Relationship Between Lateral Root Primordia in Different Ranks

Comparisons with theoretical random distributions indicate thatthe longitudinal positional relationships between lateral rootsin different ranks are random in roots of Musa acuminata, Pistiastratiotes and i, species in which lateral root primordia arisein the root tip. In Potentilla palustris, where lateral rootprimordia arise further back in the root, there is a deficiencyof close spacings, which indicates that there is some interactionbetween ranks. Musa acuminata, Pistia stratiotes, Pontederia cordat, Potentilla palustris, roots, lateral roots, pattern     

Distribution of Lateral Root Primordia in Root Tips of Musa

The distribution of lateral root primordia in the root tipsof four Musa landraces (Grande Naine, Pisang Berlin, Ngok Egomeand Yangambi Km5) grown in the field has been investigated toevaluate the range of genetic variation of lateral root initiation.In banana (Musa sp.), lateral roots are initiated in the roottip, 0.6–4 mm behind the root/cap junction and arise inseveral protoxylem-based longitudinal rows or ‘ranks’.Significant differences were observed among landraces for theposition of the most distal primordium, however the longitudinalspacing between successive primordia along the ranks was similarfor all landraces. All ranks were involved in lateral root initiation.The number of ranks also showed significant variations amonglandraces and was proportional to the stelar diameter. Hencethe density of lateral roots (roots cm^-1) was affected by stelardiameter variations. Finally, root elongation in the root tipwas landrace-specific and not necessarily exponential, unlikesuggested in previous studies. It is concluded that lateralroot initiation in Musa is not involved in the genetic variationsof root architecture in the field. A dissection of root architectureinto components which may account for these variations is proposedin relation to the improvement of root system architecture.Copyright 1999 Annals of Botany Company Lateral root initiation, root architecture, Musa, banana.     

Site, Scale and Time-course for Adjustments in Lateral Root Initiation in Wheat Following Changes in C and N Supply

The pattern of lateral root initiation in seminal roots of wheat(Triticum aestivumL. cv. Alexandria) and the location, scaleand time-course for adjustments in initiation were studied afterchanges in C and N supply. Macroscopically visible primordiaappeared in a non-acropetal sequence with the frequency (numberper unit length) increasing with distance behind the main rootapex to a maximum at 40–50 mm behind the root tip. Pruningthe root system to a single seminal axis increased the primordiafrequency by 23% within 15 h. After longer periods, the effectof root-pruning was greater. The enhanced primordia frequencywas first observed in tissue located 0–10 mm behind theapex at the start of treatment. Feeding glucose (50 mM) alsoincreased primordia frequency within 15 h, but to a greaterextent, and here additional primordia were initiated in tissuelocated 0–10and10–20 mm behind the apex at the startof treatment. Withdrawing NO₃^-from one part of a split-rootsystem, whilst maintaining the supply to the other, reducedprimordia frequency in the non-fed roots and, in some cases,a compensatory increase in the NO₃^--fed roots was observed.The location and scale of the adjustments were similar to thosefound with root-pruning and glucose-feeding, but were slightlyslower to appear. In spite of some differences in detail, therewas a broad similarity in site, scale and time-course for adjustmentsin lateral root initiation with these treatments, which is consistentwith the operation of a common mechanism. Whenever an increasein primordia frequency was observed, it was associated withan increase in the ethanol-soluble sugar content of the tissue.However, the reduction in frequency in NO₃^--deprived roots wasalso accompanied by an increase in sugar content. There wasno consistent relationship between total N content of the tissueand primordia frequency, but there was between primordia frequencyand the rate of net NO₃^-uptake. The possible mechanisms controllinglateral root initiation are discussed. Compensatory growth; correlative growth; glucose; initiation; lateral root; nitrate; primordium; split-root; Triticum aestivum; wheat     

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     

Genetic Diversity of the Wild Banana Musa acuminata Colla in Malaysia as Evidenced by AFLP

