An Automata-theoretical Model of Meristem Development as Applied to the Primary Root of Zea mays L.

Observations were made of the sequence of division within thecellular packets (groups of cells of common descent) which comprisethe cell files that run the length of the central cortex ofthe primary root meristem ofZea mays. These sequences, and alsothe relative lengths of the cells within the packets recordedat various times during root growth, indicate that cell-filedevelopment can be expressed using one, or a limited number,of deterministic ‘bootstrap’ L-systems which assigndifferent lifespans to sister cells of successive cell generations.The outcome is a regular pattern of divisions from which daughtercells emerge usually with unequal, but definite, lengths. Inthe immediately post-germination stage of root growth, one divisionpathway is especially common in the cortex and generates sequencesof unequal daughters having a particular basi-apical orientation.Later in root growth, the cellular pattern in the cortex indicatesthat this pathway is replaced by another where unequal divisionsare not so marked, but which nevertheless continues to maintaina regular arrangement of differently sized cells. This latterpathway is characteristic of a zone close to the initial cellsof the cortex. It is present at all stages of root growth andspreads along the length of the cortex as the descendants ofthese initials proliferate. The development of the whole corticalcell file can be simulated from knowledge of the growth functionsof the bootstrap systems. The files so generated contain allthe observed cell patterns. The growth functions also predictthe sequence in which cells cease dividing near the proximalmargin of the meristem, but for this it is necessary to incorporatea counter for the number of divisions that will be accomplishedin the cell file. Cytological requirements for the propagationof unequal divisions, together with a consideration of the natureof the division counter, as well as the significance of theswitch in division pathways encountered during early root growth,are discussed in the context of this deterministic model ofcell division. Cell division; root meristem; L-systems; Zea mays     

Cell genealogies in a plant meristem deduced with the aid of a 'bootstrap' L-system

Abstract. The primary root meristem of maize ( Zea mays L.) contains longitudinal files of cells arranged in groups of familial descent (sisters, cousins, etc.). These groups, or packets, show ordered sequences of cell division, which are transverse with respect to the apico-basal axis of the root. The sequences have been analysed in three zones of the meristem during the course of the first four cell generations following germination. In this period, the number of cells in the packets increases from one to 16. Theoretically, there are 48 possible division pathways that lead to the eight-cell stage, and nearly 2- × 10⁶ that lead to the 16-cell stage. However, analysis shows that only a few of all the possible pathways are used in any particular zone of the root. This restriction of pathways results from inherited sequences of asymmetric cell divisions, which lead to sister cells of unequal length. All possible division pathways can be generated by deterministic 'bootstrap' L-systems which assign different life spans to sister cells of successive generations and hence specify their subsequent sequence of divisions. These systems simulate propagating patterns of cell divisions, which agree with those actually found within the growing packets that comprise the root meristem. The patterns of division are specific to cells originating in various regions of the meristem of the germinating root. The importance of such systems is that they simulate patterns of cellular proliferation where there is ancestral dependency. They can therefore be applied in other growing and proliferating systems where this is suspected.     

Experimental Modification of Cell Division Patterns in the Root Meristem of Zea mays

In the meristem of the young primary root of maize seedlingsthe first transverse division in the cortex 250 µm fromthe root apex results in two daughter cells of distinctly unequalsize. This division could be rendered equal by raising the seedlingsin up to 7.5% methanol. The pattern of the subsequent two orthree transverse divisions in the cortex, as revealed by thearrangement of the newly divided cells in the resultant cellularpackets, was acropetal in the methanol-treated roots but basipetalin the control roots. The sequence of division within a cellularpacket tended to follow the distribution of cell sizes - largercells divided earlier than smaller cells. A temporary arrestof cell division by exposing roots to cold (5 °C) conditionshad no effect on the sequence of divisions that followed whenthe roots were allowed to recover at 20 °C. The resultssuggest that the normally asymmetric position of the cell wallformed at cytokinesis is subject to active regulation and thatmethanol interferes with this process. The cytoplasm of certaincells in the root meristem was also found to be unequally distributed,as judged by Azure B staining, between the two ends of the cell.Cytoplasmic asymmetry was not directly correlated with inequalityof division, although it too was affected by methanol. Cell polarity, root meristem, unequal division, Zea mays     

