The Behaviour of Two Cell Populations in the Pericycle of Allium cepa, Pisum sativum, and Daucus carota During Early Lateral Root Development

The length of cells of the pericycle, endodermis and middlecortex not actively involved in lateral root primordia (LRP)development was measured in primary roots of Allium cepa, Pisumsativum and Daucus carota. The presence of two cell populationsin the pericycle was demonstrated in all three species. In Alliumcepa and Pisum sativum, pericyclic cells located opposite xylempoles were significantly shorter than cells lying opposite phloempoles. In both species, LRP originated opposite xylem poles.Our results, furthermore, strongly suggest that in regions ofthe root far from the apical meristem, numerous pericyclic cellsundergo transverse division both previous to and during LRPinitiation, decreasing in mean length throughout this period.In Daucus carota, LRP begin to form in pericyclic cells locatednext to the phloem poles, such cells were significantly shorterthan those opposite xylem poles, even in areas of the primaryroot located close to the root tip. Cells also appear to dividetransversely in regions far from the root tip in this species,leading to a conspicuous drop in the mean length of those cellslocated in portions of the pericycle destined to give rise toLRP. Two different cell populations can also be distinguishedin the endodermis of Allium cepa and Pisum sativum, althoughobservations were less conclusive in Daucus carota. In all threespecies, length of cortical cells was unaffected by their positionopposite xylem or phloem poles Allium cepa, carrot, cell division, cell length, Daucus carota, endodermis, lateral root development, onion, pea, pericycle, Pisum sativum     

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     

Pericycle proliferation pattern during the lateral root initiation in adventitious roots ofAllium cepa

Summary Far from the apical meristem of adventitious roots ofAllium cepa, the pericycle shows great proliferative activity related to lateral root initiation. A group of mother pericycle cells undergoes asymmetrical transverse and periclinal divisions following a well-established pattern. Successive asymmetrical transverse divisions, progressing from one end of the cell to the other, divide the original mother cell into very short derivatives. Later, these short derivative cells undergo periclinal divisions. This proliferative activity starts nearly simultaneously in two elongated and highly vacuolated pericycle cells located in the same column in front of one of the xylem poles. Then proliferation expands centrifugally towards other pericycle cells in the same and adjacent columns. The proliferative activity of the pericycle cells decreases progressively outwards. Only the most central of these cells produce derivatives which contribute to the future lateral root.     

Changes in cell length and mitotic index in vascular pattern-related pericycle cell types along the apical meristem and elongation zone of the onion root

Summary Three pericycle cell types (opposite xylem, opposite phloem and intervening) distinguished by their location in relation to different elements of the vascular system were studied in the adventitious root ofAllium cepa L. Changes in cell length and mitotic index were analysed in these cells along the apical meristem and elongation zone of the root. The opposite phloem and intervening pericycle cells are significantly shorter than the opposite xylem pericycle cells in the apical half of the meristem. Between 1,200 and 1,400 m behind the tip, length became similar in all three pericycle cell types, while in more proximal zones the opposite phloem cells were significantly longer. These results suggest that the number of transverse divisions is different in the three types of pericycle cells. In the apical half of the meristem, mitotic index increased in intervening and opposite xylem cells but remained unchanged in opposite phloem cells, a fact likely to account for the relative lengthening of the latter. In the proximal half of the meristem, mitotic index fell in all three cell types until cell division had ceased. However, mitotic index in opposite xylem cells remained high for longer than in the other two cell types, implying that increase of the mean cell length in the former was slower. These results suggest that differences in mean cell length between the three pericycle cell types are due to different rates of proliferation.     

