THE ROOT APEX OF MALVA SYLVESTRIS. I. STRUCTURAL DEVELOPMENT

The apical organization of the primary root of Malva sylvestris was analyzed at several growth stages, beginning with the embryo, to determine the structural changes that occurred during growth. Seeds were germinated, and plants were grown under controlled conditions. There were three discrete groups of initials in the embryonic root: those of the central cylinder, cortex, and secondary columella. The secondary columella initials consisted of a plate of cells flanked by a ring of cortical initials. The lateral portion of the rootcap shared a common origin with the epidermis. During growth both the initials of the secondary columella and outer cortex produced rootcap cells. The first indication of the outer cortical initials participating in rootcap formation was observed in roots 3 cm long. In 6-, 9-, and 16-cm roots the cellular continuity between the outer cortex and rootcap was marked, but in 23- and 33-cm roots the histogenic continuity between the outer cortex and rootcap was not evident. In all growth stages the initials of the central cylinder and inner cortex retained their histogenic integrity.     

ONTOGENETIC REORGANIZATION OF THE ROOT MERISTEM IN THE COMPOSITAE

The organization of the root meristem in selected Compositae was investigated to determine whether changes in the pattern of cell arrangement occurred during root growth in species other than Helianthus annuus. Embryonic, short, and long primary roots of one species of each of twelve genera were prepared for microscopic examination. Additional intermediate growth stages were prepared for Echinacea pallida. The meristem of embryonic roots showed layers of initials typical for dicotyledons. The meristem in many of the short roots of eight species was reorganized by the development of a secondary columella. The long roots showed patterns similar to the embryonic roots. In three species which maintained closed meristems, two layers of cortical initials were common in the embryonic root, and as a general trend, a single layer of cortical initials became more common during root elongation. The cellular changes that resulted in the initiation of a secondary columella are characterized by the conversion of cortical initials to secondary columella initials by a shift in their plane of cell division. It is proposed that the size and shape of the quiescent center changes as the conversion takes place. No intermediate stages were observed which could account for the reduction of two layers of cortical initials to one layer.     

THE ROOT APEX OF LINUM

Several developmental stages of primary roots of 13 species of Linum were studied. The basic pattern of root apex organization in three species consisted of a single tier of cortical initials. Ten species had a basic pattern of two tiers of cortical initials. Variations, manifested by either an increase or a decrease in the tiers of cortical initials, were observed in some roots in those species that had a basic pattern of two tiers of cortical initials. Although these variations were interpreted as being ontogenetic, there was no total reorganization of the root apex. There was anatomical and cytological evidence that a quiescent center is established in Linum roots.     

ADVENTITIOUS ROOT DEVELOPMENT FROM THE COLEOPTILAR NODE IN ZEA MAYS L.

Department of Botany and Bacteriology, University of Arkansas, Fayetteville, Arkansas 72701 Zea mays L. root development from the coleoptilar node was observed by light and electron microscopy. Roots developed opposite collateral vascular bundles in the coleoptilar nodal region. Three distinct histogens (stelar, cortical-protoderm, and root cap) became evident in early development. In median sections of the young roots, root cap and cortical regions formed a “hat” configuration over the stelar region. As the root matured, this “hat” developed centripetally to encapsulate the stelar region. Central core cells of the root cap were characterized by having numerous dictyosomes, amyloplasts, vacuoles, and thin cell walls. As these cells matured into outer or peripheral cap cells, the Golgi vesicles became hypertrophied. These hypertrophied vesicles contained a granular PAS-positive material which accumulated between the plasma membrane and the cell wall and formed a thick layer. As the PAS-positive material passed through the cell wall, it changed to a fibrillar texture. A PAS-positive material similar to that in the outer root cap cells was found adjacent to the outer walls of the protodermal cells. In median sections, PAS-positive material was not present in the promeristem region. Root cap cells as well as parent cortical cells were crushed as the young root forced its way through the parent tissue.     