The Response of the Primary Root Meristem of Zea mays L. to Various Periods of Cold

Primary roots of maize (Zea mays L.) grown in nutrient solutionat 5?C elongate at about 1% of the rate found at 20?C. The apicalmeristem becomes shorter and shows little proliferative activityat 5?C, but following transfer to 20?C mitoses increase in frequencyand the meristem regrows to its original length. Both the amountby which the meristem shortens and the time for its completeregrowth are related to the period spent at 5?C. The shorteningof the meristem suggests that at the lower temperature the balancewhich normally exists between cell production and differentiationis altered, the latter continuing at a relatively faster ratethan the former. A new, steady-state balance between the twoprocesses is re-established during the recovery period. Themeristem recovers as a result not only of its own mitotic activitybut also through stimulation of cell division in the quiescentcentre. The degree to which the quiescent centre is activated,as judged by its mitotic index and the number of nuclei labelledby feeding with tritiated thymidine, increases as the durationof the preceding cold treatment increases. The close relationshipbetween proliferative activity in the quiescent centre and theminimum length of the meristem following the cold treatmentsuggests that there is communication between these two zoneswhich co-ordinates their respective rates of cell productionand helps to maintain a normal meristem structure. The resultsemphasize the importance of the quiescent centre as a reservoirof cells that can re-establish a meristem rendered non-functionalthrough the impact of unfavourable environmental conditions. Key words: Meristem, quiescent centre, root, temperature, Zea mays     

Cell Displacement Through the Columella of the Root Cap of Zea mays L

Exposing roots of Zea mays to a solution of caffeine for 1 hinduces a small population of binucleate cells in the meristem.The progress of the binucleate cell population was then followed,in time, as it was displaced along the length of the cap columella.Since this method of marking cells seems to have no effect onthe subsequent pattern of cell proliferation in the cap meristem,the movement of the binucleate cells through the cap is inferredto be similar to the movement of cells in an undisturbed cap.The binculeate cells that persist in the cap are believed tobe cells that were engaged in their final mitosis at the timeof the caffeine treatment, so the time that it takes for themto appear at the edge of the cap is a measure of the periodfor which a cell is contained in the non–dividing portionof the tissue before being lost from the cap surface. In rootsof Zea grown at 22 °C cells take about 7 days to reach thetip of the cap columella and about 2 to 3 days to reach theflanks of the cap following their displacement from the capmeristem. Zea mays, root cap, cell displacement, binucleate cells     

Effect of Root and Shoot Meristem Temperature on Shoot to Root Dry Matter Partitioning and the Internal Concentrations of Nitrogen and Carbohydrates in Maize and Wheat

Maize (Zea mays L.) and spring wheat (Triticum aestivum L.)were grown in nutrient solution at uniformly high air temperature(20 °C), but different root zone temperatures (RZT 20, 16,12 °C). To manipulate the ratio of shoot activity to rootactivity, the plants were grown with their shoot base includingthe apical meristem either above (i.e. at 20 °C) or withinthe nutrient solution (i.e. at 20, 16 or 12 °C). In wheat, the ratio of shoot:root dry matter partitioning decreasedat low RZT, whereas the opposite was true for maize. In bothspecies, dry matter partitioning to the shoot was one-sidedlyincreased when the shoot base temperature, and thus shoot activity,were increased at low RZT. The concentrations of non-structuralcarbohydrates (NSC) in the shoots and roots were higher at lowin comparison to high RZT in both species, irrespective of theshoot base temperature. The concentrations of nitrogen (N) inthe shoot and root fresh matter also increased at low RZT withthe exception of maize grown at 12 °C RZT and 20 °Cshoot base temperature. The ratio of NSC:N was increased inboth species at low RZT. However this ratio was negatively correlatedwith the ratio of shoot:root dry matter partitioning in wheat,but positively correlated in maize. It is suggested that dry matter partitioning between shoot androots at low RZT is not causally related to the internal nitrogenor carbohydrate status of the plants. Furthermore, balancedactivity between shoot and roots is maintained by adaptationsin specific shoot and root activity, rather than by an alteredratio of biomass allocation between shoot and roots.Copyright1994, 1999 Academic Press Wheat, Triticum aestivum, maize, Zea mays, root temperature, shoot meristem temperature, biomass allocation, shoot:root ratio, carbohydrate status, nitrogen status, functional equilibrium     

