Anatomical Changes Which Occur in Cuttings of Agathis australis (D. Don) Lindl 1. Wounding Responses

The basal and sub-basal regions in cuttings of Agathis australisundergo a complex series of anatomical changes. Many of theseare categorized as wound responses and include cell divisionsassociated with the cut base and the proliferation of tracheidsand phloem which arise in the interfascicular region about 4mm above the cut base. The vascular tissue arcs outwards anddownwards through the cortex. It may develop as isolated strandsonly a few cells wide or as sheets involving a number of cells.The precise pattern of vascular development appears to be determinedby its extent at the point of origin and by the presence ofobstacles such as primary and secondary resin canals which arelocated to the outside of the vascular bundles in the stem.Secondary resin canals are produced only in the rooting zonein cuttings that show extensive cell division. They arise schizogenouslyand do not form an interlinking network. Root primordia arise in the cortex at the end of isolated strandsof newly developed vascular tissue. Primordia never form inassociation with sheets of tracheids or after the convergenceof strands. In some cases virtually the entire sub-base is filledwith vascular tissue as a result of cell division and the differentiationof parenchymatous tissue. Root primordia never appear in thissituation. Agathis australis (D. Don) Lindl, kauri, cuttings, wound responses, vascular development, resin canals, root primordia, cellular differentiation     

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     

The Anatomy of Root Initiation in Cuttings of Griselinia littoralis and Griselinia lucida

The initiation of root primordia in Griselinia littoralis andG. lucida occurred a few millimeters above the cut base. Inall cases the first cellular event was the activation of specificregions of the cambium which were associated with traces fromleaves which themselves were particularly vigorous root-formersif detached. In cuttings from seedlings, the first cells cutoff continued to divide and gave rise to root primordia in situ,whereas those from mature plants produced files of six to 22rows of cells and it was only when the cells first cut off hadbeen displaced outwards towards the fibre caps that the cambiumstopped dividing. Primordium development may then occur at theadvancing front, possibly as a result of a stimulus originatingfrom damaged cells. Thus the same cells were involved in organizedprimordium formation both in seedling and mature cutting, butthe location differed. The reduced rooting and the long lag phase in cuttings frommature plants of G. lucida may be due to the presence of a completering of fibres. This does not seem to present a mechanical barrierto emergence - since elongating primordia can force throughthe ring - nor can it affect the receipt of any root-inducingstimulus by the target cells since it is considered that thisarrives via the vascular tissues. However, evidence suggeststhat it is the initial phase of cambial activation which isprevented in cuttings from mature plants. This suggested eitherthat the initial stimulus was less effective or that the targetcells were less responsive than those of seedlings, which havevery few fibres, or of mature G. littoralis, which only hasfibm associated with the vascular bundles. Griselinia littoralis Raoul Choix, Griselinia licida Forst. f. Prodr., adventitious roots, cuttings, primordium initiation and location     

Adventitious root initiation in hypocotyl and epicotyl cuttings of eastern white pine (Pinus strobus) seedlings

The present paper reports results of experiments to develop a system for studying adventitious root initiation in cuttings derived from seedlings. Hypocotyl cuttings of 2-week-old eastern white pine (Pinus strobus L.) seedlings were treated for 5 min with 0, 100, 200, 300, 400, 500 or 600 mg l^?1 (0, 0.54, 1.07, 1.61, 2.15, 2.69 or 3.22 mM) 1-naphthaleneacetic acid (NAA) to determine the effect on root initiation. The number of root primordia per cutting was correlated with NAA concentration and the square of NAA concentration. Thus, the number increased from less than one per cutting in the 0 NAA treatment to approximately 40 per cutting at 300 mg l-1 NAA, above which no substantial further increase was observed. The larger number of root primordia formed in response to increasing concentrations of NAA was due to the formation of primordia over a larger proportion of the hypocotyls. Histological analysis of the timing of root primordium formation in hypocotyl cuttings revealed three discernible stages. Progression through these stages was relatively synchronous among NAA-treated hypocotyl cuttings and within a given cutting, but variation was observed in the portion of different cuttings undergoing root formation. Control-treated hypocotyl cuttings formed root primordia at lower frequencies and more slowly than NAA-treated cuttings, with fewer primordia per cutting. Epicotyl cuttings from 11-week-old seedlings also formed adventitious roots, but more slowly than hypocotyl cuttings. NAA treatment of epicotyl cuttings caused more rapid root initiation and also affected the origin of adventitious roots in comparison with nontreated cuttings. NAA-treated epicotyl cuttings formed roots in a manner analogous to that of the hypocotyl cuttings, directly from preformed vascular tissue, while control-treated epicotyl cuttings first formed a wound or callus tissue and subsequently differentiated root primordia within that tissue. This system of inducing adventitious roots in pine stem cuttings lends itself to studying the molecular and biochemical steps that occur during root initiation and development.     