Histological analysis of adventitious rooting in Arabidopsis thaliana (L.) Heynh seedlings

ABSTRACT

Adventititous rooting is essential for the post-embryonic growth of the root apparatus in various species. In Arabidopsis thaliana, adventitious rooting has been reported in some mutants, and auxin seems to be the inducer of the process. The objective of the study was to identify the tissues involved in adventitious rooting in the most commonly used ecotypes for molecular and genetic studies (i.e. Columbia, Wassilewskija and Landsberg erecta) both in the presence and absence of exogenous auxin. Seedlings of the three ecotypes were grown under various conditions. When grown under 16 hours light/day for 11 days, all seedlings showed adventitious roots, both with and without auxin, however, both adventitious and lateral rooting were enhanced by exogenous auxin (2 µM naphthaleneacetic acid). Independently of the presence of auxin and of the ecotype, the hypocotyl pericycle produced adventitious roots directly (i.e., according to the same pattern of lateral root formation by the pericycle cells in the primary root). However, in the presence of auxin, roots of indirect origin also, and mainly, formed and their formation was preceded by the exfoliation of the tissues external to the stele. Exfoliation was caused by cell hypertrophy, separation, and disintegration, which mainly involved the endodermis. At the exfoliation site, the pericycle, with a minor contribution of a few endodermal cells, produced the callus from which indirect roots arose. The finding that adventitious rooting occurs in the absence of auxin (all ecotypes) indicates that this process is part of the normal root apparatus in Arabidopsis, with the hypocotyl pericycle as the target tissue of the process. Exogenous auxin alters adventitious rhizogenesis mainly affecting the endodermis response.     

Origin of Epidermis and Development of Root Primordia in Pistia, Hydrocharis and Eichhornia

All three floating plants have roots bearing laterals derivedfrom both pericycle and endodermis. In Pistia and Eichhornialaterals arise within the meristem of the mother root; in Hydrocharisthey arise from mature tissue. In Pistia and Hydrocharis theepidermis becomes anatomically discrete between cortex and cap:in Pistia it is derived from the endodermis of the mother root,in Hydrocharis from the pericycle. The epidermis is not discretein Eichhornia and is derived from the pericycle of the motherroot with the cortex. Stathmokinetic data were used to construct timetables of developmentwhich show how the differences arise. In Pistia the first periclinaldivision of the endodermis-derived tissue individualizes theepidermis and occurs early, before a quiescent centre forms.In Hydrocharis the epidermis also becomes discrete before thepole of the meristem becomes quiescent, but it does so by apericlinal division of the pericycle-derived tissue. In Eichhorniapericlinal divisions occur in the outermost layer of the pericycle-derivedtissue long after quiescence has set in at the pole and afterthe fourth periclinal division in the endodermis derived cap.Its epidermis therefore never becomes anatomically discretethough it becomes functionally discrete because its polar cellsstop dividing as in the other plants. The involvement of the endodermis of mother roots in the formationof laterals is discussed in relation to the state of differentiationat sites of primordium formation, discreteness of the epidermisand subsequent fate of primordia. Pistia stratiotes L., Hydrocharis morsus-ranae L., Eichhornia crassipes Solms., primordia, lateral root, discrete epidermis, development, chimera, stathmokinetics     

ORIGIN AND DEVELOPMENT OF LATERAL ROOTS IN TYPHA GLAUCA

Lateral roots of Typha glauca arose from the pericycle of the parent adventitious root. Periclinal divisions of the pericycle gave rise to two layers; the outermost initially produced the ground meristem and protoderm, and the innermost produced the procambium. The immature endodermis of the parent root contributed to the early stages of the root tip as an endodermal covering. Prior to emergence, the ground meristem/protoderm produced cells into the endodermal covering. After emergence, the endodermal covering was replaced by a calyptrogen, which was derived from the ground meristem/protoderm and which, in turn, formed the rootcap. A typical monocotyledonous three-tiered meristem was then produced. An outer ground meristem also arose before emergence to form a hypodermis in many lateral roots; in these, crystalliferous cell production began in midcortex cells before emergence, and a small aerenchyma developed in their cortices. The rootcap columella stored small amounts of starch shortly after emergence. Lateral roots of T. glauca were smaller than their parental adventitious roots; they normally had only two to six poles of xylem and phloem, and the cortex was less than six cells across. During 1–3-cm elongation, the lateral root apical meristem and mature regions narrowed, stored starch disappeared, fewer crystals formed, aerenchyma production ceased, and the roots stopped elongation.     

