Root Initiation in Cuttings of Pinus radiata Seedlings: I. DEVELOPMENTAL SEQUENCE

Adventitious root initiation in hypocotyls of Pinus radiataD. Don consists of three distinct phases. In the pre-initiativephase, following selection of the cuttings, no histologicalchanges are observed. The initiative phase begins with swellingof a single parenchymatous cell, the meristematic locus. Distinctcytoplasmic changes within the meristematic locus are followedby cytoplasmic migration and asymmetric division in the surroundingcortical or peripheral cells. The third, or post-initiative,phase includes continued division of derivatives of the peripheralcells to form meristemoids which subsequently differentiateinto root primordia.     

Auxin and Ethylene-stimulated Adventitious Rooting in Relation to Tissue 9

We have previously shown that both endogenous auxin and ethylenepromote adventitious root formation in the hypocotyls of derootedsunflower (Helianthus annuus) seedlings. Experiments here showedthat promotive effects on rooting of the ethylene precursor,1-aminocyclopropane-l-carboxylic acid (ACC) and the ethylene-releasingcompound, ethephon (2-chloro-ethylphosphonic acid), dependedon the existence of cotyledons and apical bud (major sourcesof auxin) or the presence of exogenously applied indole-3-aceticacid (IAA). Ethephon, ACC, aminoethoxyvinylglycine (an inhibitorof ethylene biosynthesis), and silver thiosulphate (STS, aninhibitor of ethylene action), applied for a length of timethat significantly influenced adventitious rooting, showed noinhibitory effect on the basipetal transport of [³H]IAA. Theseregulators also had no effect on the metabolism of [³H]IAA andendogenous IAA levels measured by gas chromatography-mass spectrometry.ACC enhanced the rooting response of hypocotyls to exogenousIAA and decreased the inhibition of rooting by IAA transportinhibitor, N-1-naphthylphthalamic acid (NPA). STS reduced therooting response of hypocotyls to exogenous IAA and increasedthe inhibition of rooting by NPA. Exogenous auxins promotedethylene production in the rooting zone of the hypocotyls. Decapitationof the cuttings or application of NPA to the hypocotyl belowthe cotyledons did not alter ethylene production in the rootingzone, but greatly reduced the number of root primordia. We concludethat auxin is a primary controller of adventitious root formationin sunflower hypocotyls, while the effect of ethylene is mediatedby auxin. Key words: Auxin, ethylene, adventitious rooting, sunflower     

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     

Adventitious rooting in hypocotyls of sunflower (Helianthus annuus) seedlings. IV. The role of changes in endogenous free and conjugated indole‐3‐acetic acid

We have studied the role of endogenous auxin on adventitious rooting in hypocotyls of derooted sunflower (Helianthus annuus L. var. Dahlgren 131) seedlings. Endogenous free and conjugated indole-3-acetic acid (IAA) were measured in three segments of hypocotyls of equal length (apical, middle, basal) by using gas chromatography-mass spectrometry with [¹³C₆]-IAA as an internal standard. At the time original roots were excised (0 h), the free IAA level in the hypocotyls showed an acropetally decreasing gradient, but conjugated IAA level increased acropetally; i.e. free to total IAA ratio was highest in the basal portion of hypocotyls. The basal portion is the region where most of root primordia were found. Some primordia were seen in this region within 24 h after the roots were excised. The quantity of free IAA in the middle portion of the hypocotyl increased up to 15 h after excision and then decreased. In this middle region there were fewer root primordia, and they could not be seen until 72 h. In the apical portion the amount of free IAA steadily increased and no root primordia were seen by 72 h. Surgical removal of various parts of the hypocotyl tissues caused adventitious root formation in the hypocotyl regions where basipetally transported IAA could accumulate. Reduction in the basipetal flow of auxin by N-1-naphthylphthalamic acid and 2,3,5-tri-iodobenzoic acid resulted in fewer adventitious roots. The fewest root primordia were seen if the major sources of endogenous auxin were removed by decapitation of the cotyledons and apical bud. Exogenous auxins promoted rooting and were able to completely overcome the inhibitory effect of 2,3,5-tri-iodobenzoic acid. Exogenous auxins were only partially able to overcome the inhibitory effect of decapitation. We conclude that in sunflower hypocotyls endogenously produced auxin is necessary for adventitious root formation. The higher concentrations of auxin in the basal portion may be partially responsible for that portion of the hypocotyl producing the greatest number of primordia. In addition to auxins, other factors such as wound ethylene and lowered cytokinin levels caused by excision of the original root system cuttings must also be important.     