ROOT CONTRACTION IN HYACINTH. I. EFFECTS OF IAA ON DIFFERENTIAL CELL EXPANSION

Contractile roots of Hyacinthus orientalis L. cv ‘Pink Pearl’ shorten as a result of growth of inner cortical cells which expand radially and contract longitudinally. Brief treatment with IAA (indole-3-acetic acid—0.5 and 1.0 mg/1) induces subapical swelling, root cap proliferation and decreased rates of elongation in potentially contractile roots. Growth resumes with removal of IAA from the culture medium and contraction subsequently occurs. The pattern of subsequent contraction is affected by prior IAA treatment; contraction occurs in the normal manner both acropetal and basipetal to the points of IAA-induced swelling, but does not occur in the swollen region itself. Microscopic examination of the swollen region reveals that cells of the middle and outer cortex are radially expanded and longitudinally shortened relative to middle and outer cortical cells of contracted and uncontracted portions of the same root and control roots. In contrast, inner cortical cells in swollen regions of IAA-treated roots show approximately 50% less radial expansion than inner cortical cells of control contracted roots. Middle and outer cortical cells in the swollen region of IAA-treated roots undergo radial expansion, while middle and outer cortical cells in adjacent contracting zones are compressed by radially expanding inner cortical cells. Average volumes of cortical cells in the IAA-induced swollen region increased approximately two-fold when contraction occurred in adjacent regions. These results suggest that in hyacinth roots, under certain circumstances, inner and outer cortical cells alike possess the ability for growth reorientation and expansion. However, during the usual course of contractile root development, cells of the outer cortex are restricted in this ability, through an as yet unknown mechanism, and are passively compressed by the radially expanding inner cortical cells.     

TIME-LAPSE PHOTOGRAPHIC OBSERVATIONS OF MORPHOGENESIS IN ROOT NODULES OF COMPTONIA PEREGRINA (MYRICACEAE)

Seedlings of the sweet fern Comptonia peregrina (L.) Coult. were grown aeroponically with their roots bathed in a nutrient mist lacking nitrogen except for 10 ppm N at the outset. The initiation and early development of root nodules capable of fixing atmospheric nitrogen were recorded with time-lapse photography through early development to the establishment of highly branched, roughly spherical nodules. In Comptonia multiple primary nodule lobes are formed at or near the site of infection with as many as 10 primary lobes occurring together. On the shoulders of the swollen primary lobes new primordia develop, forming secondary nodule lobes, which may persist without nodule root elongation, giving a coralloid appearance. The tips of the lobes may elongate, forming nodule roots which grow vertically upward, or, if disturbed, in random orientation. Nodule roots occasionally form lateral roots. The root axis upon which the nodule forms undergoes secondary thickening on the proximal side of the nodule attachment; the distal portion of the root shows no secondary thickening and later atrophies. Thus, nodules are perennial structures on a woody root system. The endophyte infects and occupies the basal cortical tissues of the primary nodule lobes and successive nodule lobes as they are formed, being restricted to the swollen bases and not infecting the elongate nodule roots. Development of the nodule is interpreted in terms of complex host-endophyte interactions involving the initiation of multiple primordia forming nodule lobes, the active inhibition of nodule lobes and finally nodule root elongation. Anatomical evidence for the endogenous origin of nodule primordium formation substantiates the view obtained from time-lapse photomacrography.     

INITIATION AND DEVELOPMENT OF ROOT NODULES OF CASUARINA (CASUARINACEAE)

Roots of seedlings of the “beefwood” tree, Casuarina cunninghamiana Miq. grown in nitrogen-free nutrient solution were inoculated with a suspension prepared from crashed root nodules taken from mature plants. Marked deformation of root hairs was evident but no infection threads were observed in root hairs. The mode of infection remains undetermined. Root nodules were initiated within three weeks and thereafter numerous upward-growing nodule roots developed from each nodule. Nodules in this symbiotic nitrogen-fixing plant resulted from an infection caused by an unidentified actinomycete-like soil microorganism. Anatomical analysis of nodule formation showed that nodules are the result of repeated endogenous lateral root initiations, one placed upon another in a complexly branched and truncated root system. The endophyte-infected cortical tissues derived from successive root primordia form the swollen nodular mass. Nodule roots develop from nodule lobes after escaping from the initial inhibitory effects of the endophyte. Included is a discussion of the anatomical similarities between nodules of Casuarina which produce nodule roots and those of Alnus which form coralloid nodules usually lacking nodule roots.     

