The Ultrastructure and Electron-probe Microassay of Silicon Deposits in the Endodermis of the Seminal Roots of Sorghum bicolor (L.) Moench

Si deposits in the endodermis of the seminal root of Sorghumbicolor (L.) Moench., following culture in nutrient solutioncontaining 100 ppm SiO₂ for 7 days, were investigated by transmissionelectron microscopy. Si microassay was carried out by meansof the EMMA-4 system. Endodermal ultrastructure is discussed. EM micrographs of theSi deposits reveal information regarding their formative processesand indicate a considerable involvement with the cellulosicstructure of the inner tangential wall (ITW). The initial depositsare believed to be composed structurally of particles designatedas primary spherical units. The EMMA-4 conclusively indicated that Si was localized andconfined to the ITW, and the implications of this result forprevious studies are considered. Current evidence from studies of endodermal function, as wellas root translocation physiology, is utilized in arriving attentative mechanisms for silicic acid transport and subsequentSi deposition on the ITW.     

The Interaction Between Silicon and Aluminium in Sorghum bicolor (L.) Moench: Growth Analysis and X-ray Microanalysis

Seeds of Sorghum bicolor (L.) Moench. were germinated on moistfilter paper for 6 d, before the seedlings were transferredto pots containing 500 µmol l^-1 Ca(NO₃)₂ for 2 d. Theseedlings were then treated with 0 or 100 µmol l^-1 Alin factorial combination with 0, 1400 or 2800 µmol l^-1Si for 8 d. The background solution used throughout was 500µmol l^-1 Ca(NO₃)₂. Aluminium treatment reduced root growthand caused a significant increase in shoot/root ratio. The presenceof silica in the solution significantly ameliorated the effectsof aluminium on root growth. Three treatment were selected for a microanalytical investigationof the basal region of the root: 2800 µmol l^-1 Si only;100 µmol l^-1 Al only; and a combination of the two. Inthe 2800 µmol l^-1 treatment silica was deposited in theendodermis, with the greatest accumulation being in the innertangential wall (ITW). When plants were treated with 100 µmoll^-1 Al only, aluminium concentration was highest in the outertangential wall (OTW) of the epidermis. The element was presentin the hypodermal walls and OTW of the endodermis and was notdetectable in the stele. With both 2800 µmol l^-1 Si and100 µmol l^-1 Al in the nutrient solution the two biomineralizationsites were the ITW of the endodermis, where silicon was themajor element deposited, and atypically in the OTW of the epidermiswhere both aluminium and silicon were present. The sequestrationof aluminium in the Al-Si deposit in the OTW of the epidermismay represent the mechanism that allows greater root growthin this treatment.Copyright 1993, 1999 Academic Press Sorghum bicolor (L.) Moench., aluminium, silicon, calcium, root, toxicity, biomineralization, X-ray microanalysis, freeze substitution     

The Distribution of Silicon Deposits in the Roots of Molinia caerulea (L.) Moench. and Sorghum bicolor (L.) Moench.

The occurrence of silica in relation to meristematic zones andthe thickening of the endodermis in the roots of Molinia caerulea(L.) Moench. and Sorghum bicolor (L.) Moench. has been investigatedby means of the electron-probe microanalyser and the scanningelectron microscope. In proximal regions of mature roots ofM. caerulea, the central strengthening tissue of the stele,the vessel walls, the endodermis and the sub-epidermal sclerenchymaare areas of heavy accumulation. The distal regions of suchroots are relatively free of silicon and show little thickeningof the inner tangential walls of the endodermis or of the cellsof the strengthening tissues. The thickening of these elementsis shown to be associated with their location and the age ofthe root. In the proximal regions of S. bicolor, silicon is detected andlargely confined to the inner tangential walls of the endodermiswhich display some thickening. In addition, discrete and evenly-distributeddeposits varying in size partly fill the lumen of this layer.Some cells exhibit a number of smaller protrusions. High magnificationsof these lumen deposits show a distinct granular structure incontrast to the very uniform pattern of the wall deposits. The results are compared with deposits in grass leaves and inflorescencebracts and in woody perennials. The presence of silicon in additionto suberin, lignin and polyphenols in the thickened endodermalwall is also discussed in relation to the recognized functionof the endodermis.     

