Determination of Stelar Elements in Roots of Pisum sativum L.

Cytochemical studies of esterase activity in 0.5 mm segmentsfrom root tips of Pisum sativum explanted for up to 9 days inbasal culture medium containing 2 per cent sucrose showed retentionof this activity. During this time, all segments from the secondand third 0.5 mm segments of the root tip developed xylem elementsas did the proximal end of the first segment. No xylem elementswere found in the 12–14 cells behind the quiescent centre.It is concluded that the central group of meristem cells aregenerally programmed to form tissues of the stele immediatelyon leaving the quiescent centre, and that the programming forxylem and phloem elements occurs as a second step. Pisum sativum L., garden pea, determination, histochemistry, esterases, stele, root     

Meristem Initial Cells in Irradiated Roots of Vicia faba: With one Figure in the Text

Beans (Vicia faba) were germinated for 24 hours and irradiatedwith X-rays. Primary roots were fixed after 9, 11, and 21 days.Aberrant chromosome complements, the result of chromosome changesinduced by irradiation, were present. They were used as cellmarkers to estimate the number of cell types present in themeristem initial cells of the growing root and the number ofprimordium initial cells from which root regeneration occurred.Up to 9 cell types occur as meristem initials. From the relativefrequencies of the different cell types, it is estimated thatthere are at least 32 actual meristem initial cells in regeneratingirradiated primary roots. This result is compatible with observationsmade on normal roots. The chimaerical nature of the regeneratingroot apparently does not interfere with the normal organizationof the meristem, but it serves to reveal what part of the organizationis.     

The First Appearance of Polyploid Nuclei in Primary Roots of Two Diploid Angiosperms

In Anchusa capensis, where the polyploid divisions are restrictedto a particular time and place in germination, cytophotometricanalysis showed that the polyploid cells arise at the time ofconversion of the globular-shaped embryo to the heart-shapedembryo. In Spinacia oleracea, where the divisions occur throughoutthe growth of the root, these polyploid cells do not arise duringembryogenesis but during germination and subsequent growth. Anchusa capensis Thunb., summer forget-me-not, Spinacia oleracea L., spinach, root meristem, polyploid divisions, determinate divisions, indeterminate divisions, germination, embryogenesis, cytophotometry     

Apical Meristem Activity and Lateral Root Anlage Development in Excised Roots of Pisum sativum L.

The initiation and subsequent emergence of lateral roots fromthe tissues of the primary has been examined in attached andexcised roots of Pisum. Lateral anlage inception took placein cultured roots which were 1 and 4 cm in length at the timeof excision. However, whereas a few primordia completed theirdevelopment and grew out as emerged laterals from those excisedroots which were 4 cm long, at the onset of culturc none appearedon those which had an initial length of 1 cm. The changes which took place in the rate of cell proliferationin the apical millimetre of each batch of cultured roots withtime following excision, were followed and related to the appearanceor otherwise of secondary roots on the cultured primaries. Pisum sativum L., garden pea, anlage, primodium, emerged lateral, apical meristem, cell proliferation     

Origin of Exogenous Adventitious Roots in Rhizomes of the Fern Microsorium scolopendria (Polypodiaceae)

Adventitious roots of rhizomes of the fern Microsorium scolopendriaarise exogenously from apical initials which have their originin the cell layer directly beneath the epidermis. Divisionsof the apical initial and its derivatives produce the adventitiousroot apical meristem. Cells between the apical cell and themeristele of the rhizome develop into the root trace. Anatomy, light microscopy, Microsorium scolopendria Burm.) Copel     

Proliferating and Quiescent Cells in the Apical Meristem of Elongating Lateral Roots of Vicia faba L.