DEVELOPMENTAL ANATOMY IN ROOTS OF IPOMOEA PURPUREA. I. RADICLE AND PRIMARY ROOT

The developmental anatomy of the primary root of Ipomoea purpurea was studied at several growth stages, beginning with the radicle. The radicle is generally composed of three superimposed tiers of initials, which produce the vascular cylinder, cortex, and columella; and a peripheral band of lateral rootcap-epidermal initials. The radicular cortex contains 16–19 immature laticifers; none of the tissue regions in the radicle contains mature cells. Following germination and during the first 2–3 cm growth of the primary root the apical meristem and its derivative tissues undergo a series of modifications. Root apical diameter decreases as cells in lateral portions of the rootcap elongate; meanwhile, the columella enlarges vertically. The relationship between cortical and columellar initials changes as fewer mitoses occur in the former while the latter remain active. In longer roots the columellar initials are directly in contact with the vascular initials. Cortical size diminishes during early root growth as cortical laticifers and their associated cells cease to be produced by the outer cortical initials and ground meristem. Early procambium, at the level of vascular pattern initiation, decreases in diameter by cellular reorientation, and the vascular cylinder decreases in overall diameter although the tetrarch pattern remains unchanged.     

THE COMPARATIVE AND DEVELOPMENTAL MORPHOLOGY OF THE ROOT SYSTEM OF SELAGINELLA SELAGINOIDES (L.) LINK

All of the roots of Selaginella selaginoides are attached laterally to the base of the shoot, which has monopolar growth as is characteristic of Selaginella. The first three roots are produced by meristematic activity in the cortex of the hypocotyl as in several other species of Selaginella. The fourth root is produced in the same way as the first three, except that not all of the cortical cells which become meristematic mature into root tissue. Some of the meristematic tissue remains undifferentiated and continues to produce additional roots. Potentially an unlimited number of roots could be produced, but no plant was found to have more than eight. There is some secondary growth in the cortex of the basal swelling on the hypocotyl, but no secondary vascular tissue is produced and no cambium of any sort is ever organized. On the basis of comparisons with other living species of Selaginella. the centralized root system of S. selaginoides is interpreted as having been modified from a noncentralized type of root system by the persistence of the juvenile mode of root production.     

SEQUENCE AND PATTERN OF LATERAL ROOT FORMATION IN FIVE SELECTED SPECIES

The proximal-distal distribution of the lateral roots of five species was studied. A detailed investigation was carried out on two of the five species, Ceratopteris thalictroides and Cucurbita maxima. A definite pattern of lateral root arrangement, with a degree of variability related to the number of protoxylem poles, was found in all of the species studied. In the fern Ceratopteris, lateral root initiation was found to be related to the segmentation of the apical cell, which in turn determines the distribution of the laterals. In this species the lateral roots occur in a predictable sequence and they are grouped in pairs. In the angiosperms studied, the pattern of lateral root distribution seemed to depend primarily upon a rather strict longitudinal relationship between the lateral root primordia formed opposite any one protoxylem pole. In Cucurbita maxima, 93.7 ± 5.02% of the lateral root primordia observed were in a specific sequence. The laterals of this species are also arranged in groups. In the other plants studied, Arachis hypogaea, Victoria trickeri, and Eichhornia crassipes, the laterals were not as regularly arranged, but nevertheless they were found to be arranged in groups along the main root axis and not randomly dispersed. Factors controlling the spacing of lateral root primordia include their relationship with the developing vascular system, a direct effect of the parent root apex, and an effect of older lateral root primordia in the same sector of the root.     