X-ray Microanalysis of the Seminal Root of Sorghum bicolor with Particular Reference to Silicon

Sorghum bicolor (L.) Moench. cv. P508.GB plants were grown inwater culture for 1 week, when the seminal roots were harvestedand sampled at five positions starting from the base: 0.0, 0.25,0.50 and 0.75 of the axis length, and a sub-apical position,11 mm behind the tip. Mineral distribution in bulk frozen rootsegments was investigated using SEM and X-ray microanalysis.The elements detected were potassium, chlorine, sulphur, sodium,phosphorus, calcium and silicon. The first four occurred inall root zones. Phosphorus was ubiquitous, but appeared to accumulatein the pericycle protoplasm. Calcium and silicon exhibited themost variation along the seminal axis. Calcium was present inall tissues at the base, but decreased acropetally, being detectedin only the outer cortical and epidermal walls of the subapicalzone. Silicon was present at low levels in protoplasts and wallsof most root tissues, and accumulated in the endodermal protoplastand walls. Deposition in walls is initiated coincident withthe earliest stages of secondary wall thickening. Silicon contentof the inner tangential wall of the endodermis exhibits a decreasingacropetal gradient along the axis length. It is absent frommost cell walls of the sub-apical zone. Silicon pathways inthe root, and silica aggregate formation in relation to thesurrounding ionic environment, are discussed. Sorghum bicolor (L). Moench, seminal root, cryostage, SEM, X-ray microanalysis, ion localization, silicon, endodermis     

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     

The Ultrastructure and Analytical Microscopy of Silicon Deposits in the Roots of Saccharum officinarum (L.)

Silicon accumulation in the endodermis of the ‘set’and ‘shoot’ roots of Saccharum officinarum (L.)were investigated by scanning electron microscopy and electron-probemicroanalysis. Silicon microassay was also carried out by meansof the Corinth analytical microscope (CORA). Aggregates arelargely associated with the inner tangential wall (ITW) of theendodermis and their formation is basically similar to thoseseen in Sorghum bicolor (L.) Moench. In contrast to Sorghumthe earliest deposits in Saccharum appear in wall strata wellwithin the cell wall cytoplasm interface. An additional layerof silicon was also located along the endodermal pericycle boundaryextending some distance along the middle lamella of the radialwalls. The results are discussed in relation to those of previous studiesof silicon accumulation in endodermal cells and to possiblefactors affecting such accumulations.     

Radial Hydraulic Conductivity of Individual Root Tissues of Opuntia ficus-indica (L.) Miller as Soil Moisture Varies

The constraints on water uptake imposed by individual root tissueswere examined forOpuntia ficus-indicaunder wet, drying, andrewetted soil conditions. Root hydraulic conductivity (L_P) andaxial conductance (K_h) were measured for intact root segmentsfrom the distal region with an endodermis and from midroot witha periderm;L_Pwas then measured for each segment with successivetissues removed by dissection. Radial conductivity (L_R) wascalculated fromL_PandK_hfor the intact segment and for the individualtissues by considering the tissue conductivities in series.Under wet conditions,L_Rfor intact distal root segments was lowestfor the cortex; at midroot, where cortical cells are dead,L_Rforthe cortex was higher and no single tissue was the predominantlimiter ofL_R.L_Rfor the endodermis and the periderm were similarunder wet conditions. During 30d of soil drying,L_Rfor the distalcortex increased almost three-fold due to the death of corticalcells, whereasL_Rfor the midroot cortex was unaffected;L_Rforthe endodermis and the periderm decreased by 40 and 90%, respectively,during drying. For both root regions under wet conditions, thevascular cylinder had the highestL_R, which decreased by 50–70%during 30d of soil drying. After 3d of rewetting, new lateralroots emerged, increasingL_Rfor the tissues outside the vascularcylinder as well as increasing uptake of an apoplastic tracerinto the xylem of both the roots and the shoot. The averageL_Rforintact root segments was similar under wet and rewetted conditions,but the conductivity of the tissues outside the vascular cylinderincreased after rewetting, as did the contribution of the apoplasticpathway to water uptake. Opuntia ficus-indica; prickly pear; root hydraulic conductivity; endodermis; periderm; apoplast; lateral root emergence     

Endodermal Silicification in Mature, Nodal Roots of Sorghum bicolor (L.) Moench.