Cell Wall Thickening of the Xylem Tracheary Elements in Different Zones of the Adventitious Root of Allium cepa L.

Cell wall thickness of the xylem tracheary elements was measuredin the proto- and metaxylem of the Allium cepa L. adventitiousroot. Measurements were taken in root fragments of known age(1, 3, 5, 7 and 9 d) located in either the basal or medio-apicalzone. Tracheary elements in the protoxylem matured within ashorter period of time than those in the metaxylem. Final cellwall thickness was greater in metaxylem than in protoxylem components.The cell wall thickening in the tracheary elements in both proto-and metaxylem was more rapid in the basal zone of the root thanin the medio-apical zone. Additionally, cell walls of the maturetracheary elements were thicker in the basal zone than in areasfurther from the bulb. Allium cepa, onion, root, cell wall, xylem maturation     

Occurrence of cell surface arabinogalactan-protein and extensin epitopes in relation to pericycle and vascular tissue development in the root apex of four species

Monoclonal antibodies recognizing two classes of developmentally regulated plant cell surface components – arabinogalactan-proteins (AGPs) and extensins – have been used to immunolabel cells at the root apices of four species with different characteristics of pericycle and vascular tissue development. Root apices of pea (Pisum sativum L.), radish (Raphanus sativus L.), carrot (Daucus carota L.) and onion (Allium cepa L.) were immunolabelled with the anti-AGP monoclonal antibodies JIM4 and JIM13 and anti-extensin monoclonal antibodies JIM11, JIM12, JIM19 and JIM20. All of these antibodies recognized subsets of pericycle cells in at least one, but never all, of these species. The restricted patterns of epitope occurrence also reflected vascular cell development. The differences in patterns of antibody recognition in the four species are discussed in relation to the possible roles of these cell surface molecules in cell differentiation and root patterning events. Received: 11 March 1997 / Accepted: 20 May 1997     

Lateral root initiation by asymmetrical transverse divisions of pericycle cells in adventitious roots ofAllium cepa

Summary In onion adventitious roots cellular events have been identified that indicate that lateral root initiation occurs earlier and nearer the apex than previously documented. Lateral roots are not initiated when a pericycle cell divides periclinally but earlier, when a pair of neighbouring pericycle cells in the same column divide transversely and asymmetrically, with both mitoses close to the end towards the neighbouring pericycle cell. Each cell therefore produces two cells of unequal length. The shorter cells produced by the mother pericycle cells are adjacent, while the longer cells are located above and below the shorter cells. This objective morphological criterion allows clear identification of the site of lateral root initiation. Subsequent to these asymmetric divisions, both the longer pericycle cells again divide transversely and asymmetrically producing more short cells adjacent to the previous ones. The first periclinal division occurs in one of these short pericycle cells.     

Morphogenesis in Yucca schidigera Roezl. Root Organ Cultures

Root development in suspension cultures of Yucca schidigerawas light-mediated. The green cultures consisted of roots, smalltissue aggregates and suspension cells. Roots possessed an apicalmeristem with a root cap, meristematic region and region ofdifferentiating tissues. Phloem, xylem vessels and tracheidsoccurred in discrete polyarch vascular bundles. Xylary wallthickening was reticulate, and endodermis and pericycle werepresent. Roots of intact Y. schidigera plants had a similardistribution of vascular tissues. Dark-grown cultures were cream-colouredand contained only lobed tissue aggregates and suspension cells. Yucca schidigera Roezl., tissue cultures, morphogenesis, root organ, light/dark     

The Anatomy of Contractile Roots in Eucomis L'H?rit

The anatomical changes in the different tissues during rootcontraction were studied in two species of Eucomis (Liliaceae).It is evident that the shortening of the root is brought aboutby radial and tangential broadening and longitudinal shorteningof the perivascular cortical cells. This tissue is hence calledcontractile parenchyma. As a result of this contraction thecell walls of the exodermis, endodermis, pericycle, phloem andpith become buckled longitudinally while the annular and spiralthickenings of the xylem are pressed together.     

