Sucrose Utilization via Invertase and Sucrose Synthase with Respect to Accumulation of Cellulose and Callose Synthesis in Wheat Roots under Oxygen Deficiency

The sucrose cleavage by sucrose synthase (SuSy) and neutral invertase was studied in wheat roots (Triticum aestivum L.) subjected to hypoxia or anoxia for 4 days. By in situ activity staining, increased SuSy activity was observed in the tip region and stele of root axes while the activity of invertase decreased. Cellulose content significantly increased in hypoxically treated roots. The cellulose deposition was correlated with regions of high SuSy activity, being mainly located in the pericycle and endodermis. Invertase activity was distributed along the root without clear difference between cortex and stele. Under root hypoxia, a significant increase in the structural carbohydrates, callose and especially cellulose, was shown. Increasing levels of soluble carbohydrates were partially used to synthesize cellulose for secondary wall thickening and callose to counteract the tissue injury following low-oxygen stress. Under strict anoxia, the roots were much more injured but sustained a high level of cellulose and callose while the soluble carbohydrates almost disappeared.     

Aluminum stress in the roots of naked barley

Phytotoxicity of aluminum (Al) is the major limiting factor for the crops grown in acid soils rapidly inhibiting root elongation. In this study, changes in root growth, total activity and isozyme patterns of antioxidant enzymes such as peroxidase, ascorbate peroxidase, catalase and glutathione reductase by Al stress were investigated in the roots of naked barley (Hordeum vulgare L. cv. Kwangwhalssalbori). As Al concentration increased up to 500 M, the rooting rate and root elongation substantially decreased. Growth results suggested that this cultivar is an Al-sensitive species. Total activities of antioxidant enzymes generally increased at lower Al concentrations and then gradually decreased at higher Al concentrations. They also increased when the exposure time to Al was extended up to 48 hr. Changes in the isozyme patterns of antioxidant enzymes were investigated byin situ enzyme activity staining on a non-denaturing PAGE gel. They generally coincided with the changes in the total activity in parallel. Changes in the total activity of antioxidant enzymes also coincided with the changes of the root growth. Since growth reduction in the roots by Al stress could be related with the changes in the activities of antioxidant enzymes, these results suggested that Al might cause the oxidative stress in the roots of this cultivar of naked barley.     

The importance of anatomical structural study of roots for the understanding of physiological processes

Abstract

Besides being species‐specific, the inner structure of the root is influenced by the place and time of origin during the growth period. From the root tip up to the base of a particular root, the zones of cell division, cell elongation, formation of root hairs and root branching can be distinguished. The root tip that is covered by a root cap and mucilage is protected against evaporation and water contact. From the end of the lateral parts of the root cap, the cells become exposed to the surrounding environment. The cells can elongate by water uptake or can shrink by water loss. All processes of geotropic growth take place there. In this study, some differences are illustrated using Zea mays plants. Radicle and roots emerging from several nodes of the shoot as well as lateral roots are compared. The distances from the tip and from the base of the root are also very important for characterization of particular root functions. Distinctive features such as root diameter, size of the stele and of the cortex, ratio of cortex to stele, number and width of the xylem vessels, size of cells, special thickenings and stage of lignification as well as symptoms of maturation are observed.     

A comprehensive analysis of root morphological changes and nitrogen allocation in maize in response to low nitrogen stress

The plasticity of root architecture is crucial for plants to acclimate to unfavourable environments including low nitrogen (LN) stress. How maize roots coordinate the growth of axile roots and lateral roots (LRs), as well as longitudinal and radial cell behaviours in response to LN stress, remains unclear. Maize plants were cultivated hydroponically under control (4 mm nitrate) and LN (40 μm ) conditions. Temporal and spatial samples were taken to analyse changes in the morphology, anatomical structure and carbon/nitrogen (C/N) ratio in the axile root and LRs. LN stress increased axile root elongation, reduced the number of crown roots and decreased LR density and length. LN stress extended cell elongation zones and increased the mature cell length in the roots. LN stress reduced the cell diameter and total area of vessels and increased the amount of aerenchyma, but the number of cell layers in the crown root cortex was unchanged. The C/N ratio was higher in the axile roots than in the LRs. Maize roots acclimate to LN stress by optimizing the anatomical structure and N allocation. As a result, axile root elongation is favoured to efficiently find available N in the soil.     