Musa acuminata Colla (Musaceae), the wild progenitor of thecultivated banana, is highly variable in Malaysia and presentsseveral unresolved nomenclatural problems. AFLP was employedto distinguish among three subspecies of Musa acuminata(subsp.truncata and subsp. malaccensis from peninsular Malaysia andsubsp. microcarpa from Borneo) and to examine whether subsp.truncata is a distinct taxon. Eight primer combinations revealedmolecular markers specific for each of the three taxa. UPGMAcluster analysis showed the three taxa were distinct. Subspeciesmalaccensis which is endemic in peninsular Malaysia and subsp.microcarpa which is endemic in Borneo were found to be moresimilar to each other in their DNA patterns than they are tosubsp. truncata, which is endemic to peninsular Malaysia. Sincesubsp. truncata is genetically separate from subsp. malaccensisand subsp.microcarpa , it cannot be regarded as synonymous witheither of these subspecies. This paper sheds light on the nomenclatureof the three subspecies of Musa acuminata. Copyright 2001 Annalsof Botany Company Musa acuminata Colla, truncata, malaccensis, microcarpa, Musaceae, wild banana, genetic diversity, AFLP, DNA fingerprinting     

Lateral Root Development in Attached and Excised Roots of Zea mays L. Cultivated in the Presence or Absence of Indol-3-yl Acetic Acid

The initiation of lateral root primordia and their subsequentemergence as secondary roots have been examined in attachedand excised roots of Zea mays grown in the presence or absenceof indol-3-yl acetic acid (IAA). Exposure to IAA enhanced anlageinception in both batches of roots. In the attached roots, theIAA-induced stimulation of primordium initiation was followedby a similar increase in lateral emergence. IAA treatment, however,had no effect on the number of laterals produced, per centimetreof root, in the excised primaries. Thus, exposure to IAA didnot directly enhance lateral emergence in the attached rootsnor did it stimulate such emergence in the excised ones. Nocorrelation was found between proliferative activity in themeristem at the apex of the primary or the rate of root elongationon the one hand, and either the number of primordia initiated,or the number of laterals produced, per centimetre of primary,on the other. Zea mays, maize, root, primordium, lateral, indol-3-yl acetic acid, meristematic activity     

Lateral Root Anlage Development in Excised Roots of Vicia faba L., Pisum sativum L., Zea mays L. and Phaseolus vulgaris L.

The development of lateral root primordia has been investigatedin excised roots of Vicia faba, Pisum sativum, Zea mays andPhaseolus vulgaris cultured in White's medium supplemented with2 per cent sucrose and compared with previously published dataon such development in primaries of the corresponding intactplants (control roots). Primordia were produced in each batchof excised roots over the 6 day culture period but at a lowerrate (number day^–1) than in the controls. Such primordia in cultured roots of Zea and Phaseolus completedtheir development and grew out as lateral roots over a periodsimilar in length to that found in the controls, but with acell number of only about 33 per cent of that attained at thetime of secondary emergence in the primaries of the latter roots.These lower cell numbers were at least partly a reflection ofincreases in mean cell doubling time over the period of anlagedevelopment investigated in the excised roots relative to thecorresponding values found in the controls. Primordia initiated in excised roots of Pisum and Vicia didnot complete their development in culture, i.e. no lateral rootsemerged and arrest took place with cell numbers of only 37 (Pisum)and 17 (Vicia) per cent of the numbers determined at the timeof secondary root emergence in the controls. Such arrested primordiahad few nuclei in S and none in mitosis. Moreover, at leastin Pisum, the frequency distribution of the relative DNA contentof the nuclei in the latter primordia approximated that foundin the apical meristem of primary roots following the establishmentof the stationary phase under conditions of carbohydrate starvation. It has also been demonstrated in the course of these investigationsthat lateral root primordium development in all four speciesis at least biphasic and possibly triphasic. Vicia faba L., broad bean, Pisum sativum L., garden pea, Zea mays L., maize, Phaseolus vulgaris L., dwarf bean, root primordia, anlage, cell doubling time, lateral root emergence     

Studies on the Roots of Musa acuminata cv. Gros Michel: 1. The Anatomy and Development of Main Roots

The anatomy of the adventitious roots of Musa acuminata cv.Gros Michel is described. Interesting anatomical features arethe occurrence of lysigenous lacunae in the cortex and the presenceof very large metaxylem vessels and internal phloem strandsin the stele. The organization of the root meristem is consideredin terms of present concepts. Histological preparations andautoradiographs suggest the existence of an area of relativequiescence in the meristem. The use of modified histogen terminologyin describing the apex is recommended. The development of thetissue systems is also described with emphasis on the differentiationof the vascular tissue.     