Whole root systems of Vicla faba were continuously exposed to³H-TdR for periods of up to 72h, following which LI was determinedin the cap initials, in the quiescent centre or in that partof the meristem in which a quiescent centre will develop, andin the stele and in the cortex-epidermis at intervals alongthe apical 800 µm basal to the junction between the capinitials and the rest of the meristem, in newly-emerged (NE),0.2 and 4.0 cm long lateral roots, after each exposure period.Cell doubling time (Td), mean cycling time (Tc) and the sizeof the growth fraction (GF) were then calculated for each partof the meristem investigated in each batch of roots, from thecurves recording increase in labelling index (LI) with increasein duration of the period of exposure to ³H-TdR and from therate of increase in LI over the initial l-12h labelling period.Since it is extremely difficult to eliminate all sources oferror in calculating GF from the values obtained for LI in continuouslabelling experiments, it is emphasized that the values of GFreported in the present paper may not be totally accurate. Thisis also true of the results obtained for Tc as Tc was derivedfrom the product of the corresponding values for Td and GF. Cell doubling time and mean cycling time were both longer inthe cells forming the quiescent centre in the 0.2 and 40 cmlong roots than in any other part of the apical meristem examined.The size of the GF was found to decrease basally along the steleand the cortex-epidermis from the most apical to the most basalsegment examined in the NE, while Td increased in duration.Similar changes took place along the stele of the 0.2 cm longlaterals, but not in the cortex-epidermis of these roots or,to any great extent, in any of the tissues examined in the 4.0cm secondary roots. No consistent trend was apparent in theduration of Tc basally along either tissue examined in theseroots. It was concluded from these results, and from supportingdata in the literature, that, as the laterals elongated fromNE to 4.0 cm, the apical meristem increased in length.     

A Histological Study of the Development of Buds,Roots, and Embryos in Organ Cultures of Petunia inflata R. Fries

Organ cultures of Petunia inflata R. Fries have displayed adiversity of morphogenetic responses such as rhizogenesis, budformation, and embryo development. Studies oriented towardsdetermining the histological pathways preceding the morphogeneticexpression have indicated that bud formation and rhizogenesiswas always preceded by the establishment of a peripheral meristem.Embryos originated from cytoplasm-rich embryonal initials andthe pattern of their development simulated relatively closelywith the zygotic embryogenesis.     

Growth Substances in the Roots of Vicia faba II. THE EFFECTS OF AGEING AND EXCISION OF THE MAIN TAP-ROOT MERISTEM

A quantitative chromatographic study has been made of the changesin the activity and distribution of the main ether-soluble acidauxins of Vicia faba seedling root systems during developmentand resulting from excision of the main tap-root meristem. AnIAA-like auxin (AP (ii)) is apparently synthesized predominantlyin apical and at a lower rate in lateral meristems. Productionseems to stop when meristematic growth stops. Its concentrationin mature extended cells is much lower and may fall to zeroin old cells, suggesting active degradation by an auxin-oxidase.Excision of the main tap-root tip gradually results in a greatlyaugmented production of AP (ii) in lateral meristems, conceivablythe result of correlative growth promotion. A second auxin and root-growth inhibitor (AP (iii)) is presentat higher activity levels than AP (ii) in tap-root meristemsand at the same level in lateral meristems. In mature cellsits activity is much lower than that of AP (ii). In contrastto AP (ii) it accumulates in both tap-root meristems and maturetissue as the root system ages. It could also be produced duringmeristematic growth but is not subsequently degraded. As withAP (ii), excision of the tap-root tip brings about a great increasein its concentration in lateral tips. A third auxin and root-growth accelerator (AP (i)) (accelerator?) is present in lower concentrations in both meristem and maturetissue. Its concentration tends to decrease with ageing and,in lateral meristems, is not affected by tap-root tip excision. It is suggested that AP (ii) produced by the meristem is normallyat suboptimal levels in the extending cells and may be the principalhormone controlling extension growth. AP (iii) accumulationmay account for growth deceleration on ageing. The role of AP(i) remains obscure. It is unlikely that correlative effectsof the taproot tip on lateral root growth are exercised directlyvia these auxins.     