EFFECTS OF SOIL WATER DISTRIBUTION ON THE LATERAL ROOT DEVELOPMENT OF THREE SPECIES OF CALIFORNIA OAKS

The responses of seedling root systems of three species of oaks in California to two experimental soil moisture regimes were studied by comparing lateral root development, root and shoot weights, and root: shoot ratios. In the first soil moisture treatment the taproot was allowed to extend into moist soil throughout the duration of the experiment (control), while in the second treatment (shallow) the taproot grew into a dry substrate below 30 cm of moist soil. The treatments were intended to approximate soil moisture conditions experienced by oak seedlings in the field when deep soil water sources vary in their accessibility (control: accessible, shallow: inaccessible). Lateral root growth of Quercus agrifolia did not increase significantly when the primary root tip died in the shallow treatment, resulting in an overall decrease in the percent of the root system composed of lateral roots. Q. douglasii and Q. lobata increased lateral root weights by 80% and 70%, respectively, on the upper 30 cm of the primary root when the primary root tip died. Q. lobata was the only species that decreased in shoot and root weight (25% and 21%, respectively) with the loss of the root tip, indicating that, unlike the other species, it was dependent on the primary root for maximum growth. The morphological responses of these species correspond with their distributions and also may be a factor that influences their interactions with other species.     

COMPARATIVE ANATOMY OF ENDOGENOUS BUD AND LATERAL ROOT FORMATION IN CONVOLVULUS ARVENSIS ROOTS CULTURED IN VITRO

The anatomy of endogenous bud and lateral root formation was studied in Convolvulus arvensis roots cultured in vitro. Although early stages in the initiation of buds and roots were found to be identical, continued development of primordia into roots or buds was accompanied by differences in participating primary root tissues, in rates of development, and in orientation of cell divisions. Evidence for the existence of a primordium capable of developing into either a bud or root is discussed with reference to Convolvulus roots and other plant parts shown to have similar morphogenetic potentials.     

小麦种子根的发育解剖

小麦胚胎发育过程中通常形成5条幼根(少数可形成6条),这些根统称为种子根,中间最先发生的为初生根.初生根的原基在胚胎发育的早期就在胚轴的一侧发生,原基细胞由不规则到规则排列。侧生种子根的原基在胚胎发育后期才出现,通常成对发生,并且是由胚轴上的节(盾片节和胚芽鞘节)维管束外方的细胞形成。侧生种子根的发育明显较初生根的快,分化能力也较强,后生木质部导管母细胞出现早,数目较多.因此,小麦胚胎发育过程中从胚轴上形成的这些侧生的种子根,形态上,仍应看作是一些不定根,其结构特征与后来形成须根系的不定根的比较近似。     

THE ROOT APICAL MERISTEM OF OSMUNDA REGALIS

Roots of the Osmundaceae differ from most ferns in having more than one apical cell. The size of the apical initial group, which includes cells that are considered to be apical cells, varies directly with root diameter in Osmunda regalis L. Mitotic indices were 6.63% for apical cells, 7.45% for the entire apical group, 6.25% for distal derivatives, and 7.15% for developing cortical cells. Cytophotometric measurements of Fuelgen-stained nuclei indicated no endopolyploidy in the populations of cells studied. These findings demonstrate that there is no quiescent center in the roots of O. regalis.     

