

Glandular Trichomes on the Leaves and Flowers of Plectranthus ornatus: Morphology, Distribution and Histochemistry

The types of glandular trichomes and their distribution on leavesand flowers of Plectranthus ornatus were investigated at differentstages of their development. Five morphological types of glandulartrichomes are described. Peltate trichomes, confined to theleaf abaxial surface, have, in vivo, an uncommon but characteristicorange to brownish colour. Capitate trichomes, uniformly distributedon both leaf surfaces, are divided into two types accordingto their structure and secretory processes. In long-stalkedcapitate trichomes, a heterogeneous secretion (a gumresin) isstored temporarily in a large subcuticular space, being releasedby cuticle rupture, whereas, in the short-stalked capitate trichomes,the secretion, mainly polysaccharidic, is probably exuded viamicropores. On the leaves, digitiform trichomes, which do notshow a clear distinction between the apical glandular cell andthe subsidiary cells, occur with a similar distribution to thecapitate trichomes. They do not develop a subcuticular space,and secrete small amounts of essential oils in association withpolysaccharides. The reproductive organs, particularly the calyxand corolla, exhibit, in addition to the reported trichomes,unusual conoidal trichomes with long unicellular conical heads.A large apical pore, formed by tip disruption, releases thesecretion (a gumresin) stored in a rostrum-like projection.On the stamens and carpels, digitiform, capitate and conoidaltrichomes are absent, but peltate trichomes are numerous. Theyoccur between the two anther lobes, on the basal portion ofthe style, and between the four lobes of the ovary. The resultspresented are compared with those of other studies on Lamiaceaeglandular trichomes. Copyright 1999 Annals of Botany Company Plectranthus ornatus Codd, Lamiaceae, glandular trichomes, morphology, histochemistry, essential oils and mucilage secretion.     

A New Glabrous Gene (csgl3) Identified in Trichome Development in Cucumber (Cucumis sativus L.)

Spines or trichomes on the fruit of cucumbers enhance their commercial value in China. In addition, glabrous mutants exhibit resistance to aphids and therefore their use by growers can reduce pesticide residues. Previous studies have reported two glabrous mutant plants containing the genes, csgl1 and csgl2. In the present study, a new glabrous mutant, NCG157, was identified showing a gene interaction effect with csgl1 and csgl2. This mutant showed the glabrous character on stems, leaves, tendrils, receptacles and ovaries, and there were no spines or tumors on the fruit surface. Inheritance analysis showed that a single recessive gene, named csgl3, determined the glabrous trait. An F₂ population derived from the cross of two inbred lines 9930 (a fresh market type from Northern China that exhibits trichomes) and NCG157 (an American processing type with glabrous surfaces) was used for genetic mapping of the csgl3 gene. By combining bulked segregant analysis (BAS) with molecular markers, 18 markers, including two simple sequence repeats (SSR), nine insertion deletions (InDel) and seven derived cleaved amplified polymorphism sequences (dCAPs), were identified to link to the csgl3 gene. All of the linked markers were used as anchor loci to locate the csgl3 gene on cucumber chromosome 6. The csgl3 gene was mapped between the dCAPs markers dCAPs-21 and dCAPs-19, at genetic distances of 0.05 cM and 0.15 cM, respectively. The physical distance of this region was 19.6 kb. Three markers, InDel-19, dCAPs-2 and dCAPs-11, co-segregated with csgl3. There were two candidate genes in the region, Csa6M514860 and Csa6M514870. Quantitative real-time PCR showed that the expression of Csa6M514870 was higher in the tissues of 9930 than that of NCG157, and this was consistent with their phenotypic characters. Csa6M514870 is therefore postulated to be the candidate gene for the development of trichomes in cucumber. This study will facilitate marker-assisted selection (MAS) of the smooth plant trait in cucumber breeding and provide for future cloning of csgl3.     

