STUDIES ON THE GERMINATION OF SEEDS OF THE ROOT PARASITES,ALECTRA VOGELII AND STRIGA GESNERIOIDES. I. ANATOMICAL CHANGES IN THE EMBRYOS

Anatomical changes in the radicle and shoot meristems of embryos of germinating seeds of the obligate root parasites, Alectra vogelii and Striga gesnerioides were studied. When germination of seeds was stimulated by cowpea (Vigna unguiculata) root exudate, growth occurred mainly in the radicular pole of embryos and minimally in the plumular pole, resulting in seedlings with elongated radicles. Maximum radicle elongations of about 3 mm in A. vogelii and 2 mm in S. gesnerioides were recorded during a period of 8 and 11 days, respectively. Analysis of the radicular tip during the course of seed germination revealed that the activity of the meristematic tissue progressively decreased until it completely disappeared. When germinated seeds were cultured on nutrient agar media, the radicle meristem of A. vogelii continued to grow producing a normal root with a root cap. On the other hand, the radicles of cultured S. gesnerioides seeds elongated only slightly before meristematic activity ceased. During continued growth of seedlings of both species on agar media, lateral roots whose tips had typical angiosperm root topography, were initiated from the radicle.     

TEMPERATURE-INDUCED CAVITIES AND SPECIALIZED PARENCHYMA CELLS IN THE VASCULAR CYLINDER OF PEA ROOTS

Pea seeds (Pisum sativum L.) of six cultivars were planted in the field, in the greenhouse, or in growth chambers, in five different media, in light or dark, and at various temperatures (10–32 C). Under all conditions above 15 C the central portion of the vascular cylinder, in all cultivars except “Ageotropum,” tended to form cavities in almost every primary root examined. These cavities then became filled by the ingrowth of specialized parenchyma cells (SP cells). The formation of cavities and SP cells was temperature dependent since the roots grown below 15 C always formed central metaxylem tracheary elements (MTEs), without cavities and SP cells. Cavities and SP cells did not form over the entire root length. When the roots were longer than 3 cm, they started to form cavities and SP cells and continued for an additional 10–30 cm. After that, late MTEs formed in the central vascular cylinder, and no cavities and SP cells occurred regardless of temperature. Within an individual root grown above 15 C, cavities and SP cells tended to form during periods of fast growth, while during periods of slow growth large central MTEs formed instead.     

THE ULTRASTRUCTURE OF THE QUIESCENT CENTER IN THE APEX OF CULTURED ROOTS OF CONVOLVULUS ARVENSIS L.

Cultured roots of the common bindweed, Convolvulus arvensis L. growing at the rate of 15–30 mm/day in sterile nutrient medium were fixed for electron microscopic analysis. The ultrastructure of the quiescent center, the initials of the ground meristem, and the initials of the procambium were studied in order to determine whether sequential structural changes could be correlated with models for specifying the mechanisms by which cell differentiation and cell division might be controlled. The differentiation of cells in the root proper occurs very gradually in linear files from the site of the quiescent center proximally into the different tissue regions. Major structural changes, such as the orientation and subsequent elongation of cells along the longitudinal axis of the root and cell wall changes, indicate that the control of differentiation and perhaps cell division occurs in radial gradients outwardly from the quiescent center.     

THE QUIESCENT CENTER IN CULTURED ROOTS OF CONVOLVULUS ARVENSIS L.

Cultured roots of Convolvulus arvensis were incubated in 0.2–0.3 μc/ml methyl-³H-thymidine for different intervals of time. In roots supplied with tritiated thymidine for 12 hr, 14 hr, 48 hr, or 14 hr followed by transfer to fresh medium for 24 hr, autoradiographs prepared of serial, longitudinal sections of the root tips showed the presence of a subterminal quiescent center in the root proper at the distal poles of the central cylinder and cortex. In addition, a zone of unlabelled cells in the columella, distal to the root cap initials, was present. In roots supplied continuously with tritiated thymidine for 64 hr, 96 hr, and 120 hr, the quiescent center was either reduced in size or was not present.     

Characteristics of adventitious root formation in cotyledon segments of mango (Mangifera indica L. cv. Zihua): two induction patterns, histological origins and the relationship with polar auxin transport