Some Cytological Changes accompanying the Regeneration of Meristematic Activity at the Apex of Decapitated Roots of Vicia faba L.

The changes that took place in mitotic index (MI), labellingindex (LI) and the relative proportions of interphase nucleiwith different amounts of DNA have been investigated duringthe regeneration of meristematic activity at the apex of rootsof Vicia faba over the 144 h period following removal of thecap and apical mm of the meristem. Measurements were also madeof the corresponding changes that took place as cells were displacedbasally along the root from the apex over the experimental period.In both parts of the root, MI and the relative proportions ofnuclei with different DNA contents changed from levels similarto those at the apex of the controls at the start and end ofthe experiment to levels resembling those found in more matureparts of the root at 24 and 48 h. In contrast to these results,LI declined over the experimental period. These cytologicalchanges were aresult of the development of lateral root primordiain both the apical 2 mm of the decapitated roots and as cellswere displaced out of the meristem into more basal parts ofthe root. It was concluded that the events leading to the regenerationof meristematic activity at the apex of roots from which thecap and apical mm of the meristem were removed, are no differentfrom those which result in lateral formation as cells are displacedbasally along the primary root from the apex, and they takeplace over the same time interval in both systems.     

Developmental Responses to Drought and Abscisic Acid in Sunflower Roots: 2. MITOTIC ACTIVITY

Mitotic activity was studied in the root apices of aeroponicallygrown sunflower seedlings (Helianthus annum L. var. RussianGiant) which were draughted or treated with abscisic acid (ABA)over a 7 d period. Labelling index (LI) and mitotic index (MI)were scored from autoradiographs of median longitudinal sectionsof [³H] methyl-thymidine treated root apices. Both drought stressand ABA-treatment (at a concentration of 10^–2 mol m^–3inhibited DNA synthesis and mitosis within the first 6 h oftreatment. The depression of mitotic activity was first evidentin the proximal regions of the meristem (1000–1500 µmfrom the cap junction). This was followed by a general depressionof mitotic activity throughout the meristem which was, in turn,followed by a partial recovery of mitotic activity in the distalregions of the meristem. The beginning of this partial recoverywas concurrent with the activation of the quiescent centre (QC).Treatment with lower concentrations of ABA (10^–3 mol m^–3and 10^–4 mol m^–3) also inhibited mitotic activity.Exogenous supplements of sucrose to the plant did not alleviatethe inhibition of mitotic activity by drought or ABA. Thesefindings support the hypothesis that ABA mediates drought-inducedchanges in the primary development of sunflower roots. Key words: Abscisic acid, drought, mitotic activity     

Cellular Growth in Roots of a Gibberellin-Deficient Mutant of Tomato (Lycopersicon esculentum Mill.) and its Wild-type