Development of Lateral Primordia in Decapitated Adventitious Roots of Allium cepa

In decapitated adventitious roots of Allium cepa L. var. French,the time taken for lateral primordia to appear in five sectorshas been studied. This time is approx. 4–5 d after theapex of the adventitious root gave rise to the primordial initialcells. Allium cepa, adventitious root, lateral primordia     

Reserve Lipids in Stem Root Primordia of Poplar

Lipids are reserve substances in stem root primordia of poplarand occur as droplets approximately uniformly in all cells ofroot primordia. In ultrathin sections they appear as sphericalelectron-dense particles (0.5–1.4 µm diameter),filling up virtually all the available space in the cytoplasm.The presence of lipids in stem root primordia was demonstratedin several species of the genus Populus from the section Aigeirosand Tacamahaca, as well as in another representative of thefamily Salicaceae, viz. in the genus Salix.     

The RHG Gene Is Involved in Root and Hypocotyl Gravitropism in Arabidopsis thaliana

In higher plants, shoots show negative gravitropism and rootsshow positive gravitropism. To elucidate the molecular mechanismsof root and hypocotyl gravitropism, we segregated the secondmutation from the original phyB-1 mutant line which impairedboth root and hypocotyl gravitropism and characterized thisnovel mutation named rhg (for root and hyzypocotyl gravitropism).The rhg is a single recessive nuclear mutation and it is mappedon the lower part of the chromosome 1. Analyses on the gravitropicresponses of the rhg mutant indicate that root and hypocotylgravitropism are severely impaired but inflorescence stem gravitropismis not affected by the rhg mutation. In the rhg mutant seedlings,amyloplasts (statoliths for gravity-perception) were presentin the presumptive statocytes of roots and hypocotyls. Phototropismby roots and hypocotyls was not impaired in the rhg mutant.These results suggest that the RHG gene product probably actson the gravity-perception and/or the gravity-signal transductionin root and hypocotyl gravitropism. This is the first reportabout the genetic locus specifically involved in both root andhypocotyl gravitropism but not inflorescence stem gravitropism,supporting our hypothesis that the mechanisms of gravitropismare genetically different between hypocotyls and inflorescencestems. (Received March 11, 1997; Accepted April 17, 1997)     

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     

Adventitious Root Formation in Veronica Spp.

Species of the genus Veronica differ in habitat preferences,growth form and in adventitious root production. The annualspecies rarely or never produce adventitious roots in intactplants in the field but some, for example V. persica and V.arvensis will root vigorously from single node stem segmentsin culture. Others, such as V. agrestis require the presenceof IAA for substantial levels of root formation to occur incultured stem segments. Veronica hederifolia cuttings rarelyproduce roots. Stem cuttings of the perennial species, in general,rooted more vigorously than those of annual plants. Both V.fihiformis and V. serpyllifolia root very strongly. The position of root production from the stem cuttings differedfrom species to species. Roots arose either from the node, theregion of the base or at some intermediate point. Veronica arvensis,V. chamaedrys and V. persica rooted mainly from the basal regionwhereas V. filiformis rooted mainly from the node. Veronicaserpyllifolia cuttings rooted at both of these locations. Veronica filiformis, a perennial species that is infertile inBritain, produces root primordia in intact plants at nodes whichare close to the shoot apex. Thus, even very young stem segmentshave ‘preformed’ root primordia. For this reason,detached stem segments of V. filiformis root very rapidly andthis probably has been of great significance in its successfulinvasion and spread in lawns and short turf areas. Veronica spp., adventitious roots, indol-3-ylacetic acid, root primordia, vegetative reproduction     

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     

Growth and Histological Changes During Transition to Dormancy in the Regeneration Buds of Hordeum bulbosum L.