Oxygen distribution and movement,respiration and nutrient loading in banana roots (Musa spp. L.) subjected to aerated and oxygen-depleted environments

Excessive soil wetness is a common feature where bananas (Musa spp.) evolved. Under O₂ deficiency, a property of wet soils, root growth and functions will be influenced by the respiratory demand for O₂ in root tissues, the transport of O₂ from the shoot to root and the supply of O₂ from the medium. In laboratory experiments with nodal roots of banana, we examined how these features influenced the longitudinal and radial distributions of O₂ within roots, radial O₂ loss, solute accumulation in the xylem, root hydraulic conductivity, root elongation and root tip survival. In aerated roots, the stele respired about 6 times faster than the cortex on a volume basis. Respiratory O₂ consumption decreased substantially with distance from the root apex and at 300–500 mm it was 80% lower than at the apex. Respiration of lateral roots constituted a sink for O₂ supplied via aerenchyma, and reduced O₂ flow towards the tip of the supporting root. Stelar anoxia could be induced either by lowering the O₂ partial pressure in the bathing medium from 21 to 4 kPa (excised roots) or, in the case of intact roots, by reducing the O₂ concentration around the shoot. The root hair zone sometimes extended to 1.0 mm from the root surface and contributed up to a 60% drop in O₂ concentration from a free-flowing aerated solution to the root surface. There was a steep decline in O₂ concentration across the epidermal-hypodermal cylinder and some evidence of a decline in the O₂ permeability of the epidermal-hypodermal cylinder with increasing distance from the root apex. The differences in O₂ concentration between cortex and stele were smaller than reported for maize and possibly indicated a substantial transfer rate of dissolved O₂ from cortex to stele in banana, mediated by a convective water flow component. An O₂ partial pressure of 4 kPa in the medium reduced net nutrient transfer into the vascular tissue in the stele within 1 or 2 h. Hypoxia also caused a temporary decrease in radial root hydraulic conductivity by an order of magnitude. In O₂ deficient environments, the stele would be among the first tissues to suffer anoxia and O₂ consumption within the root hair zone might be a major contributor to root anoxia/hypoxia in banana growing in temporarily flooded soils.     

白鲜根的发育解剖学研究

应用半薄切片、常规石蜡切片并结合离析法,对药用植物白鲜(Dictamnus dasycarpus Turcz.)根的发生发育过程进行了研究。结果表明:白鲜根的发生发育过程包括4个阶段,即原分生组织阶段、初生分生组织阶段、初生结构阶段以及次生结构阶段。原分生组织位于根冠内侧及初生分生组织之间,衍生细胞分化为初生分生组织。初生分生组织由原表皮、基本分生组织以及中柱原组成。原表皮分化为表皮,基本分生组织分化为皮层,中柱原分化为维管柱,共同组成根的初生结构;在初生结构中,部分表皮细胞外壁向外延伸形成根毛,皮层中分布有油细胞,内皮层有凯氏带,初生木质部为二原型或偶见三原型,外始式;根初生结构有髓或无。次生结构来源于原形成层起源的维管形成层的活动以及中柱鞘起源的木栓形成层的活动;白鲜次生韧皮部宽广,其中多年生根中可占根横切面积的85%,另外除基本组成分子外,还分布有油细胞;周皮发达,木栓层厚;初生皮层、次生木质部和次生韧皮部薄壁细胞中常充满丰富的淀粉粒。     

Regulation of Growth Anisotropy in Well-Watered and Water-Stressed Maize Roots (I. Spatial Distribution of Longitudinal,Radial, and Tangential Expansion Rates)

As a system to study the regulation of growth anisotropy, we studied thinning of the primary root of maize (Zea mays L.) occurring developmentally or induced by water stress. Seedlings were transplanted into vermiculite at a water potential of approximately -0.03 MPa (well-watered) or -1.6 MPa (water-stressed). The diameter of roots in both treatments decreased with time after transplanting; the water-stressed roots became substantially thinner than well-watered roots at steady state, showing that root thinning is a genuine response to water stress. To analyze the thinning responses we quantified cell numbers and the spatial profiles of longitudinal, radial, and tangential expansion rates separately for the cortex and stele. The results showed that there was no zone of isotropic expansion and the degree of anisotropy varied greatly with position and treatment. Thinning over time in well-watered roots was caused by rates of radial and tangential expansion being too low to maintain the shape of the root. In response to low water potential, cell number in both tissues was unchanged radially but increased tangentially, which shows that thinning was caused wholly by reduced cell expansion. Water stress substantially decreased rates of tangential and radial expansion in both the stele and cortex, but only in the apical 5 mm of the root; basal to this, rates were similar in well-watered and water-stressed roots. By contrast, as in previous studies, longitudinal expansion was identical between the treatments in the apical 3 mm but in water-stressed roots was inhibited at more basal locations. The results show that expansion in longitudinal and radial directions can be regulated independently.     