Anatomy and Histochemistry of the Root System of the Kiwifruit Vine, Actinidia deliciosa var. deliciosa.

The kiwifruit vine is a species which has been newly introducedinto cultivation and little is known of its comparative physiologyand anatomy. In this study we found that fibrous, 'magnolioid'roots, which have undergone secondary vascular development butwhich retain the cortex and develop a suberized epidermis, comprisethe greater part of the root system (95% of total length). Newlyinitiated roots with primary development conform to norms establishedin other woody plant species. However, the structural roots,like the fibrous roots, also retain a cortex and phellodermwhich is initiated by hypodermal cells within the cortex andnot by the pericycle which is the common progenitor tissue inother species. This phellogen produces new cells centrifugallyonly. The cortex is a relatively small component of the structuralroot and the bulk of the tissue is vascular in origin, as inthe roots of other plant species. The endodermis is retainedand continues to divide periclinally to accommodate the increasein circumference with growth.Copyright 1993, 1999 Academic Press Actinidia deliciosa, root anatomy, ontogony, histochemistry, exodermis, endodermis     

DEVELOPMENTAL ANATOMY IN ROOTS OF IPOMOEA PURPUREA. II. INITIATION AND DEVELOPMENT OF SECONDARY ROOTS

In seedlings of Ipomoea purpurea secondary roots are initiated in the primary root pericycle opposite immature protoxylem. Cells derived from immature endodermis, pericycle, and incipient protoxylem and stelar parenchyma contribute to the primordium. The derivatives of the endodermis become a uniseriate covering over the tip and flanks of the primordium and emerged secondary root; the endodermal covering is sloughed off when the lateral root reaches 1–5 mm in length. A series of periclinal and anticlinal divisions in the pericycle and its derivatives gives rise to the main body of the secondary root. The initials for the vascular cylinder, cortex, and rootcap-epidermis complex are established very early during primordium enlargement. After emergence from the primary root, the cortical initials undergo significant structural modifications related to enlargement of the ground meristem and cortex, and the rootcapepidermal initials are partitioned into columellar initials and lateral rootcapepidermal initials. Procambium diameter increases by periclinal divisions in peripheral sectors. The mature vascular cylinder is comprised of several vascular patterns, ranging from diarch to pentarch, that are probably related ontogenetically. Cells derived from incipient protoxylem and stelar parenchyma cells of the primary root form the vascuar connection between primary and secondary roots.     

Morphological changes and transversal growth kinetics along the apical meristem in the pericycle cell types of the onion adventitious root

Summary In the apical meristem of the adventitious root ofAllium cepa, all pericycle cells show a marked increase in cross-sectional area between 400 and 800 m behind the tip, this transversal growth ceasing in the 1,200–1,400 m interval. However, different pericycle cell types (opposite xylem, intervening and opposite phloem) show different transversal growth kinetics. Along the meristem, the opposite xylem cells are narrower than both the intervening and opposite phloem cells, and these latter are similar in cross-sectional area. Another relevant difference is in the polarity of the transversal expansion, which in turn gives rise to changes in cell shape. In fact, in apical most portions of the meristem, the opposite phloem cells mainly expand tangentially, while the intervening cells do so radially, and the opposite xylem cells undergo a similar tangential and radial expansion. By contrast, in basal most portions of the meristem, radial expansion continues in the opposite phloem cells when it has ceased in the intervening cells. These latter expand tangentially once again when tangential expansion has ceased in the opposite phloem cells. As a consequence of this transversal growth, the opposite xylem cells, which can initiate lateral root primordia, retain their isodiametric transversal shape along the meristem, whereas the transversal shape of the opposite phloem and intervening cells initially changes from isodiametric to markedly enlarged tangentially (opposite phloem) or radially (intervening), after which both cell types tend to become more rounded in shape.     