Patterns of Distribution of Lateral Root Primordia

The distribution of lateral root primordia has been examinedin Pistia stratiotes and Potentilla palustris, and re-examinedin Pontederia cordata. The major feature of the distributionpattern is a rather regular spacing along protoxylem-based ranks.There appears to be some correlation of activity between ranks,but simulations of patterns based on the data from Pontederiasuggest that the observed correlation is spurious. Pistia stratiotes, Pontederia cordata, Potentilla palustris, roots, lateral roots, pattern     

Waterlogging Tolerance Among a Diverse Range of Trifolium Accessions is Related to Root Porosity, Lateral Root Formation and 'Aerotropic Rooting'

Waterlogging tolerance, root porosity and root anatomy wereevaluated for 20 Trifolium accessions (species and sub-species,all annuals) selected from the eight Sections of the genus.Nine accessions were sensitive [relative growth rate (RGR) reducedby up to 80%] to waterlogging, nine accessions were tolerant(RGR not reduced), and in two accessions RGR increased (up to1.9-fold), when compared to drained controls. Growth of themain (i.e. tap) root axis was severely reduced in all accessionswhen waterlogged. Lateral roots formed the bulk of the rootsystem of tolerant accessions when grown in waterlogged soil.Lengths of the longest lateral roots were up to three-timeslonger than the main root axis. Root porosity varied from 0.7–12%among accessions when grown in aerated solution and from 1.1–15.5%in plants grown in hypoxic (0.031–0.045 mol O₂m^-3) solution.In some accessions aerenchyma formed by cell lysigeny; in othersit formed by schizogenous cell separation, or a combinationof both processes. O₂consumption rates of expanded lateral roottissues varied by up to 1.7-fold (on a mass basis) among thesix accessions tested and was reduced by an average of 24% forroots of plants grown in hypoxic solution prior to measurements.Accessions with the highest root porosity tended to have longerroots when grown in waterlogged soil. Three accessions formed‘aerotropic roots’ and the lateral root lengthsof these plants exceeded those of all other accessions, suggestingenhanced O₂movement to the submerged lateral root axis via theaerotropic roots. Waterlogging-tolerant accessions were identifiedin seven of the eight Sections in Trifolium, and the tolerantaccessions tended to be those with extensive lateral root systemsof relatively high porosity. Copyright 2001 Annals of BotanyCompany Waterlogging, Trifolium, aerenchyma, hypoxia, flooding, root respiration, clover, root anatomy, root porosity, pasture, aerotropic roots     

Effect of Main Root Decapitation on Root Branching in Flax Seedlings

Root branching patterns in intact and decapitated flax (Linum usitatissimumL.) roots were compared. The number of initiated primordia in the control and decapitated roots was similar, but decapitated roots produced an increased number of lateral roots owing to an increase in the number of primordia developed into the laterals. It is suggested that the apical meristem influences lateral root development only at the stage of root emergence from the parent root.     

Root Distribution, Growth, Respiration, and Hydraulic Conductivity for Two Highly Productive Agaves

Cultivated Agave mapisaga and A. salmiana can have an extremelyhigh above-ground dry-weight productivity of 40 Mg ha^–1yr^–1. To help understand the below-ground capabilitiesthat support the high above-ground productivity of these Crassulaceanacid metabolism plants, roots were studied in the laboratoryand in plantations near Mexico City. For approximately 15-year-oldplants, the lateral spread of roots from the plant base averaged1.3 m and the maximal root depth was 0.8 m, both considerablygreater than for desert succulents of the same age. Root andshoot growth occurred all year, although the increase in shootgrowth at the beginning of the wet season preceded the increasein growth of main roots. New lateral roots branching from themain roots were more common at the beginning of the wet season,which favoured water uptake with a minimal biomass investment,whereas growth of new main roots occurred later in the growingseason. The root: shoot dry weight ratio was extremely low,less than 0.07 for 6-year-old plants of both species, and decreasedwith plant age. The elongation rates of main roots and lateralroots were 10 to 17 mm d^–1, higher than for various desertsucculents but similar to elongation rates for roots of highlyproductive C₃ and C₄ agronomic species. The respiration rateof attached main roots was 32 µmol CO₂ evolved kg^–1dry weight s^–1 at 4 weeks of age, that of lateral rootswas about 70% higher, and both rates decreased with root age.Such respiration rates are 4- to 5-fold higher than for Agavedeserti, but similar to rates for C₃ and C₄ agronomic species.The root hydraulic conductivity had a maximal value of 3 x 10^–7ms^–1 MPa^–1 at 4 weeks of age, similar to A. deserti.The radial hydraulic conductivity from the root surface to thexylem decreased and the axial conductivity along the xylem increasedwith root age, again similar to A. deserti. Thus, although rootsof A. mapisaga and A. salmiana had hydraulic properties perunit length similar to those of a desert agave, their highergrowth rates, their higher respiration rates, and the greatersoil volume explored by their roots than for various desertsucculents apparently helped support their high above-groundbiomass productivity Key words: Crassulacean acid metabolism, productivity, root elongation rate, root system, water uptake     