A Maize Mutant with an Altered Vascular Pattern

A recessive maize mutant with disrupted seedling developmentwas isolated following transpositional mutagenesis with Mutator.This mutant, initially identified during germination on thebasis of abnormal growth of the scutellar node, was designatedlsn1 (l arge s cutellar n ode). The mutant seedling exhibitsan enlarged primary root with a longitudinal groove and multipleseparate root tips. The mutant root is shorter than normal,because of defective cell elongation, and lacks lateral roots.The mutant plant shows defective leaves and reduced internodeelongation. Histological analyses on primary root, shoot, scutellarnode and juvenile leaves revealed a series of defects, all relatedto an irregular differentiation of vascular elements. In addition,in situ hybridization of mutant leaves demonstrates an abnormalpattern of expression of Knotted-1, a marker of meristem function.The presence of multiple roots fused together can be interpretedas suppression of the negative control responsible for the differentiationof only one root primordium. Therefore, the data obtained onseedlings of lsn1 point to a relationship between meristem activity,vascular differentiation and auxin polar transport, and mayallow the identification of a gene which is active during embryogenesis.Copyright2000 Annals of Botany Company Auxin, defective seedling, maize, meristem activity, vascular differentiation, Zea mays.     

Development of Lateral Root Primordia in Vicia faba, Pisum sativum, Zea mays andPhaseolus vulgaris: Rates of Primordium Formation and Cell Doubling Times

Lateral root primordium development has been examined in primaryroots of Vicia faba L., Pisum sativum L., Zea mays L. and Phaseolusvulgaris L. Following their initiation from an estimated minimumnumber of 77–162, 20–57, 17 and 12 cells respectivelyin Vicia, Phaseolus, Pisum and Zea, the primordia rapidly increasedin cell number to emerge as secondary roots about 2.8–3.6days later depending on the species being examined. Cell doublingtimes were estimated directly from cell numbers at differenttimes following primordium inception and were found to increasewith increase in primordium size in each of the species investigated. The number of primordia formed per cm of root growth per daywas greatest in Zea and least in Pisum. A comparison of thedata obtained for Vicia with that in the literature led to theconclusion that although the number of primordia produced percm of root growth was independent of the rate of primary elongation,the number produced per day increased in a linear fashion withincrease in the rate at which the primary lengthened. Vicia faba L, Pisum sativum L, Zea mays L, Phaseolus vulgaris L, broad bean, garden pea, maize, dwarf bean, root primordia, cell division, cell doubling time     

Image Analysis of Maize Root Caps--Estimating Cell Numbers from 2-D Longitudinal Sections

The cap of the primary root of maize produces several thousandborder cells that are shed from the outside of the cap eachday. Border cell production is important in the penetrationof soil by roots, and in influencing the activity of both beneficialand pathogenic organisms in the rhizosphere. To improve understandingof the dynamics of border cell production, it is desirable toknow the number of cells in different parts of the root cap.An image analysis procedure was used to quantify cell dimensionsand locations in the median longitudinal section of maize (Zeamays L.) root caps. Calculations based on root symmetry werethen used to estimate the number of cells in 3-dimensions. Ourestimation procedure was tested initially using regular arraysof identical square and hexagonal shapes to represent cells.It was then tested using two different tissues showing analogousarrays: a transverse section through the maize root cap junction,and a transverse section through a barley root. Good linearcorrelations were obtained between the number of cells estimatedand the number of cells actually counted in the microscope.The numbers of cells in the whole maize root cap (8870 ±390) were then estimated from longitudinal sections. These numbersof cap cells agreed with values that had been estimated formaize by other methods. In the first tier of the cap meristem,ten-times more meristematic cells were located in the cap flanks(>500 cells) than were present in the columella portion.Similarly, only 7% of cells in the outermost layer of the rootwere associated with the columella region of the cap, a fractionwhich compared well with previous measurements of sloughed cellsextracted from rhizosphere sand. This present technique canbe applied to estimate the numbers of cells in any cylindricallysymmetrical tissue from two-dimensional sections. Copyright2001 Annals of Botany Company Anatomy, border cells, cell production, image analysis, maize, rhizosphere, root cap, sloughing, stereology, Zea mays L     