Opaline Silica Deposits in the Leaves of Bidens pilosa L. and their Possible Significance in Cancer

The occurrence of silica deposits in suitably treated leavesof Bidens pilosa L. of African origin was investigated usinglight and scanning electron microscopy and X-ray analyses. Numerousfibres of amorphous silica were located at the edges and surfacesof the leaves. The deposits were always associated with theepidermal microhairs and were confined to the walls. The significance of these accumulations in a dicotyledonousspecies and in relation to possible deposition mechanisms isdiscussed. Their possible role as a cause of cancer is reviewedin the light of recent investigations involving Bidens pilosa. Bidens pilosa L., opaline silica, oesophageal cancer     

An Analysis of the Mussell and Morre Quantitative Bioassay for Polygalacturonases using Pectate trans-eliminase from Erwinia atroseptica

The Mussell and Morré method for estimating the activityof chain-splitting pectic enzymes was tested with sections ofparenchyma from cucumber fruit and potato tubers. It provedto be a very sensitive test for the pectate trans-eliminaseof Erwinia atroseptica, between 10 and 100 times as sensitiveas a viscometric method based on degradation of polypectate.Loss of fresh weight was readily detected within a few minutesafter immersion in enzyme solution and continued until the sectionof tissue had lost coherence. For some time after immersion,loss of weight was mainly due not to loss of cells from thesurface of the section but to loss of water and solutes, presumablybecause of the effect of the trans-eliminase in decreasing thepermeability of the protoplasts. In the later stages loss ofcells became more important. Potato sections lost weight atabout half the rate of cucumber sections but the rate of increaseof absorbance at 235 nm of the ambient solution was about athird higher for potato than for cucumber. Potato sections lostelectrolytes much more rapidly than did cucumber sections. Thesedifferences probably reflect, at least in part, the much higherwater content of cucumber sections. The advantages of the Mussell and Morré method are itssensitivity and the fact that the natural substrates of chain-splittingenzymes can be used for assessing activity.     

Glandular Trichomes on Vegetative and Reproductive Organs of Leonotis leonurus (Lamiaceae)

The types of glandular trichomes, their ontogeny and patternof distribution on the vegetative and reproductive organs ofLeonotis leonurus at different stages of development, are studiedby light and scanning electron microscopy. Two morphologicallydistinct types of glandular trichomes (peltate and capitate)are described. Peltate trichomes, at the time of secretion,are characterized by a short stalk, which is connected witha large spherical head composed of eight cells in a single layer.Capitate trichomes can be divided into various types. Generally,they consist of a four-celled head supported by one or threestalk cells. The two kinds of trichomes differ in the secretionprocess. In the peltate trichomes, the secretory product seemsto remain accumulated in a subcuticular space, unless an externalfactor damages it. In the capitate trichomes, this product probablybecomes released through micropores. On the leaves peltate andcapitate trichomes are abundant, while on the flowers only thepeltate trichomes are numerous and the capitate are rare orabsent.Copyright 1995, 1999 Academic Press Leonotis leonurus R. Br., lion's ear, lion's tail, Lamiaceæ, glandular trichomes, morphology, ontogeny     

Glandular Trichomes of Fagonia L. (Zygophyllaceae) Species: Structure, Development and Secreted Materials

The glandular trichomes ofFagoniaconsist of one secretory celland a multicellular stalk, which develops by division, elongationand elevation of epidermal cells. The latter become seperatedfrom the mesophyll and a subepidermal chamber is formed. Thelength of the stalk, which differs in the various species orvarieties is determined by the number of cell divisions and/or the extent of cell elongation. Although the basic morphologyand development of the trichomes of the species and varietiesexamined are similar, two types of mature trichomes can be distinguished:one occurs in the two examined varieties ofF. mollisand thesecond inF. glutinosaandF. arabica. The secretory cells of thesecond type possess a very thick wall and bear a porous cupuleon their top. Histochemical tests revealed that the sticky substancesecreted by the secretory cells contains mainly polysaccharidesand lipophilic compounds. The secreted material exhibits autofluorescence.InF. mollisvar.hispidachanges in the amount and shape of thefluorescent material inside the secretory cell, during the courseof development, have been observed. The contribution of theglandular trichomes inFagoniaspecies to survival in hot desertconditions is discussed. Fagonia; glandular trichomes; subepidermal chamber; secreted material; adaptation to desert conditions; stalk; fluorescence