The role of gibberellins in regulating the growth of tomatoroots was investigated by comparing various cellular parametersin cultured roots of the gibberellin-deficient mutant gib-l/gib-lwith those in roots of the near-isogenic wild-type. In addition,wild-type roots treated with 0?1 µM 2S,3S paclobutrazol,an inhibitor of gibberellin biosynthesis, and mutant roots treatedwith 0?1 µM GA₃ were also compared: the former roots constitutea phenocopy of the mutant, whereas the latter roots appear tobe ‘normalized’ and similar to wild-type. The elongationof mutant and phenocopied roots were similar, their maximumelongation rates being about half or two-thirds that of wild-typeor GA₃-treated mutant roots, respectively. These rates wereinterpreted in terms of the numbers and lengths of cells withinthe meristematic and non-meristematic portions of the elongationzone. Mean meristem length tended to be shorter in both themutant and the 2S,3S paclobutrazol-treated wild-type roots thanin the other two types of root. A major difference between thetwo pairs of mutant and normal roots was their mean final celllengths: mean lengths of cortical cells of the mutant and 2S,3Spaclobutrazol-treated roots were, respectively, 39% and 25%shorter than the mean length of wild-type roots. Final celllength in the GA³-treated mutant roots were similar to wild-type.By contrast, the diameters of mature cortical cells of the mutantand phenocopy were about 20% greater than the diameters of equivalentwild-type or ‘normalized’ mutant cells. The meanvolumes of cortical cells in all four types of roots showedno significant differences. Knowledge of the distribution ofcortical cell lengths, widths and volumes along the root axis,together with information about the rate of root elongation,permitted comparisons of the relative elemental growth ratesof each of these three cellular parameters. The available evidence suggests that the level of endogenousgibberellins in mutant roots is lower than in wild-type roots.The present results, therefore, suggest that endogenous gibberellinsare necessary for normal growth of cultured tomato roots andthat they regulate the relative amounts of growth at the longitudinaland transverse walls of the cells which, in turn, affects theshape of the elongating cells. Key words: Cell growth, cultured roots, gibberellin, gib-l mutant, Lycopersicon esculentum, 2S,3S paclobutrazol, relative elemental growth rate, root meristem     

Growth Pattern and Movement of Epidermal Cells within Leaves of Asphodelus tenuifolius Cav.

The pattern of growth (velocity field) in the intercalary growthzones of monocotyledon leaves can be determined from patternsof cell number density (number per unit length of cell file)and leaf elongation rates using theory based on a cell numberconservation equation. The case where elongation rate is non-steadywhile the pattern of cell number density is steady is discussedand a method for extending calculations into the meristem usingobservations of numbers of mitotic cells is outlined. Applicationof these methods is illustrated using data for epidermal cellsin the first leaf of Asphodelus tenuifolius Cav. During earlyleaf development, leaf elongation rate increased exponentiallybut cell number density and mitotic number density were steady.Cells 0.1 mm from the base of the leaf when leaves were 3.2mm long took 8.3 d to move through the growth zone. In leavesthat were 4 d older, similar cells took 5.1 d to traverse thegrowth zone. Increases in the rates of leaf elongation and ofcell movement appeared to be associated mainly with increasesin total rates of cell production in the epidermal meristem. Asphodelus tenuifolius Cav., Asphodelus fistulosus L., velocity field, meristem, mitotic cell number density, extension-only zone     

Some Observations on Invertase Activity in Roots of Vicia faba L.

Invertase activity has been determined at intervals along primaryroots of Vicia faba as they elongated from 0·5 to 8 cm.Little activity was evident in 0·5–1·0 cmlong primaries but in those 2–8 cm in length the mainpeak of enzyme activity was associated with the region of cellelongation. Changes took place in the pattern of invertase activityalong the primary roots as they lengthened and these changeshave been correlated with fluctuations in both the rate of rootelongation and the supply of sucrose to the root from the cotyledons.The presence of a root cap did not increase the activity ofthis enzyme in the apical 1 mm of these roots. Invertase activity was higher in lateral root primordia thanin most parts of the primary root basal to the meristem, presumablybecause of the presence of sucrose in the adjacent cavity inthe cortex of the primary root. The peaks of invertase activityfound basal to the region of cell elongation in 3–8 cmlong primary roots probably resulted from the development ofroot pnmordia in these parts of the root.     

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     

Aluminium Toxicity in Roots: An Investigation of Spatial Sensitivity and the Role of the Root Cap