The Difference Between Open and Closed Meristems

An open and a closed root meristem have been compared by investigatingthe cell kinetics of small regions of the apices of Helianthusand Zea. The cells of the stelar pole are quiescent in both and thereis no exchange of cells between stele and cortex or stele andcap. The immediately distal cells in the closed meristem (Zea)are also quiescent and the few divisions that do occur can betransverse or longitudinal. In the open meristem (Helianthus)these cells are not quiescent, but they go out of cycle transiently,prolonging the potential cell-doubling time. Their divisionsare transverse. It is a consequence of these differences thatclosed meristems form root caps discrete from the cortex whereasopen meristems force instability in the boundary between theperipheral part of the cap and the cortex. Another consequencein roots with open meristems is a succession of columella complexestransversely displaced from each other by the state of fluxin the meristem during the non-cycling phase of the proximaltier of cells, those immediately distal to the stelar pole. The results are discussed in relation to the ontogenetic onsetof quiescence and the evidence for switches between open andclosed operation of meristems. meristem, root apex, Helianthus annuus, Zea mays L.     

Nuclear and Cell Sizes in Different Regions of Root Meristems of Zea mays L.

Nuclear volumes and cell areas were determined for seven regionsof the meristem of roots of Zea mays. Roots were fixed in 10per cent neutral buffered formalin, in 3 per cent glutaraldehydeor in acetic acid/alcohol; they were prepared as sections oralls were teased apart. Mean volumes of interphase nuclei weresimilar in all regions of the root except the vascular tissueof the stele. Mean nuclear volumes and the overall range ofvolumes were similar in sub-populations of cells with differentproportions of G¹, S and G² cells, e.g. in row I of root capinitials, whose cells lack a G¹ phase, and in quiescent centrecells, which are mainly in G¹. Nuclear volume does not appearto be closely correlated with DNA content. Nuclear volumes covereda 6 to 12-fold range within a meristem and even within specificregions, in which cells are part of the same cell lineages,there was a 4- to 9-fold range. Nuclear volumes were comparedin sister cells in rows I and II of the root cap initials. In10 per cent of the pairs, sister nuclei had identical volumes;the other pain had different volumes and mean difference was68 µm³. Mechanisms by which this variability could begenerated are discussed, particularly asymmetry, at mitoses,of factors that regulate nuclear growth. Zea mays L., nuclear volume, cell size, root mcristem, DNA content, mitosis     