Chloramphenicol effects on adventitious root production by radish hypocotyls

Abstract

The excision of the root accelerates greatly the formation of adventitious roots in the hypocotyl of etiolated radish seedlings, but if the seedlings develop in CAP 1×10^?4M, no adventitious root are induced after cutting. IAA either alone or associated with CAP, significantly increases the number of primordia in normal hypocotyls if given at the moment of cutting, while it has not stimulatory effect on the hypocotyls of seedlings grown in CAP. IAA has significant effect on both elongation and tickening of hypocotyl segments prepared from seedlings grown in CAP, and this could indicate a specific action of the inhibitor either on a particular process or on particular cells.

The endodermis and the pericycle, which are the two cell layers implicated in the formation of the adventitious roots, could be the mediators of this particular effect of CAP in rooting.     

Calcium Movement, Graviresponsiveness and the Structure of Columella Cells and Columella Tissues in Roots of Allium cepa L.

Roots of Allium cepa L. cv. Yellow are differentially responsiveto gravity. Long (e.g. 40 mm) roots are strongly graviresponsive,while short (e.g. 4 mm) roots are minimally responsive to gravity.Although columella cells of graviresponsive roots are largerthan those of nongraviresponsive roots, they partition theirvolumes to cellular organelles similarly. The movement of amyloplastsand nuclei in columella cells of horizontally-oriented rootscorrelates positively with the onset of gravicurvature. Furthermore,there is no significant difference in the rates of organellarredistribution when graviresponsive and nongraviresponsive rootsare oriented horizontally. The more pronounced graviresponsivenessof longer roots correlates positively with (1) their caps being9.6 times more voluminous, (2) their columella tissues being42 times more voluminous, (3) their caps having 15 times morecolumella cells, and (4) their columella tissues having relativevolumes 4·4 times larger than those of shorter, nongraviresponsiveroots. Graviresponsive roots that are oriented horizontallyare characterized by a strongly polar movement of ⁴⁵Ca²⁺ acrossthe root tip from the upper to the lower side, while similarlyoriented nongraviresponsive roots exhibit only a minimal polartransport of ⁴⁵Ca²⁺. These results indicate that the differentialgraviresponsiveness of roots of A. cepa is probably not dueto either (1) ultrastructural differences in their columellacells, or (2) differences in the rates of organellar redistributionwhen roots are oriented horizontally. Rather, these resultsindicate that graviresponsiveness may require an extensive columellatissue, which, in turn, may be necessary for polar movementof ⁴⁵Ca²⁺ across the root tip. Allium cepa, onion, root, columella tissue, columella cell, gravitropism, calcium, ultrastructure     

ADVENTITIOUS ROOT DEVELOPMENT FROM THE SEEDLING HYPOCOTYL OF LYCOPERSICON ESCULENTUM

Tomato seedlings five through ten days old were used for this investigation. Adventitious roots were initiated from the pericycle of the tomato hypocotyl. The position of adventitious root development was irregular in the rhizogenic hypocotyl; however, the cellular pattern of individual root development was very regular. Four layers of pericycle derivatives participated in root histogenesis and a bi- or triseriate endodermal cover was derived from the endodermis. Fluorescent microscopy showed that Casparian strips on the meristematic endodermal cell walls were not removed biochemically but were displaced around the root primordium by anticlinal divisions and cell enlargement. Casparian strips were not synthesized by endodermal cover cells. The emergent root had a typical three tiered or closed pattern of apical organization, and quiescent centers were present in all emergent roots longer than 0.5–0.6 cm.