The primary symptom of aluminium (Al) toxicity in higher plantsis inhibition of root growth. In this study, we investigatedthe spatial sensitivity of maize (Zea mays L.) roots to Al.A divided-chamber technique indicated that only exposure ofthe terminal 10 to 15 mm of the root to Al resulted in inhibitionof growth. Application of Al to all but this apical region ofthe root had little or no effect on growth for 24 h and causedminimal damage to the root tissue. Small agar blocks infusedwith Al were then applied to discrete areas of the apex of maizeroots to determine which section (root cap, meristem or elongationzone) was more important to Al-induced inhibition of growth.The terminal 20 to 30 mm of root (root cap and meristem) mustbe exposed to Al for inhibition. Application of Al to the 30mm of root proximal to this terminal zone (elongation zone)resulted in damage to the root tissue but no significant inhibitionof growth. Therefore, the visible injuries incurred by rootsduring Al-stress are not associated directly with the inhibitionof root growth. Furthermore, removal of the root cap had noeffect on the Al-induced inhibition of root growth in solutionexperiments and argues against the root cap providing protectionfrom Al stress or serving an essential role in the mechanismof toxicity. We suggest that the meristem is the primary siteof Al-toxicity. Key words: Aluminium, toxicity, root growth, root cap     

Experimental and Analytical Studies of Pteridophytes: XXXIX. Morphogenetic Investigations of Sori in Leptosporangiate Ferns

The fact that there is a demonstrable unity of origin of themeristematic tissues in ferns, and that developments of an adventitiouscharacter are rare or absent, has led to the view that the youngsoral receptacle, or placenta, is a special kind of meristem,now described as a sporogenous meristem. These meristems originateas growth centres in the leaf marginal meristem, the latterbeing now recognized as an organized formative region sharingmany of the properties and functions of the shoot apical meristem.According to the nature of the growth relationships in the marginalmeristem after the onset of the reproductive phase, sporogenousmeristems may remain in a marginal or intramarginal position,or they may come to occupy a superficial position on the abaxialside of the leaf. In those species in which the further growthof the leaf margin is more or less completely inhibited as sorusdevelopment proceeds. the mature sori occupy marginal or intramarginalpositions: on the other hand, in species in which marginal growthis sustained, the sori eventually occupy superficial positionson the abaxial leaf surface. Thus the phylogenetic shift ofsori from a marginal to a superficial position, as envisagedby evolutionary morphologists of an earlier period, has nowbeen shown to take place in the inidividual species ofSuperficialsas an onto-genetic development. Information of this kind isconsidered to be of special value and interest in that it admitsof a convergence of the ideas that have emerged from both morphogeneticand phylogenetic studies.     

Ontogeny of Tendrils in Antigonon leptopus H. & Arn.

In Antigonon leptopus the main tendrillar and axillary branchis a modified inflorescence axis. It usually bears 6–7lateral bracts out of which 3–4 lower ones are small andleaf-like while the upper 2–3 are tendrillar; 2–3tendrils are also present at its terminal end. The vegetativeshoot apex shows a single layer of tunica and an inner massof corpus without any cytohistological zonation. The earliestaxillary bud or tendril meristem arises at the second node andit elongates due to rib meristem activity. The bract primordiaarise in an acropetal succession. The initiation of the bract-tendriland the leafy bract is similar. In the development of the bract-tendril,marginal meristem activity is absent or reduced and the differentiationof the apical and subapical initials is absent. The terminalbract-tendrils arise as lateral appendages and the residuumof the apical meristem of the main axillary tendril persistsfor some time. In the flowering period the floral buds arisein the axils of the bracts and bract-tendrils. No flowers arepresent in the axils of terminal bract-tendrils.     

Estimation of Growth Fractions in Meristems of Zea mays L.

Three ways of estimating the fraction of cycling cells in ameristematic population have been examined region by regionthroughout the root meristem of Zea mays. Only in the quiescentcentre and in the nearby proliferating regions did the threemethods produce similar results. The estimate based on the ratioof the average cycle duration of cycling cells and the cell-doublingtime is the most reliable. The two methods that use labellingindex either immediately after a short pulse of ³H-thymidineor after a long labelling period exaggerate the growth fractionby counting endoreduplicating cells which become prominent evenin the proximal half of the meristem and in the second and subsequenttiers of the cap meristem. The nature of the non-cycling cellsis discussed.