Tissue zinc distribution in maize seedling roots and its action on growth

Zinc (Zn) distribution over tissues and organs of maize (Zea mays L.) seedlings and its action on root growth, cell division, and cell elongation were studied. Two-day-old seedlings were incubated in the 0.25-strength Hoagland solution containing 2 or 475 μM Zn(NO₃)₂. Zn toxicity was assessed after the inhibition of primary root increment during the first and second days of incubation. The content of Zn was determined by atomic absorption spectrometry in the apical (the first centimeter from the root tip) and basal (the third centimeter from the kernel) root parts. Zn distribution in various tissues was studied by histochemical methods, using a metallochromic indicator zincon and fluorescent indicator Zinpyr-1 and light and confocal scanning fluorescent light microscopy, respectively. To evaluate Zn effects on growth processes, the average length of the meristem; the length of fully elongated cells; the number of meristematic cells in the cortex row; and duration of the cell cycle were measured. When the Zn concentration in the solution was high, the Zn content per weight unit was higher in the basal root part due to its accumulation in lateral root primordial. Zn was also accumulated in both the meristem apoplast and cell protoplasts. In the basal and middle root parts, Zn was detected essentially in all tissues predominantly in the apoplast. Zn inhibited both cell division and elongation. Under Zn influence, the size of the meristem and the number of meristematic cells decreased, which was determined by an increase in the cell cycle duration. The length of the fully elongated cells was also reduced. A comparison of Zn distribution and growth-suppressing activity with other heavy metals studied earlier allows a conclusion that toxic action of heavy metals is mainly determined by physical and chemical properties of their ions and specific patterns of their transport and distribution. As a result, two basic processes determining root growth, e.g., cell division and elongation, could be affected differently.     

Characterizing Pathways by Which Gravitropic Effectors Could Move from the Root Cap to the Root of Primary Roots of Zea mays

Plasmodesmata linking the root cap and root in primary rootsZea mays are restricted to approx. 400 protodermal cells borderingapprox. 110000 µm² of the calyptrogen of the root cap.This area is less than 10% of the cross-sectional area of theroot-tip at the cap junction. Therefore, gravitropic effectorsmoving from the root cap to the root can move symplasticallyonly through a relatively small area in the centre of the root.Decapped roots are non-responsive to gravity. However, decappedroots whose caps are replaced immediately after decapping arestrongly graviresponsive. Thus, gravicurvature occurs only whenthe root cap contacts the root, and symplastic continuity betweenthe cap and root is not required for gravicurvature. Completelyremoving mucilage from the root tip renders the root non-responsiveto gravity. Taken together, these data suggest that gravitropiceffectors move apoplastically through mucilage from the capto the root. Calyptrogen, open meristem, protoderm, root cap, root gravitropism, Zea mays     

Regeneration of the Root Cap of Zea mays L. and Pisum sativum L.: A Study with the Scanning Electron Microscope

Removal of the root cap from a root apex initiates regenerationof a new cap. The process has been followed using scanning electronmicroscopy. Quantitative data have been obtained for the growthin area of the exposed acroscopic surface of the quiescent centre(QC) and the increase in volume of the regenerating cap tissue.In Zea the surface of the QC shows an initial rapid increasein area followed by a slower increase. In Pisum the surfacearea increases uniformly, a rapid initial phase being absent.Together with observations on the behaviour of an incision atthe exposed surface, the results indicate that in Zea the capnormally imposes a constraint upon radial growth at the acroscopicsurface of the QC; in Pisum the QC appears not to be so constrained.The different responses may be related to the different arrangementsof cells at the apex of the meristem of these two species. Zea mays, Pisum sativum, maize, peao, scanning electron microscopy, root apex, regeneration     

Root Gravitropism in a Cultivar of Zea mays Whose Columella Cells Contain Starch-deficient Amyloplasts

Starch occupies 4.2 per cent of the volume of plastids in calyptrogencells in primary roots of Zea mays L. cv. vp-7 wild type. Plastidsin calyptrogen cells are distributed randomly around large,centrally located nuclei. The differentiation of calyptrogencells into columella cells is characterized by cellular enlargementand the sedimentation of plastids to the bottom of the cells.Although sedimented plastids in columella cells do not containsignificantly more starch than those in calyptrogen cells, primaryroots are graviresponsive. The onset of root gravicurvatureis not associated with a significant change in the distributionof plastids in columella cells. These results indicate thatin this cultivar of Z. mays (1) the sedimentation of plastidsin columella cells is not based upon their increased densityresulting from increased starch content alone, (2) starch-ladenamyloplasts need not be present in columella cells for rootsto be graviresponsive, and (3) the onset of root gravicurvaturedoes not require a major redistribution of plastids in columellacells. Columella cell, gravitropism (root), plastids, root cap, Zea mays     