Transport of [3H]IAA label in gravistimulated primary roots of maize

The curvature of roots in response to gravity is attributed to the development of a differential concentration gradient of IAA in the top and bottom of the elongation region of roots. The development of the IAA gradient has been attributed to the redistribution of IAA from the stele to cortical tissues in the elongation region. The gravistimulated redistribution of IAA was investigated by applying [³H]IAA to the cut surface of 5 mm apical primary root segments. The movement of label from the stele-associated [³H]IAA into the root, tip, root cap, and cortical tissues on the top and bottom of the elongation region was determined in vertically growing roots and gravistimulated roots. Label from the stele moved into the region of cell differentiation (root tip) prior to accumulating in the elongation region. Little label was observed in the root cap. Gravistimulation did not increase the amount of label moving from the stele; but gravistimulation did increase the amount of label accumulating in cortical tissues on the lower side of the elongation region, and decreased the amount of label accumulating in cortical tissues on the upper side of the elongation region. Removal of the cap prior to or immediately following gravity stimulation rendered the roots partially insensitive to gravity and also prevented gravity-induced asymmetric redistribution of label. However, removal of the root cap following 30 min of gravistimulation did not alter root curvature or the establishment of an IAA asymmetry across the region of root elongation. These results suggest that a signal originating in the root cap directs auxin redistribution in tissues behind the root cap, leading to the development of an asymmetry of IAA concentration in the elongation region that in turn causes the differential growth rate in the elongation region of a graviresponding root.     

Transport of [3H]IAA label in gravistimulated primary roots of maize

The curvature of roots in response to gravity is attributed to the development of a differential concentration gradient of IAA in the top and bottom of the elongation region of roots. The development of the IAA gradient has been attributed to the redistribution of IAA from the stele to cortical tissues in the elongation region. The gravistimulated redistribution of IAA was investigated by applying [³H]IAA to the cut surface of 5 mm apical primary root segments. The movement of label from the stele-associated [³H]IAA into the root, tip, root cap, and cortical tissues on the top and bottom of the elongation region was determined in vertically growing roots and gravistimulated roots. Label from the stele moved into the region of cell differentiation (root tip) prior to accumulating in the elongation region. Little label was observed in the root cap. Gravistimulation did not increase the amount of label moving from the stele; but gravistimulation did increase the amount of label accumulating in cortical tissues on the lower side of the elongation region, and decreased the amount of label accumulating in cortical tissues on the upper side of the elongation region. Removal of the cap prior to or immediately following gravity stimulation rendered the roots partially insensitive to gravity and also prevented gravity-induced asymmetric redistribution of label. However, removal of the root cap following 30 min of gravistimulation did not alter root curvature or the establishment of an IAA asymmetry across the region of root elongation. These results suggest that a signal originating in the root cap directs auxin redistribution in tissues behind the root cap, leading to the development of an asymmetry of IAA concentration in the elongation region that in turn causes the differential growth rate in the elongation region of a graviresponding root.     

Comparative Analysis of Endogenous Hormones in Leaves and Roots of Two Contrasting Malus Species in Response to Hypoxia Stress

Plant hormones play important roles in regulating developmental processes and signaling networks involved in plant responses to biotic and abiotic stresses. We comparatively studied the growth and endogenous hormonal levels in leaves and roots in two Malus species (M. sieversii and M. hupehensis) differing in hypoxia tolerance under normoxic and hypoxia stress. The results showed that hypoxia stress inhibited growth of seedlings of both Malus species, but with significant differences in intensity. Exposure to hypoxia altered the levels of endogenous hormones in leaves and roots in both Malus seedlings. Leaf and root abscisic acid (ABA) contents increased in response to hypoxia stress in both genotypes despite different extents. Compared with M. hupehensis, M. sieversii was more responsive to hypoxia stress, resulting in larger increases in leaf and root ABA contents. The changes in leaf and root ABA contents correlating with the different tolerance levels of the genotypes confirm the involvement of this hormone in plant responses to hypoxia stress. Gibberellins (GAs; GA₁ + GA₄) continuously increased in leaves and roots during the whole period of stress, whereas indole-3-acetic acid (IAA) showed a sharp increase at the early stage in both Malus seedlings. In addition, zeatin riboside (ZR), dihydrozeatin riboside (DHZR), and isopentenyl adenine (IPA) differed in their pattern of changes in both Malus seedlings under hypoxia stress. Based on variations in endogenous hormonal levels in both Malus species that differ in their ability to tolerate hypoxia, we conclude that not a single hormone but multiple hormones and their interplay are responsible for hypoxia tolerance.     