Cotyledon segments derived from zygote embryos of mango (Mangifera indica L. cv. Zihua) were cultured on agar medium for 28 days. Depending on different pre-treatments with plant growth regulators, two distinct patterns of adventitious roots were observed. A first pattern of adventitious roots was seen at the proximal cut surface, whereas no roots were formed on the opposite, distal cut surface. The rooting ability depended on the segment length and was significantly promoted by pre-treatment of embryos with indol-3-acetic acid (IAA) or indole-3-butyric acid (IBA) for 1 h. A pre-treatment with the auxin transport inhibitor 2,3,5-triiodobenzoic acid (TIBA) completely inhibited adventitious root formation on proximal cut surfaces. A second pattern of roots was observed on abaxial surfaces of cotyledon segments when embryos were pre-treated with 2,700 μM 1-naphthalenacetic acid (NAA) for 1 h. Histological observations indicated that both patterns of adventitious roots originated from parenchymal cells, but developmental directions of the root primordia were different. A polar auxin transport assay was used to demonstrate transport of [³H] indole-3-acetic acid (IAA) in cotyledon segments from the distal to the proximal cut surface. In conclusion, we suggest that polar auxin transport plays a role in adventitious root formation at the proximal cut surface, whereas NAA levels (influx by diffusion; carrier mediated efflux) seem to control development of adventitious roots on the abaxial surface of cotyledon segments.     

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 DEVELOPMENTAL ANATOMY OF OPUNTIA BASILARIS. I. EMBRYO,ROOT, AND TRANSITION ZONE

The large seeds of Opuntia basilaris Engelm. & Bigel. show an unusually high percentage of germination, followed by a rapid development of the seedling during the first 30 days of growth. The primary root has six xylem arms alternating with six phloem poles around a large central pith. Development of metaxylem opposite each of the primary phloem poles results in the formation of eight collateral bundles. Secondary and tertiary roots have four xylem and phloem poles with xylem developing to the center of the stele. The transition zone is characterized by a gradual disappearance of all but two of the primary xylem arms of the root. Metaxylem development in the central portion of the transition zone interconnects the protoxylem poles forming a primary xylem cylinder around the central pith. The collateral bundles pass through the transition zone essentially without change.     

MORPHOLOGY OF HUMULUS LUPULUS. II. SECONDARY GROWTH IN THE ROOT AND SEEDLING VASCULARIZATION

Miller , Robert H. (U. Nevada, Reno.) Morphology of Humulus luppulus. II. Secondary growth in the root and seedling vascularization. Amer. Jour. Bot. 46(4): 269–277. Illus. 1959.—In the primary state the roots of Humulus lupulus L. have a diarch xylem plate with 2 strands of primary phloem lying on either side of the primary xylem. Secondary histogenesis is described for the primary root. Fibrous and fleshy storage roots are developed by the hop plant and their respective developmental and anatomical structures are described. Lateral roots are initiated in the pericycle opposite the protoxylem poles. The architecture of these secondary roots is similar to that of the primary root. The seedling develops a fleshy storage organ through secondary growth of the primary root and the hypocotyl. The hypocotyl eventually resembles a fleshy taproot throughout most of its extent. The vascular cambium differentiates large amounts of parenchymatous tissues. A relatively smaller amount of tracheary tissue is formed. The secondary phloem comprises a high percentage of phloem parenchyma and ray cells containing numerous large starch grains, and constitutes the larger portion of the fleshy storage root. Numerous thick-walled lignified fibers occur throughout the secondary vascular tissues. Resin and tannin cells are abundantly distributed. A phellogen is differentiated from the pericycle and develops a persistent periderm on the outer surface of the fleshy storage organ. A relatively short transition region occurs in the upper part of the hypocotyl. The transition takes place from a radially alternate arrangement of the vascular tissues in the root to a collateral arrangement in the cotyledons.     

ADVENTITIOUS ROOT DEVELOPMENT IN TYPHA GLAUCA,WITH EMPHASIS ON THE CORTEX

Adventitious root development was investigated in Typha glauca plants grown under experimental conditions with the previous year's dead, sterile stalk either emerged above or submerged below the surface of Hoagland's solution. Adventitious roots emerged from buds in which most primordia had been earlier formed. Most roots elongated to 14–19 cm in 3–4 weeks and produced abundant lateral roots to their tips. Root apical meristem organization was typically monocotyledonous with a single tier of ground meristem/protoderm over the procambium. The ground meristem had zones of periclinal divisions in its innermost and outermost layers; the innermost layer initiated the endodermis and midcortex, and the outermost layers initiated the hypodermis. Crystalliferous cells with raphides were produced in the midcortex, and aerenchyma resulted from the radial expansion of schizogenous air spaces and some lysigeny in the midcortex. The procambium produced a vascular cylinder with 10–13 phloem and xylem poles, 6–9 large metaxylem elements, and central sclerenchyma. As roots stopped elongating, they narrowed, the vascular cylinder diminished in size, typical aerenchyma was lost from the cortex, crystal production ceased, and the rootcap diminished in size with its storage starch used up. Growth was determinate in these adventitious roots. The results suggested that a periclinally derived outer ground meristem was a prerequisite for a hypodermis, which, in turn, was necessary as a structural framework for aerenchyma. Without a hypodermis, typical aerenchyma was not present.     