Cortical Cell Breakdown and Lateral Root Primordium Development in Vicia faba L

The effect of the formation of a cavity in the cortex of theprimary root of Vicia faba adjacent to lateral root primordiaon root development has been investigated. Premature exposureof such primordia to the external medium by removing the overlyingtissues of the primary root has no effect on primordium developmentif that primordium was within 48 h of emerging as a lateralroot. Similar exposure of primordia which were at an earlierstage of development and consisted of between 3400 and 7000cells resulted in the generation of a stationary phase, withmost of the nuclei arrested in G1 (presynthetic interphase),48–72 h after exposure began, followed by nuclear degenerationby 96 h. Since no mature vascular tissue was found in theseprimordia until after they emerged as secondary roots, all ofthe nutrients necessary for the maintenance of cell proliferationin these meristems must reach them by simple diffusion fromthe surrounding medium. A preliminary analysis of the liquidcontents of the cavity next to developing primordia demonstratesit to be rich in carbohydrates and it is clear, from the resultsreported in this paper, that cell proliferation in primordia,consisting of a mean number of 5400 cells, is largely dependenton the substances present in the cavity fluid, although somematerials reach the primordium by diffusion from the cells ofthe primary root to which the primordium remains attached.     

欧美杂种山杨微扦插不定根发生过程的解剖学研究

采用石蜡切片技术,以欧美杂种山杨插穗基部茎段为实验材料,连续解剖观察插穗不定根发生发育过程,分析根原基发生部位与扦插生根的关系。结果显示:欧美杂种山杨插穗不定根的发生过程分为4个时期,为根原基诱导期,不定根起始期、表达期和伸长生长期。根原基诱导期维管形成层产生具有分生组织特点的薄壁细胞;不定根起始期,维管形成层及附近的薄壁细胞脱分化,形成不定根原基发端细胞;不定根表达期,根原基发端细胞不断分裂成具有方向性的根原基,根原基穿过韧皮射线和皮层,向皮孔方向发展;不定根伸长生长期,根原基从皮孔伸出,其内部的维管系统开始发育,形成不定根。研究认为,欧美杂种山杨为皮部诱导生根类型,不定根原基起源于维管形成层区,起源部位单一,扦插难生根。     

Factors Influencing Adventitious Root Production in Cuttings of Griselinia littoralis and Griselinia lucida

Griselinia littoralis roots quickly and vigorously from cuttingsof seedlings and mature plants and also forms roots on detachedleaves. Cuttings root in the dark but leaves must be present.In contrast G. lucida roots vigorously only from cuttings takenfrom seedlings. Light is essential for root formation on oldermaterial. Detached leaves will not root. Although callus formationat the cut base commonly occurs in both species it is not directlyassociated with root formation. Griselinia littoralis Raoul Choix, Griselinia lucida Forst. f. Prodr., adventitious roots, stem cuttings, leaf cuttings, woody plants     

红皮云杉茎的解剖结构与插条不定根形成的研究

１９９２年７－８月定时固定红皮云杉插条基部材料于ＦＡＡ液中,石蜡制片法室内解剖研究不定根的发生。结果表明：红皮云杉插条诱发根原基的来源有两种途径。一种是愈伤组织生根型,在愈伤组织的再生形成层处,或茎的维管形成层诱发根原基;另一种是非愈伤组织生根型,在插条切口处的维管形成层、皮层或初生木质部与次生木质部间的薄壁组织较深的部位,直接产生纵向不定根原始体,有的在距离切口０．１－０．５ｃｍ以上茎的维管形成层,维管形成层与木射线的交界处及叶隙等薄壁组织产生径向不定根。不同个体间产生的不定根数量及发育的早晚差异较大。     

胡杨不定根原基发生的分生细胞结构特征及内源激素变化分析

以胡杨实生苗去根插穗为试材,采用显微技术、气质(GC/MS)联用技术和同工酶分析技术观察和研究了外源3-吲哚丁酸(IBA)对其不定根原基(干细胞)形成的影响.结果表明:(1)胡杨插穗培养在无IBA的培养基上,36 h时有大核细胞发生,60 h时大核细胞聚集成团形成根原基,72 h时生成不定根;而添加了IBA的培养基上插穗培养60 h时大核细胞分散,出现许多薄壁细胞,72 h时薄壁细胞变大,没有根原基的形成和根的发生,表现出细胞组织愈伤化,不再分化出干细胞.添加外源IBA抑制了胡杨插穗不定根形成.(2)组织化学观察显示,在无IBA的培养基上的插穗,60 h时具双环状核仁的干细胞细胞质浓,富含蛋白质;而此时添加了IBA的插穗,具双环状核仁的细胞细胞质很少,蛋白质含量也少.(3)整个胡杨不定根形成的过程中,内源激素IAA和ABA可能作为诱导根原基发生的重要信号分子;内源激素IAA和GA3处于一个较稳定状态,有利于不定根的形成.(4)过氧化物酶(POD)2b酶带的持续表达有利于根原基的诱导;淀粉酶在60 h时表达增强,是根原基发育的标志;根原基诱导时不需要酯酶,而根原基发育时需要酯酶.     