INCREASE IN SIZE AND CELL NUMBER OF LATERAL ROOT PRIMORDIA IN THE PRIMARY OF INTACT PLANTS AND IN EXCISED ROOTS OF PISUM SATIVUM AND VICIA FABA

Increase in cell number, and in anlage volume and length have been investigated during the development of lateral root primordia in roots of intact plants of Pisum sativum and Vicia faba and in excised roots of both species cultured in White's medium supplemented with 2% sucrose. With the exception of primordia in excised roots of Vicia, the general equation which best described increase in each aspect of primoridium growth measured against time was that for exponential growth. When the times necessary for cell number and primordium volume and length to double were determined at intervals over the period of development studied, however, they were found to vary. Similarly, estimates of the size of the proliferative fraction of cells at different times during anlage development indicated that this index of meristematic activity also fluctuated over the developmental period investigated, i.e., increase in cell number and in primordium volume and length do not occur in a truly exponential fashion as the primordia increase in size and cell number. One difference between anlage development in the roots of intact plants and in those grown in culture was that whereas the former primordia completed their development and emerged as lateral roots over the period of the investigation, the latter did not. Moreover, cell doubling time and anlage volume and length doubling times were longer, and the proliferation fraction of cells lower, over the whole period of, and at intervals during, primordium development in the excised roots compared with the results obtained for the roots of the corresponding intact plants.     

THE EFFECTS OF TRITIATED THYMIDINE INCORPORATION ON SECONDARY ROOT PRODUCTION BY PISUM SATIVUM

Hummon , Margaret R. (Montana State U., Missoula.) The effects of tritiated thymidine incorporation on secondary root production by Pisum sativum. Amer. Jour. Bot. 49(10): 1038–1046. Illus. 1962.—Most studies of effects of radiation on plants have involved a general exposure of all the cells and tissues of an organ or entire plant. Tritiated thymidine offers a tool for selective irradiation of the nucleus with little effect on the cytoplasm of a cell. Furthermore, differential incorporation due to variation in the pattern of DNA synthesis permits selective irradiation of cells and tissues. In this study, developing primary roots of Pisum sativum were submerged for brief periods in a solution of tritiated thymidine. This resulted in an alteration of the lateral root pattern. In the area corresponding to the region of elongation during treatment, subsequent lateral root production was suppressed. This correlated with the portion of the root in which there was incorporation of tritiated thymidine into a high percentage of pericycle nuclei. Abnormal development of vascular tissues also occurred, with evidence of altered polarity in the xylem. Although incorporation also occurred in the apical meristem, the latter was not affected at this level of exposure. Thus, differential sensitivity as well as differential incorporation may have been involved in producing the temporary alteration of lateral root production.     

OBSERVATIONS ON THE ROOT ANATOMY OF RICE (ORYZA SATIVA L.)

Rice plants were grown hydroponically and roots were prepared for light and electron microscopy using standard techniques. The roots are bounded by an epidermis, exodermis, and fibrous layer. The exodermis has a suberin lamella along its inner tangential wall. The fibrous layer is composed of thick-walled lignified cells with little pitting. The cortical parenchyma is compact when young, but expands and separates to form a zone of cell walls and air spaces in a spoked arrangement. Supporting columns of living parenchyma cells are occasionally present, particularly near lateral roots. The endodermis is typical for grasses with Casparian strips, suberin lamellae, and tertiary state walls with numerous pits. The pericycle and pith become sclerified. Protoxylem elements alternate with protophloem in the young root; later, early metaxylem, late metaxylem, and metaphloem proliferate. The exodermis, fibrous layer, lacunate cortex, and endodermis appear to present a formidable barrier to radial ion movement in the mature portions of the root.     

SEEDLING GROWTH AND ROOT CONTRACTION IN THE SOAP PLANT,CHLOROGALUM POMERIDIANUM (LILIACEAE)