ON THE CORRELATION OF PRIMARY ROOT LENGTH,MERISTEM SIZE AND PROTOXYLEM TRACHEARY ELEMENT POSITION IN PEA SEEDLINGS

Pea seedlings (Pisum sativum L. cv. Alaska) were darkgrown in vermiculite. Roots of various lengths were cleared, stained and measured to determine the relative meristem height (MH), width and volume and the distance to the most proximal protoxylem tracheary element (PTE). A correlation was found between root length, MH and PTE position as follows: in roots from 4–40 mm as the root elongated the MH lengthened and PTE position increased its distance from the body/cap juncture; in roots 40–80 mm MH and PTE position remained approximately constant; in longer roots (80–120 mm) MH became shorter, and PTE position closer to the tip as the root elongated. The relationship, using our measurement procedure was that for every 0.19 mm change in MH, the PTE position changed by 1 mm. This response was partially growth rate dependent since short roots (4–80 mm) grew at a constant rate and longer roots (80–140 mm) grew slower. Root manipulations and trifluralin treatment, to inhibit cell division, caused tip swelling and modulated the position of the PTE toward the root tip. The control of the spatial relationship between meristem size and maturation position is discussed.     

DEVELOPMENTAL MORPHOLOGY OF THE EMBRYO AND SEEDLING OF RHIZOPHORA MANGLE L. (RHIZOPHORACEAE)

The embryo of Rhizophora mangle L. is initially attached to the integument by a long multiseriate suspensor. Its basal cells lyse, and intrusive growth of the endosperm envelops the embryo, forces the micropyle open, and often carries the embryo out of the integument. Thus, “germination” is effected by growth of the endosperm rather than of the embryo. The surface of the endosperm differentiates into a layer of peculiar transfer cells. The cotyledonary body initiates as a toroidal primordium, which later becomes lobed; most of the free portions ultimately fuse. After “germination,” the axis of the viviparous seedling grows by a diffuse intercalary meristem below the cotyledonary node. Before seedling abscission, the shoot apex produces three pairs of leaves, the first of which aborts, leaving the rest of the plumule protected by their stipules. The (immersed) radicle apex is nearly inactive, but lateral roots arise early in seedling development; these are usually the first or only roots to grow during establishment. Ten provascular strands “differentiate” in the cotyledons; a hollow provascular cylinder develops in the hypocotyl. Initial vascular differentiation in the latter is of many alternate poles of xylem and phloem; later, de novo differentiation of metaxylem opposite the protophloem poles, and vice versa, produces collateral bundles. Xylem maturation is endarch over most of the length of the hypocotyl, but tangential and random series of metaxylem vessels occur in the radicle end.     

Does salinity reduce growth in maize root epidermal cells by inhibiting their capacity for cell wall acidification?

The reduction in growth of maize (Zea mays L.) seedling primary roots induced by salinization of the nutrient medium with 100 millimolar NaCl was accompanied by reductions in the length of the root tip elongation zone, the length of fully elongated epidermal cells, and the apparent rate of cell production: Each was partially restored when calcium levels in the salinized growth medium were increased from 0.5 to 10.0 millimolar. We investigated the possibility that the inhibition of elongation growth by salinity might be associated with an inhibition of cell wall acidification, such as that which occurs when root growth is inhibited by IAA. A qualitative assay of root surface acidification, using bromocresol purple pH indicator in agar, showed that salinized roots, with and without extra calcium, produced a zone of surface acidification which was similar to that produced by control roots. The zone of acidification began 1 to 2 millimeters behind the tip and coincided with the zone of cell elongation. The remainder of the root alkalinized its surface. Kinetics of surface acidification were assayed quantitatively by placing a flat tipped pH electrode in contact with the elongation zone. The pH at the epidermal surfaces of roots grown either with 100 millimolar NaCl (growth inhibitory), or with 10 millimolar calcium ± NaCl (little growth inhibition), declined from 6.0 to 5.1 over 30 minutes. We conclude that NaCl did not inhibit growth by reducing the capacity of epidermal cells to acidify their walls.     

夏侧金盏花幼苗初生维管系统的解剖学研究

夏侧金盏花(Adonis estivalis L.)为毛莨科(Ranuncula-ceae)侧金盏花属(Adonis)植物。其幼苗可明显地分为上胚轴苗区、子叶节区和下胚轴根区。幼苗以子叶节区为中心,往上其子叶节区中部和上部为子叶节—茎过渡区;向下其下胚轴为子叶节—根过渡区。目前对毛莨科某些属幼苗初生维管系统的个体发育研究已有一些报道。但夏侧金盏花幼苗的子叶节—茎过渡区的转变与已报道的其他属均不同。主要表现为其子叶节区下部的中始式二原型双肩状单中柱,到达子叶节区中部,其后生木质部弦向发育成二唇形外韧维管束雏型,在中柱中央出现薄壁组织,进一步发育则形成髓;再往上,即子叶节区上部,便一分为多个内始式的外韧维管束雏型,直接形成上胚轴(茎)的真中柱。此研究为再一次验证子叶节区理论的正确性与进一步揭示被子植物初生维管系统的演化规律积累一份新资料。     