Change in Size and Cell Number during the Development of Lateral Root Primordia in Zea mays L.

Increase in cell number, anlage volume and length have beeninvestigated during the overall period of lateral root primordiumdevelopment in excised primaries and in attached roots of Zeamays L. Each of these aspects of anlage growth was found toincrease more or less exponentially during the interval betweenprimordium initiation and subsequent emergence as a lateralin both batches of roots. Values were then determined for celldoubling time (Td), the size of the proliferative fraction (Pf),and for anlage volume (Tv) and length (Tt) doubling times duringthe overall period of primordium development and at intervalsduring this period in both the excised and attached roots. Thepattern of change which took place in Td, Tv, Tl and Pf duringlateral primordium development was found to be similar in bothbatches of roots. However, the overall period of anlage developmentwas shorter in the excised roots than in the attached ones.Moreover, when laterals grew out of the excised roots they didso with fewer cells than comparable laterals emerging from theattached roots. Zea mays L., maize, root primordia, lateral emergence, cell doubling time     

Mutations in Arabidopsis multidrug resistance-like ABC transporters separate the roles of acropetal and basipetal auxin transport in lateral root development

Auxin affects the shape of root systems by influencing elongation and branching. Because multidrug resistance (MDR)-like ABC transporters participate in auxin transport, they may be expected to contribute to root system development. This reverse genetic study of Arabidopsis thaliana roots shows that MDR4-mediated basipetal auxin transport did not affect root elongation or branching. However, impaired acropetal auxin transport due to mutation of the MDR1 gene caused 21% of nascent lateral roots to arrest their growth and the remainder to elongate 50% more slowly than the wild type. Reporter gene analyses indicated a severe auxin deficit in the apex of mdr1 but not mdr4 lateral roots. The mdr1 deficit was explained by 40% less acropetal auxin transport within the mdr1 lateral roots. The slow elongation of mdr1 lateral roots was rescued by auxin and phenocopied in the wild type by an inhibitor of polar auxin transport. Confocal microscopy analysis of a functional green fluorescent protein-MDR1 translational fusion showed the protein to be auxin inducible and present in the tissues responsible for acropetal transport in the primary root. The protein also accumulated in lateral root primordia and later in the tissues responsible for acropetal transport within the lateral root, fully supporting the conclusion that auxin levels established by MDR1-dependent acropetal transport control lateral root growth rate to influence root system architecture.     

Protein Changes Associated with Auxin-Induced Stimulation and Kinetin-Induced Inhibition of Lateral Root Initiation in Lettuce (Lactuca sativa) Roots

Protein changes associated with hormonal regulation of lateralroot initiation in lettuce (Lactuca sativa L.) roots were examined.Naphthalene acetic acid (NAA) stimulates the induction of lateralroots (Maclsaac, Sawhney, and Pohorecky, 1989) and this wasaccompanied by an increase in soluble proteins as well as thesynthesis of several polypeptides, including specific polypeptidesof 32 and 31 kD. The synthesis of these polypeptides coincidedwith the onset of cell division in the pericycle of NAA-treatedroots. Application of cycloheximide at different times showedthat NAA-induced protein synthesis is essential for the initiationand development of lateral root primordia. Kinetin inhibitedthe formation of lateral roots as well as the level of solubleproteins in NAA-treated roots. In addition, kinetin-treatedroots contained 22 and 21 kD polypeptides not found in othertreatments. This study suggests that the mechanisms of NAA-stimulationand kinetin-inhibition of lateral root initiation are probablydifferent. Key words: Lactuca sativa, lateral roots, proteins     

Root Growth Inhibitors from Root Cap and Root Meristem of Zea mays L.