Growth and histological changes in a regeneration bud of Hordeumbulbosum during transition to dormancy were studied. Activeformation of new-leaf primordia on the elongating apex accompaniedby arrest of cell division in the lower leaf primordia characterizedthe first period. When activity subsequently in the distal partof the bud decreased, exillary buds and root primordia werestill actively being produced in its basal part.     

A Transgenic Mutant Defective in Cell Elongation and Cellular Organization during Both Root and Shoot Development in Lettuce, Lactuca sativa

A recessive mutation affecting both root and shoot developmentwas isolated from transformants of lettuce cv. Diana co-transformedwith maize Ac transposase and Ds. Mutant phenotype in the progenywas co-segregated with the T-DNA containing Ac transposase indicatingthat the mutation was caused by insertional mutagenesis. Mutationin this novel genetic locus, designated as ANORMAL ROOT ANDSHOOT (ARS), showed a dwarf phenotype with short thick roots,short hypocotyl and abnormal filamentous leaves without anyfurther reproductive development. The anatomical analysis revealedthat the ars mutant root phenotype is primarily due to the lackof cell elongation and to the abnormal increase in cell numberin the cortex region in the roots. ars mutants are able to initiatenormal leaf primodia, but, the cell elongation and cellularorganization of the developing leaf primodia is impaired andresults in sequentially abnormal development of the leaf. arsmutants also display photomorphogenic development in darknessby producing open cotyledons, developing vegetative leaves,and short hypocotyls. This suggests that the ARS gene may alsobe involved in the regulation of cell elongation in the hypocotylin the absence of light. Abnormal development in ars mutationscan not be normalized by exogenous application of phytohormonessuch as gibberellin and brassinosteroids, indicating that themutant is not impaired in the biosynthesis of these two hormones. (Received January 5, 1999; Accepted August 19, 1999)     

Control by Light of Hypocotyl Elongation and Levels of Endogenous Gibberellins in Seedlings of Lactuca sativa L.

Four 13-hydroxygibberellins, gibberellin A₁ (GA₁), 3-epi-GA₁,GA₁₉ and GA₂₀ were identified by full-scan GC/MS in extractsof lettuce seedlings (Lactuca sativa L. cv. Grand Rapids). Theresults suggest that the early-13-hydroxylation biosyntheticpathway to GA₁ functions in the lettuce seedlings. It was alsofound that GA₁ is active per se in the control of hypocotylelongation in lettuce seedlings. To investigate the relationshipbetween control by light of hypocotyl elongation and levelsof endogenous GAs in lettuce, endogenous levels of GAs werequantified by radioimmunoassay in seedlings that had been grownfor 5 days in the dark (5D) and in those that had been grownfor 4 days in the dark and then under white light for 1 day(4D1L). The endogenous level of GA₁ in the upper and lower partsof hypocotyls in 5D seedlings was about four times higher thanthat in 4D1L seedlings. The response of explants (hypocotylsegments with cotyledons) from dark-grown seedlings to GA₁ isknown to be similar in the dark and under white light when theexplants are treated with inhibitors of the biosynthesis ofGA. Therefore, the above information suggests that the highlevel of GA₁ in hypocotyls of dark-grown seedlings is responsiblefor the rapid elongation of hypocotyl, while irradiation bywhite light decreases the endogenous level of GA₁ in the hypocotylswith a resultant decrease in the rate of hypocotyl elongation. (Received March 13, 1992; Accepted May 21, 1992)     

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     

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.     