Nodal roots of mature, soil-grown specimens of Sorghum bicolor(L.) Moench. were investigated for their Si content by meansof the electron-probe microanalyser and the scanning electronmicroscope. No consistent accumulation of Si occurred other than in theendodermis. Indication of a lower Si content in the old as comparedto the younger nodal roots was obtained. The solid silica depositsoccurred as domeshaped silica aggregates confined to the innertangential wall. Comparisons with the similar deposits of youngseminal roots are made. It is argued that the endodermal deposits represent a specializedaspect of silicification and contrasting hypotheses to accountfor it are proposed, one involving physico-chemical factors,the other, protoplasmic control.     

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     

Silicification in sorghum (Sorghum bicolor) cultivars with different drought tolerance

Sorghum belongs to a group of economically important, silicon accumulating plants. X-ray microanalysis coupled with environmental scanning electron microscopy (ESEM) of fresh root endodermal and leaf epidermal samples confirms histological and cultivar specificity of silicification. In sorghum roots, silicon is accumulated mostly in endodermal cells. Specialized silica aggregates are formed predominantly in a single row in the form of wall outgrowths on the inner tangential endodermal walls. The density of silica aggregates per square mm of inner tangential endodermal cell wall is around 2700 and there is no significant difference in the cultivars with different content of silicon in roots. In the leaf epidermis, silicon deposits were present in the outer walls of all cells, with the highest concentration in specialized idioblasts termed 'silica cells'. These cells are dumb-bell shaped in sorghum. In both the root endodermis and leaf epidermis, silicification was higher in a drought tolerant cultivar Gadambalia compared with drought sensitive cultivar Tabat. Silicon content per dry mass was higher in leaves than in roots in both cultivars. The values for cv. Gadambalia in roots and leaves are 3.5 and 4.1% Si, respectively, and for cv. Tabat 2.2 and 3.3%. However, based on X-ray microanalysis the amount of Si deposited in endodermal cell walls in drought tolerant cultivar (unlike the drought susceptible cultivar) is higher than that deposited in the leaf epidermis. The high root endodermal silicification might be related to a higher drought resistance.     

Ion Distribution in Salt-stressed Mature Zea mays Roots in Relation to Ultrastructure and Retention of Sodium

Ultrastructural features and the distribution of soluble ionshave been examined in mature roots of Zea mays plants grownin both NaCl and Na₂SO₄ salinities. When the plants were grown in either salt, the Na concentrationincreased proximally along the root with a concomitant declinein the K concentration. Both trends were reversed in the shoot. X-ray microanalysis of deep-frozen, fully hydrated specimensshowed that in salt-treated roots Na, and Cl, or S were distributedabout stoichiometrically in the cortex and endodermis. Na wasusually less concentrated than the anion in the lumens of thevessels, but was concentrated markedly relative to either Clor S in the adjoining xylem parenchyma cells. In the older, proximal parts of seminal roots of plants grownboth without salt (controls) and in the presence of either NaClor Na₂SO₄, wall developments occurred in xylem parenchyma cellsat the half-bordered pits in which the cell wall became markedlythicker and possessed a loosely packed fibrillar structure.These structures were not comparable with the transfer-celltype of protuberances reported in the roots of other species. In the xylem parenchyma of plants grown in the presence of Na₂SO₄there were dramatic increases in the quantities of rough endoplasmicreticulum, ribosomes, and mitochondria relative both to controlsand NaCl treatments. The results are discussed in relation to the possible functionof the xylem parenchyma of the mature root in the reabsorptionof Na from the xylem sap, which may mitigate adverse effectsof salinity in salt-sensitive glycophytes.     

Increased root growth in elevated CO2: a biophysical analysis of root cell elongation

A biophysical analysis of root expansion was conducted in fourchalk downland herbs (Sanguisorba minor Scop., Lotus corniculatusL., Anthyllis vul-neraria L. and Plantago media L.) exposedto either ambient or elevated CO₂in controlled environment cabinets.Measurements of fine (F) and extra-fine (EF) root extensionrate (RER), water relations, and cell wall tensiometric extensibilityrevealed differences in the diurnal pattern of root growth betweenspecies. After 35 d of exposure to elevated CO₂, RER of bothF and EF roots increased significantly in darkness and on illuminationfor S. minor, whilst for A. vulneraria (EF roots only) and L.corniculatus a significant increase occurred at night whereasfor P. media a significant increase occurred during the day.Cells measured in the zone of elongation were longer in allspecies exposed to elevated CO₂. Water potential (     