Flow Cytometric Determination of Relative Nuclear DNA Contents in Bicellulate and Tricellulate Pollen

Nuclear DNA content in mature pollen was measured with a flowcytometer Pollen of Lilium longiflorum, Dendranthema grandiflora(syn Chrysanthemum monfolium) and Zea mays was chopped and stainedwith the DNA fluorochrome DAPI DNA levels, expressed as arbitraryC values, were compared with those of nuclei isolated from leafor root material of the same plants In mature tricellulate pollen the generative cell is dividedafter second pollen mitosis into two sperm cells Tricellulatepollen from maize and chrysanthemum gave rise to one large 1Cpeak and, only in the case of chrysanthemum, a much smallerone at the 2C level These results suggest that the haploid nucleiof the vegetative as well as both sperm cells in tricellulatepollen are arrested in the G₁ stage of nuclear division Thesmall 2C peak in the case of chrysanthemum probably arose froma fraction of pollen with the sporophytic chromosome number(2n pollen) In contrast to this, mature bicellulate lily pollengave rise to two identical peaks at the 1C and the 2C levelFrom this result it was concluded that in bicellulate pollen,the 1C peak is caused by the signal of the haploid vegetativenucleus arrested in the G₁ stage of nuclear division, whereasthe 2C peak originates from the haploid generative nucleus whichhas already undergone DNA synthesis and is arrested in G₂ Lilium longiflorumThunb, lily, Dendranthema grandiflora Tzelev (syn Chrysanthemum morifolium Ramat ), chrysanthemum, Zea maysL, maize, male gametophytic cells, vegetative cells, generative cells, sperm cells, unreduced pollen, sporophytic cells, relative nuclear DNA contents, replication stage     

Nuclear Chromatin Structure in Relation to Cell Differentiation and Cell Activation in the Cap and Quiescent Centre of Zea mays L.

Barlow, P. W. 1985. Nuclear chromatin structure in relationto cell differentiation and cell activation in the cap and quiescentcentre of Zea mays L —J. exp. Bot. 36: 1492–1503.Nuclear chromatin structure has been analysed by electron microscopyof thin sections of cells in four zones of the root cap—meristem,central, slime-secreting and outermost cells—and alsoin the quiescent centre of the root before and after decapping.The chromatin pattern has been related to the DNA and RNA syntheticactivity of the nuclei. During cap cell maturation there wasa progressive condensation of the chromatin and this was accompaniedby some reduction of RNA synthesis. The degree of condensationwas estimated from the area and number of pieces of electrondense chromatin which increased and decreased, respectively,during cap maturation. The volume fraction of condensed chromatinwas also estimated but, in the cap, was not found to be a goodindicator of nuclear activity. The outermost cells of the capshowed the greatest degree of chromatin condensation but werestill active in RNA synthesis. Microdensitometry of their nuclearDNA contents gave an indication of loss of DNA in some of thenuclei. Decapping activated DNA and RNA synthesis in the quiescentcentre and also stimulated a decondensation of chromatin: thenumber of condensed pieces of chromatin increased, and theirsize and volume fraction both decreased 4 h after decapping.The number of pores per unit length of nuclear envelope profilewas also estimated. In the cap this number increased duringcap maturation; in the activated quiescent centre the numberremained constant except for a small rise 4 h after decapping Key words: Zea mays, chromatin, root cap, quiescent centre     

Germination and Early Growth of Plants Studied Using Neutron Radiography

Seed swelling, germination, root extension, lateral root initiationand shoot growth were studied in soils of different water contents,using non-destructive, serial neutron radiography. Seeds fromthree varieties of soya beans (Glycine max L.) and one varietyeach of maize (Zea mays L.) and vetch (Vicia sativa L.) wereused. The seeds germinated when they had increased in size bya certain amount, if germination is taken as the time when theradicle first appears. The rate at which roots and shoots extendalso depend on soil water content. Glycine max L., Vicia sativa L., Zea mays L., Soya bean vetch, maize, seed germination, root extension, lateral root initiation, neutron radiography