A micro-assay based on the growth inhibition of root segmentsof the seminal roots of Zea mays has been used to investigatethe root-growth-inhibiting substances in root caps and meristemsrespectively of the roots of Zea mays. This micro-assay is sensitiveto 50 pg of IAA or less. Paper chromatography of the acid fractionof methanolic extracts shows the presence of one main inhibitorin root caps and a different main inhibitor in root meristems.Neither is IAA, whose presence in meristems is sometimes indicatedby small inhibitions (or stimulations) at the characteristicRf of IAA. A Commelina leaf-epidermis assay shows the presenceof one stomata-closing ABA-like substance in root caps and onein meristems, one corresponding in Rf to the main root-growthinhibitor from the root cap. The implications of these findingsfor the geotropic responses of roots is briefly discussed.     

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.     

Histology of Shoot Bud Ontogeny from Seedling Root Segments of Brassica napus L.

Root explants of Brassica napus cultured in vitro form adventitiousshoots. The root buds originated at the base of the newly initiatedlateral root. Cells in association with the differentiatingphloem of the developing lateral roots were the sites for rootbud formation. A nodular mass of cytoplasmic cells developedby day 7 at the base of the lateral root. This group of cellscontinued to divide an enlarge. The cells in the peripheralregion of the nodular cell mass differentiated further intoa meristematic zone. The meristematic cells grew towards theperiphery of the cortex by crushing the outer layer of corticalcells. Further development of the meristematic layer resultedin the formation of shoot primordia with organized shoot apicalmeristems and leaf primordia.Copyright 1993, 1999 Academic Press Brassica napus, canola, cultured root segments, root buds     

Effect of Localized Anoxia on Phaseolus vulgaris L. Root Growth

Heterogeneity within the root environment results in differentialgrowth within root systems. The response of five Phaseolus vulgarisL. cultivars to non-uniform root aeration was evaluated. Threetreatments were applied to a split root system for a periodof 72 h. Treatments consisted of an aerated control, a non-aeratedcontrol (both halves non-aerated, using N₂). and localized anoxia(one-half the root system aerated and the remaining half subjectedto N₂). Shoot and root growth were reduced in the anoxic controlbut not in the aerated control or localized anoxia treatment. Root growth was greatest in the aerated portion of the localizedanoxia treatment for all genotypes. Contributions of the rootcomponents to the compensatory responses differed dependingon the plant cultivar examined. The growth of branched and lateralroots present before the treatment period increased by 65% inline 31908. A 50% increase in the growth of lateral roots whichemerged during the treatment period occurred in another line(Swan Valley). Other genotypes responded in an intermediatemanner. These observations indicate differences in cultivarresponses to localized soil stress. Key words: Phaseolus vulgaris L., Anoxia, Root growth     

Pattern Formation in Cluster Roots: Some Developmental and Evolutionary Considerations

Cluster roots, also known as proteoid roots, are one of themajor adaptations for nutrient acquisition in terrestrial vascularsporophytes, occurring in many important plant families, inkey areas of biodiversity, and in significant ecological niches.Their development and function are closely linked and presentan ideal experimental system with which to investigate the basisof pattern and its morphogenetic amplification. Both meristemfunction and root initiation are controlled within a spatialand temporal framework, resulting in predictive phenotypic expression.In this paper, these patterns of development are described withinthe context of our knowledge of lateral root initiation anddevelopment. Recent work is summarized in an attempt to highlightthe issues of most importance for future research. The caseof the genus Lupinus is taken as a means of exploring the phylogeneticrelationships of species with cluster roots. The first evidenceof cluster roots having arisen only once within the Lupinusgenus is presented. Copyright 2000 Annals of Botany Company Cluster root, development, lateral root, lupin evolution, nutrient acquisition, pattern, proteoid root