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     

Involvement of the Cotyledon in Adventitious Root Initiation in Impatiens balsamina L. Seedlings

Adventitious root primordia are found in the pre-hypocotyl tissueof developing seeds of Impatiens balsamina L. by the third weekafter petal drop, and are present in the mature seed. Aftergermination, the adventitious roots emerge from a collet swellingon the hypocotyl of the young seedlings. Removal of the colletduring the first five days results in the formation of anotherat the base of the remaining hypocotyl. Older seedlings respondto the excision of the collet by producing one or more rootsnear the cut end, unless the cut is made close to the cotyledon,when, even in nine-day seedlings, a reduced collet is formedassociated with four or fewer roots. The influence of the cotyledonon collet/root regeneration diminishes in older seedlings andin these is manifested only in hypocotyl tissue adjacent tothat organ. Impatiens balsamina, balsam, cotyledon, adventitious roots, collet     

Influence of Aryl Esters of Indole-3-acetic and Indole-3-butyric Acids on Adventitious Root Primordium Initiation and Development

Synthetic aryl esters of indole-3-acetic acid (IAA) and indole-3-butyric acid (IBA) greatly enhanced adventitious root primordium initiation in bean (Phaseolus vulgaris L. cv. Top Crop) and jack pine (Pinus banksiana Lamb.) cuttings, respectively. Bean cuttings produced 95 to 154% more macroscopically visible root primordia in 2 days when treated with phenyl indole-3-acetate (P-IAA), in comparison with an equal concentration of IAA. Substantial but lesser increases occurred when treatment was done with 3-hydroxyphenyl indole-3-acetate (3HP-IAA). On a molar basis, either P-IAA or 3HP-IAA were 10 or more times as efficient as IAA in inducing adventitious root primordium initiation in bean cuttings. Methyl indole-3-acetate was no more effective than IAA in these tests. Phenyl indole-3-butyrate (P-IBA) consistently enhanced the number of rooted jack pine seedling cuttings by 11 to 12% in comparison with a 27% higher concentration of IBA. The number of elongated roots (2 mm or more) after 5 days was 165 to 276% greater for P-IAA than for IAA-treated bean cuttings. Similar but lesser increases occurred as a result of 3HP-IAA treatment. P-IBA in comparison with IBA treatment did not influence either the number of roots or length of the longest root per rooted jack pine cutting. Enzymes in bean and jack pine cuttings hydrolyzed the aryl esters. However, check experiments showed that initial integrity of the esters was required for enhanced activity in inducing root primordium initiation. Treatment of bean cuttings with hydrolysates of P-IAA, or with IAA and phenol, alone or combined, did not influence root primordium initiation or development in a manner different from treatment with IAA alone.     

The Anatomical Relationship Between Cambial Regeneration and Root Initiation in Wounded Winter Cuttings of the Apple Rootstock M.26

Root primordia differentiate remote from the existing vasculartissue of split- or incision- wounded winter cuttings. Thisis preceded by heavy callusing of the wounds and basal cut surface.Most of the callus is cortical in origin but callus is alsoformed as a result of damage to the cambium. In the incisiontreatment a new cambium differentiates within the cambial callusfrom the undamaged cambium on either side so as to form an outward-pointingsalient enclosing randomly orientated xylem. Two such salientsare formed in the split base treatment since each resultanthalf behaves like a separate cutting. The salients are subdividedhorizontally into finger-like projections due to the dispositionof the rays at the edges of the wound. Once a root primordiumforms, differentiation of procambium-like tissue between itand one or more projections proceeds rapidly, followed by outgrowthof the root. Roots emerge from the basal callus and in verticalfiles from the wound callus. The split base treatment increases16-fold the number of rooted cuttings over controls, while incisionwounds increase rooting four-fold; this is associated with thefact that more rhizogenic (salient-forming) sites are formedby splitting the base. The physical and anatomical factors involvedin cambial regeneration are discussed. Anatomy, apple, callus, cambium, cambial regeneration, M.26 rootstock, Malus pumila L., rooting, root initiation, wound response     

Endogenous Development of Adventitious Root Primordia in Lettuce Hypocotyls

Rudimentary adventitious root primordia were observed in optically-sectionedFeulgen-stained hypocotyls of 8-d-old lettuce seedlings germinatedin the dark and in a low light intensity environment. It issuggested that formation of these primordia may represent anaspect of normal development in the seedling. Adventitious, root primordia, hypocotyls, lettuce, Lactuca sativa     