Radicles and adventitious roots of the soap plant are contractile and through their activity, mature bulbs of this species are buried to depths of 20–30 cm. Experiments were performed to determine rates of contraction and displacement of the shoot apex resulting from activity of the contractile radicle and the first several adventitious roots. The average displacement was 23.2 mm over a 10-wk period and 63.8 mm over 29 weeks. Small glass beads and abortive seeds served as controls and showed no displacement through the soil column. Measurements from longitudinal and transverse sections of contracted and uncontracted portions of radicles revealed average increases of 26–64% in radial dimensions and 40–56% decreases in longitudinal dimensions of inner and middle cortical cells (excluding the endodermis) following contraction. Cells of the outermost cortex (excluding the exodermis) decreased in average longitudinal dimensions by 18–26% before becoming distorted and collapsed as contraction was completed. Average volumes of innermost cortical cells decreased by 15–54%, while two or three cell layers of the middle cortex, adjacent to collapsed outer cortical cells, increased in volume up to 75%. These middle cortical cells are identified as the “active” cells which, by their growth, are responsible for the shortening of the root. Throughout the process of contraction, the stele remains straight and undistorted, although the closer spacing of tracheary element secondary wall thickenings following contraction suggests longitudinal compression of the stele. The average number of cortical cells per transverse section does not differ in contracted and uncontracted roots and no evidence is found to support the “interdigitation” hypothesis of root contraction. However, reorientation of middle cortical cell expansion may be the mechanism of root contraction in Chlorogalum pomeridianum.     

STUDIES ON THE ROOT OF HORDEUM VULGARE L.– ULTRASTRUCTURE OF THE SEMINAL ROOT WITH SPECIAL REFERENCE TO THE PHLOEM

Seminal root tissue of Hordeum vulgare L. var. Barsoy was fixed in glutaraldehyde and osmium tetroxide and studied with the light and electron microscopes. The roots consist of an epidermis, 6–7 layers of cortical cells, a uniseriate endodermis and a central vascular cylinder. Cytologically, the cortical and endodermal cells are similar except for the presence of tubular-like invaginations of the plasmalemma, especially near the plasmodesmata, in the former. The vascular cylinder consists of a uniseriate pericycle surrounding 6–9 phloem strands occurring on alternating radii with an equal number of xylem bundles. The center of the root contains a single, late maturing metaxylem vessel element. Each phloem strand consists of one protophloem sieve element, two companion cells and 1–3 metaphloem sieve elements. The protophloem element and companion cells are contiguous with the pericycle. Metaphloem sieve elements are contiguous with companion cells and are separated from tracheary elements by xylem parenchyma cells. The protoplasts of contiguous cells of the root are joined by various numbers of cytoplasmic connections. With the exception of the pore-plasmodesmata connections between sieve-tube members and parenchymatic elements, the plasmodesmata between various cell types are similar in structure. The distribution of plasmodesmata supports a symplastic pathway for organic solute unloading and transport from the phloem to the cortex. Based on the arrangement of cell types and plasmodesmatal frequencies between various cell types of the root, the major symplastic pathway from sieve elements to cortex appears to be via the companion and xylem parenchyma cells.     

ANATOMY OF MACROZAMIA COMMUNIS LATERAL ROOTS AND ROOT NODULES FORMED IN VITRO STUDIED WITH LIGHT AND SCANNING ELECTRON MICROSCOPY

The anatomy of Macrozamia communis L. Johnson lateral roots and nodules was studied following axenic culture in light and darkness. Pointed lateral roots from dark cultures had an open apical organization similar to that of other cycads and gymnosperms. A distinct protoderm-derived epidermis was not observed. At the apex, the dermis was formed by the outer root capcortical cell layer. Subapically, the outer cortex formed the dermis. No evidence of an algal zone was observed in these roots. The stele was bounded by a distinct endodermis and contained an exarch, diarch xylem. Apogeotropic nodules which developed at the root-shoot junction in darkness, branched dichotomously and had rounded tips covered by tangentially-enlarged root cap cells. The root cap was reduced to a few cell layers and was confined to the extreme nodule apex. The central region of the apical meristem was enlarged, and meristematic cells contained differentiated amyloplasts. A presumptive algal zone was present in some but not all nodules and divided the cortex into inner and outer regions. Stelar anatomy was similar to that observed in pointed, dark-grown lateral roots, except that there was greater xylem differentiation. Nodules which developed in the light were similar to dark-formed nodules, except that root cap cells were radially enlarged and extended over the flanks of the nodule forming a persistent root cap. The heteromorphic lateral roots of M. communis formed a developmental continuum not a heterorhizic root system.