Reliable plantlet formation from embryos and seedling shoot tips of radiata pine

A method has been devised for the reliable production of plantlets from embryos and seedling shoot tips of Pinus radiata D.Don (radiata pine). Buds were induced on an agar or liquid Schenk and Hildebrandt (SH) medium containing 5.0 mg/l benzylaminopurine (BAP). Except for some abnormal buds, the buds grew into elongated shoots on an agar SH medium without cytokinin. The transfer of shoots from a SH medium to a Gresshoff and Doy (GD) medium was found to be an important pretreatment which increased the survival of the shoots when they were placed in a peat and pumice mix for root formation. Elongated shoots were induced to form roots under non-sterile conditions in a humid environment with occasional misting. An intervening 5-day treatment of shoots in an agar medium containing 2.0 mg/l indolebutyric acid (IBA) and 0.5 mg/l napthaleneacetic acid (NAA) significantly increased the percentage of shoots forming roots and the number of roots formed per shoot over control shoots placed directly in the peat:pumice mix. An enhanced level of CO₂ during root formation had no effect on the time of root formation or on the percentage of shoots forming roots. These results concerning the elongation, growth and rooting of adventitious shoots are now being applied to the development of very large numbers of plantlets starting from cotyledons from partially germinated seeds.     

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.     

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 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.     

Effect of heavy metals and temperature on the oxalate oxidase activity and lignification of metaxylem vessels in barley roots

The effect of Cd on oxalate oxidase (OxO) activity and its localisation were analysed in barley root. In Cd-treated roots OxO activity was strongly induced in the region 2–4 mm behind the root tip and in the area toward the root base. In situ analyses showed that Cd-induced OxO activity was localised to the cell wall (CW) of early metaxylem vascular bundles and surrounding parenchyma cells and was accompanied by lignification of metaxylem vessels. OxO activation was also observed during treatment with other heavy metals (HMs), salt treatment and at elevated non-optimal temperature. In contrast to HM activation of OxO and lignification, high temperature and NaCl indeed activated OxO but did not induce lignification of metaxylem vessels. These results suggest that oxalate oxidase as an H₂O₂-generating enzyme is activated in response to several stresses, however the ectopic lignification of metaxylem vessels is activated specifically by HMs. This HM-induced premature root xylogenesis due to ectopic lignification of metaxylem vessels probably causes shortening of the root elongation zone and therefore a reduction in root growth.     

Mechanisms of Reduction of the Stelar Pattern along Barley Roots

The stelar pattern along the seminal and nodal roots of barley (Hordeum vulgare L.) is gradually simplified due to a decreasing frequency of longitudinal cell division in the apical meristem. The decrease involves the proportion of stelar parenchyma, the number of vascular strands on the periphery of the stele and, in nodal roots with a more complex structure, the number of central metaxylem files. In spite of the fact that the stelar parenchyma is reduced in distal parts of the roots to approximately one half, the discontinuity of central and peripheral metaxylem is preserved. Reduction of the number of central metaxylem files is due to fusion. In the reduction of peripheral xylem and phloem strands, the development of certain xylem strands is discontinued and they are terminated blindly. Two phloem strands that had alternated radially with them, approach each other, coalesce and a single phloem strand continues to develop. In this way the regular alternation of phloem and xylem is re-established. The importance of fusions ensuring reduction of the functional continuity in vascular tissue by formation of a network structure must be stressed. This reduction mechanism is involved not only in files of the wide central metaxylem but also in phloem strands which are thus preferred over blindly terminating peripheral xylem strands.     

THE ROOT APICAL MERISTEM OF ASPLENIUM BULBIFERUM: STRUCTURE AND DEVELOPMENT

The root apical meristem of Asplenium bulbiferum Forst. f. has a prominent four-sided pyramidal cell with its base in contact with the rootcap. Derivatives (merophytes) that contribute to the main body of the root are produced from the three proximal faces of the apical cell. The rootcap has its origin from the fourth (distal) face of the apical cell. The first division in a proximal merophyte is periclinal to the root surface, separating an outer cell and an inner cell. The outer cell is the origin of the outer part of the cortex and the epidermis; the larger inner cell is the origin of the inner cortex, endodermis, pericycle, and vascular tissue. After the establishment of the basic number of cells in a unilayered merophyte, the cells undergo transverse divisions forming longitudinal files of cells. The mitotic index of the apical cell indicates that it is not a quiescent cell. Also, the first plane of division in a newly formed merophyte dictates that the apical cell is the originator of merophytes.