The Effect of Mechanically Induced Stress on the Growth of Cauliflower, Lettuce and Celery Seedlings

Brushing cauliflower, lettuce and celery seedlings with paperfor 1.5 min each day for 11–13 d, 10–12 d or 21–28d, respectively resulted in smaller, more compact, plants thanthe unbrushed controls. In all three species shoot fresh anddry weights and leaf area were reduced following brushing. Incauliflower and celery the largest growth reduction was in petiolelength. In lettuce, which has no discernible petioles, the reductionin leaf length caused by brushing was proportionally greaterthan the reduction in leaf width. Brushing reduced hypocotyllength in cauliflowers and to a lesser extent in lettuce. Petioleand hypocotyl thickness was reduced in cauliflower, whereashypocotyl thickness was increased in lettuce following brushing.Brushing increased leaf thickness in cauliflower, celery andto a lesser extent in lettuce and increased the percentage drymatter content of lettuce shoots. The weight of chlorophyllper fresh weight of leaf tissue increased following brushingin celery and lettuce and declined in cauliflowers. Root length and the number of branches per root system werereduced in all three species following brushing. Root dry weightwas reduced and the root:shoot dry weight ratio was increasedin lettuce, reduced in celery and unaffected in cauliflowers. There were different patterns of response to brushing, the reductionin leaf weight being greatest in the youngest leaf of cauliflowerand least in the youngest leaf of lettuce and celery. Growthresponses to brushing were seen several days after brushinghad ceased, noticeably in leaves which were barely visible atthe time of brushing. It is suggested that growth retardation of cauliflowers, lettuceand celery, induced by mechanical stress such as brushing mayprove valuable as a means of ‘conditioning’ theseedlings to withstand both the physical and physiological stresseswhich occur at and during transplanting. Brassica oleracea, cauliflower, Lactuca sativa L., lettuce, Apium graveolens L., celery, mechanical stress, shoot growth, root growth, chlorophyll     

角茴香根水浸液对生菜的化感潜力及机理初探

以生菜为受体植物,使用培养皿法研究了角茴香根水浸液的化感潜力,并对其可能的机理进行了初步探讨.结果表明:不同浓度角茴香根水浸液对生菜种子的萌发、幼苗的根和下胚轴的生长以及植株鲜重均有显著抑制作用,而凡处理液浓度越高,生菜生长所受到的抑制程度越深;随着角茴香根水浸液处理浓度的增大,生菜幼苗中超氧阴离子自由基的含量逐渐升高,其抗氧化酶如SOD、POD和CAT活性比对照显著增强.研究发现,角茴香根水浸液中可能含有较强活性的化感物质,使生菜幼苗受到了明显的活性氧伤害,也因此而表现出较强的化感潜力,但生菜通过上调体内抗氧化酶活性来应对这种伤害.     

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     

Are Lateral Roots of Tomato Spaced by Multiples of a Fundamental Distance?

Roots of tomato (Lycopersicon esculentum Miller) were culturedin vitro for 8-16 d at the end of which period the number ofdaughter roots (first order laterals), and their distributionon the mother root axis, were recorded. It was observed thatthe daughter roots are distributed along the mother root intwo series, and that their longitudinal distribution, in bothseries and in the total series, was non-random, in the senseof being systematically spaced out rather than systematicallyclustered. There was an approximately two-thirds probabilitythat a daughter root would be on the opposite side of the rootfrom its closest sister in the proximal direction, but thisfinding seemed explicable by the observed 'spaced-out' distributionin each series, without invoking any interaction between thetwo side series. Earlier, it had been suggested that distancesbetween successive sister roots in a series are multiples ofa quantum distance, probably because the daughter roots observedare a subset of a set of evenly spaced candidate root primordia,the rest of which are developmentally arrested. The sister-sisterdistances were analysed statistically for the presence of sucha quantum. However, although some daughter root series showeda statistically significant quantal tendency, this did not seemto be a general feature of the mother roots.Copyright 1993,1999 Academic Press Angiosperms, Lycopersicon esculentum, tomato, lateral root, root primordia, roots in vitro, cosine quantogram     

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