Anatomy, Ultrastructure and Assimilate Concentrations of Roots of Citrus Genotypes Differing in Ability for Salt Exclusion

Seedlings of Rangpur lime (Citrus reticulata var. austera hybrid?)and Etrog citron (C. medica) were treated in water culture with0, 25, 50 or 100 mol m^–3 NaCl, and in sand culture with0 or 100 mol m^–3 NaCl. Leaf chloride analyses indicatedthat Etrog citron accumulated the most chloride at all levelsof salinity. The structure, ultrastructure and concentrationsof chloride and reserve assimilates of the primary root up to50 mm back from the tip were compared between genotypes andbetween salt treatments. There were no differences in root anatomy in the absence ofsalt between the two genotypes. The hypodermal cells developedlignified and suberized walls which blocked the plasmodesmataand resulted in degeneration of the cell contents. Frequentthin-walled passage cells in the hypodermis had living contentsand may represent major sites of ion uptake into the symplasm,which was connected with the stele via plasmodesmatal connectionswith and between the cortical cells. The primary endodermalcells had lignified casparian strips and plasmodesmata in othercell wall areas. These connections were blocked by secondarysuberization of all except the endodermal passage cells oppositethe protoxylem arcs. Suberization of the hypodermis and endodermis and the appearanceof granular deposits in the vacuole occurred closer to the roottip of both genotypes after treatment with 100 mol m^–3NaCl. Levels of starch and triglyceride in 10 mm serial segmentswere similar between roots of control and salt-treated plantsand increased with distance back from the root tip. Chlorideconcentrations increased with salt treatment but values (ona tissue water basis) were similar between genotypes and betweenthe apical and proximal ends of the root. Reducing sugar concentrationsdecreased with salt treatment to a similar extent in roots ofboth genotypes. Key words: Citrus, Roots, Salinity     

The Growth of the Shoot Apex of Chenopodium rubrum L. Treated with a Florigenic Extract from Flowering Tobacco Plants: Preliminary Anatomical Observations

Anatomical changes in the shoot apex of Chenopodium rubrum L.treated with an extract from flowering tobacco plants and cultivatedin non-inductive conditions are described. They are comparedwith the anatomy of non-treated vegetative apices and with apicesof plants induced with a short day. Treatment with the extractresulted in both activation of cell division in the upper partof the apex and in apex elongation. Acceleration of leaf primordiainitiation and stimulation of branching took place. The effectcorresponds to the sequence of changes in photoperiodically-inducedplants but is more pronounced. Elongation following 10^–4M GA₃ treatment was of a differentnature; there was only a slight stimulation in the upper partof the apex in contrast with a strong stimulation of growthin length in the lower internodes. These preliminary resultssuggest a similarity between apical changes evoked by a stimulusproduced by short days and an exogenously applied floral stimulus.The changes differed from those caused by exogenous phytohormones. Key words: Chenopodium rubrum, florigen, shoot apex     

Morphological Alterations and Root Nodule Formation inAgrobacterium rhizogenes-mediated Transgenic Hairy Roots of Peanut (Arachis hypogaeaL.)

Transformed hairy roots were induced at the excised site ofthe epicotyl of dry mature seed of a Spanish type peanut(Arachishypogaea)cv. Java 13 2 weeks after inoculation with a wild typestrain ofAgrobacterium rhizogenes,MAFF-02-10266. Composite plantsconsisting of transformed roots with non-transformed shootswere cultured using pouches. Forty days after inoculation, thecomposite plant showed a root system with abundant root mass,more lateral branching and high fractal dimension compared tothe control. No differences were observed in production of rosette-typeroot hairs or the cross sectional structure between transformedand non-transformed roots. The inoculation ofBradyrhizobiumsp.A2R1 strain to the composite plants led to the induction oftransformed root nodules. These transformed root nodules showedproduction of leghaemoglobin in the bacterial zone and nitrogenaseactivity as assayed by C₂H₂reduction, and exhibited enlargementof the nodule cortex region andde novoroot formation from thenodule cortex.Copyright 1998 Annals of Botany Company Agrobacterium rhizogenes;Arachis hypogaeaL.; composite plant; peanut; transformation; root nodule.     