Morpho-Histological Investigation of Plantlet Formation In Vitro in Taiwania flousiana Gaussen

Histological events during adventitious shoot formation in cultured shoot apex of 10–12-day-old seedlings and adventitious root formation in the elongated shoot of Taiwania floudana Gaussen were examined. Ceils of the peripheral subsurface layers of the shoot apex responded to cytokinin and divided into meristematic cells from which the shoot primordia were proliferated. A few bud primordia also originated from the epidermis and hypodermis of the adaxial surface of the cotyledon. The parenchyma of leaf gap of the shoots cultured in rooting medium dedifferentiated to regain the capacity of division and form adventitious root. Besides, cells that had relatively low potential of differentiation, such as the cortex parenchyma, pith ray, phloem parenchyma and cambium zone, albeit initiated to divide, but seldom formed root primordium. The origin of the adventitious roots in the leaf gap facilitated the establishment of the vascular connection between the shoot and root.     

The Micronutrient Boron Causes the Development of Adventitious Roots in Sunflower Cuttings

Three-day-old light-grown sunflower seedlings were de-rootedand incubated in nutrient solutions that either contained orlacked boric acid (B). In the absence of B, in the majorityof the seedlings, no adventitious roots were formed. The micronutrientB caused the development of numerous roots in the lower partof the hypocotyl. The effect of B occurred without the supplyof any phytohormones. A dose-response curve of B-induced rootingyielded an optimum concentration of 0.1 mM boric acid. Histologicalstudies revealed that cell divisions occurred in the controlbut no root primordia developed. In cuttings that were incubatedin B (0.1 m M) root primordia were observed that rapidly developedinto well-differentiated adventitious roots. Sunflower cuttingsthat were planted with their cut end in vermiculite that wasmoistened with nutrient solutions without B degenerated afterseveral weeks. In the presence of B the cuttings formed numerousadventitious roots that entirely replaced the tap root systemof intact seedlings. The rooted cuttings developed into sturdyadult sunflower plants. Our results are discussed with respectto the possible role of B in the evolution of vascular fromprevascular plants.Copyright 1999 Annals of Botany Company Adventitious roots, boron, cuttings, organogenesis, sunflower seedlings.     

Root-to-shoot primordium conversion on Sesbania rostrata Brem. stem explants

The morphogenetic responses of cultured stem explants of Sesbaniarostrata Brem. from various positions along the stem axis wereanalysed after treatment with four growth regulators (BAP, NAA,kinetin, and GAJ. Internodal explants formed adventitious shootbuds when cultured on a Murashige and Skoog basal medium withoutadded growth regulators. Histological studies of regenerated shoot buds revealed thatapproximately 30% of the buds resulted from the conversion ofa preformed root primordium (characteristic of this species)into a shoot bud without a callogenesis phase. Each bud whichoriginated from a single root primordium grew into a leafy shoot.Preformed root primordia of stem explants of Sesbania rostratamay constitute an excellent model for physiological researchon plant differentiation. Key words: Organogenesis, adventitious bud, preformed root primordium, conversion, Sesbania rostrata     

Mechanisms of primordium formation during adventitious root development from walnut cotyledon explants

 In walnut (Juglans regia L.), an otherwise difficult-to-root species, explants of cotyledons have been shown to generate complete roots in the absence of exogenous growth regulators. In the present study, this process of root formation was shown to follow a pattern of adventitious, rather than primary or lateral, ontogeny: (i) the arrangement of vascular bundles in the region of root formation was of the petiole type; (ii) a typical root primordium was formed at the side of the procambium within a meristematic ring of actively dividing cells located around each vascular bundle; (iii) the developing root apical meristem was connected in a lateral way with the vascular bundle of the petiole. This adventitious root formation occurred in three main stages of cell division, primordium formation and organization of apical meristem. These stages were characterized by expression of LATERAL ROOT PRIMORDIUM-1 and CHALCONE SYNTHASE genes, which were found to be sequentially expressed during the formation of the primordium. Activation of genes related to root cell differentiation started at the early stage of primordium formation prior to organization of the root apical meristem. The systematic development of adventitious root primordia at a precise site gave indications on the positional and biochemical cues that are necessary for adventitious root formation. Received: 30 July 1999 / Accepted: 16 February 2000     

Influence of Zeatin on Flowering in Root Forming Cuttings of Anagallis arvensis L.

Applied zeatin affected flowering in cuttings of Anagallis arvensis(long-day plant). According to the zeatin concentration andthe root formation stage, zeatin inhibited flowering at differentdegrees or slightly promoted it. Thus, the root primordium settingseemed to be correlated with this effect of zeatin. (Received February 14, 1984; Accepted May 18, 1984)