Length changes in herring (Clupea harengus) larvae: effects of capture and storage in formaldehyde and alcohol

The effects on standard length of storing laboratory-rearedlarval herring (9–19 mm live length) in 4% buffered formaldehydeor 70% buffered ethanol with and without simulated capture bytowed net were assessed. Lengths during storage were consideredstable after 15 days for larvae placed directly in formaldehyde,and after 30 days for larvae placed directly into alcohol orfor larvae treated by net-capture simulation before storage.The following linear regressions described the relationshipsbetween live length and the stored lengths of larvae after thesetimes: for larvae stored directly in formaldehyde, L = 1.765+ 0.867 x X₁ for larvae stored directly in alcohol, L = 0.564+ 0.971 x X₁; for larvae subjected to net-capture simulationthen stored in formaldehyde, L = 0.984 + 0.993 x X₁; and forlarvae subjected to net-capture simulation then stored in alcohol,L = 0.532 + 0.989 x X₁ where L = live standard length and X₁= standard length after storage. The non-linear regression formulaparameterized by Theilacker (Fish. Bull US, 78,685–692,1980) for northern anchovy larvae provided a good fit to thedata for herring larvae subjected to net-capture simulationand then stored in formaldehyde. However, the model had to bere-parameterized to provide a good fit for larvae stored inalcohol. The precision achieved in length measurements usinga computer-aided measuring system is also discussed.     

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     

The Relationship Between Lateral-Root Distribution and Endodermis and Pericycle Cell Length in Allium cepa L. Adventitious Roots

Endodermis and pericycle cell lengths were measured in intactand decapitated adventitious roots of Allium cepa L. Decapitationhad no effect on cell length in mature portions of the root,although it affected more immature cells, impeding normal elongation.Cell length shows a characteristic pattern in different zonesof the adventitious root: cells in the medial region were moremarkedly elongated. The number of lateral root primordia wasalso determined in different zones of the adventitious root.The possible relationship between lateral root distributionpattern and cell length in the endodermis and pericycle is discussed. Allium cepa, onion, endodermis, pericycle, lateral root, cell length     

SILICON IN THE ROOTS OF HIGHER PLANTS

Silicon (Si) distribution in the roots of Sorghastrum nutans (L.) Nash and Sorghum bicolor (L.) Moench. was investigated by means of the electron-probe microanalyzer and scanning electron microscope. In both species, Si was confined to the inner tangential wall of the tertiary-phase endodermal cells in the form of nodular silica aggregates of similar morphology and X-ray intensity. The results are compared to those for six closely related genera, as well as to studies of Si in the roots of species of other tribes of the family Poaceae. The various types of root deposits occurring in the family are described, and their relationships discussed. It is concluded that the type of Si distribution exhibited is determined largely by the phylogenetic status of the genus, rather than by the basic pattern of root anatomy.     

The role of apical development around the time of leaf initiation in determining leaf width at maturity in wheat seedlings (Triticum aestivum L.) with impeded roots

High soil resistance to root penetration (measured as penetrometerresistance, R_s slows down leaf growth and reduces mature leafsize in wheat seedlings {Triticum aestivum L.). Underlying changesin the kinetics of cell partitioning and expansion and in thesize and organization of mature cells were reported in companionpapers (Beemster and Masle, 1996; Beemster et al., 1996). Inthe present study, the relationships between apex growth, primordiuminitiation and expansion were analysed for plants grown at contrastingR_s, focusing on a leaf whose whole development proceeded afterthe onset of root impedance (leaf 5). High R_s reduced the rates of apex and leaf development, butdid not appear to have immediate effects on the pattern of developmentof the newly initiated phytomers. During an initial short period,the rate of development of a leaf primordium and associatednode were related to plastochronic age, according to similarrelationships (slopes) at the two R_s. Effects on developmentalpatterns were first detected on phytomer radial expansion duringplastochron 2. The ontogenetic pattern of leaf elongation wasaffected later, during the next few plastochrons preceding leafemergence (‘post-primordial stage’). It is concludedthat a reduction in the number of formative divisions and inthe number of proliferative cells along the intercalary mer-istemreported earlier (Beemster and Masle, 1996; Beemster et al.,1996) is not related to the size of the apical dome at leafinitiation nor to the size and number of meristematic cellsinitially recruited to the leaf primordium, which were all unaffectedby R_s. Rather they are generated at the primordial and post-primordialstages. Key words: Wheat, apex development, leaf primodium development, mature leaf width, root impedance