Cell Expansion during the Elongation of Lateral Roots of Vicia faba L.

Cell width, breadth, length, cross-sectional area, volume andthe ratio of the volume of the nucleus to that of the cell havebeen determined in the epidermis, cortex and stele over theapical mm of lateral roots of Vicia faba as they elongated fromjust-emerged to 4 cm in length. Cell volume increased basallyalong the root in each tissue, but this was not a result ofcell expansion taking place in all dimensions in the epidermis,cortex and stele. Thus, while increase in cell length, was themajor factor involved in cell volume increase basally alongthe root in the stele, the corresponding dimensions in the epidermisand cortex were cell width and cell breadth respectively. Cellvolume was greater in the cortex than in the other tissues andusually greater in the epidermis than in the stele. Cell lengthwas greater in the stele than in the other tissues, while cellbreadth was maximal in the cortex and cell width in the epidermis.Changes took place in the various measurements made as the rootselongated. These are discussed with respect to the onset oflateral root growth and to changes in the rate of growth ofthese roots as they elongate.     

Protective Effect of External Ca2+ on Elongation and the Intracellular Concentration of K+ in Intact Mung Bean Roots under High NaCl Stress

The effects of Ca²⁺ in the external medium on intact mung beanroots under high NaCl stress were investigated. With increasingexternal concentrations of NaCl, mung bean roots showed suppressionof elongation and a decrease in the intracellular concentrationof K⁺. Addition of Ca²⁺ to the external medium alleviated theinhibition of root elongation under the high NaCl stress andmaintained a high intracellular concentration of K⁺ in the elongatingregion of the roots. This counter effect of Ca²⁺ against theNaCl stress on roots was correlated with the ratio of [Ca²⁺]/[Na⁺]²in the external medium. A value above 5.0 ? 10^–4 mM^–1resulted in almost complete recovery of root elongation undervarious high concentrations of NaCl. Root elongation for 24h under NaCl stress was correlated with the extent to whichthe intracellular concentration of K⁺ was in excess of 10 mM.Maintenance of an adequate concentration of K⁺ in root cellsis essential for root elongation under salt stress. These findingsindicate that Ca²⁺ prevents the leakage of intracellular K⁺and thereby supports the elongation of roots under salt stress. (Received November 13, 1989; Accepted June 5, 1990)     

The Effect of Coumarin on Root Growth and Root Histology

The effect of coumarin on the root growth was studied on roots from intact plants, isolated roots and isolated elongating zones. All material was cultivated aseptically. A new method was developed for sterile culture of intact plants in flowing nutrient medium. The effects on cell division and cell elongation were studied separately. An effect on both these processes can be established at all concentrations that affect the root growth. The concentration-growth curve has an “all-or-none” appearance. Coumarin inhibits the transverse divisions in all cell layers; the perivascular layers seem to be more sensitive. Also the mitotic activity that is involved in the initiation of laterals is inhibited. The longitudinal divisions within the stele are enhanced. Coumarin decreases the cell length in all cell layers, most likely with greater relative sensitivity in the perivascular layers. Studies on the time course of cell elongation in both attached corn roots and isolated elongating zones reveal that the decrease in cell length is caused exclusively by a decrease in the maximal rate of elongation, whereas the duration of the elongation is unchanged. With each decrease of the cell length, the cell diameter is increased. The two changes are intimately connected within the greater part of the active region of concentration. Studies on the time course of the radial expansion in isolated elongating zones show a strict connection in time between cell elongation and radial expansion. The radial expansion leads to unchanged or increased cell volume at most concentrations and for most cell types. Coumarin causes an inhibition of the longitudinally directed processes and a stimulation of the radially directed ones. This is interpreted as indicating that the formative system is disengaged or reorientated, i.e., the polarity of the cells is changed. Through experiments partly with isolated elongating zones and partly by disruption of the linear phase by means of mannitol, the inhibitory effect of coumarin could be localized to the first non-linear phase of the elongation. The results were compared with earlier findings in the literature. The microtubuli are proposed as a conceivable main Component in the formative system common to both cell division and cell elongation. These are assumed to be affected by changes in the SH/SS balance produced by coumarin.     

Abscisic Acid Response of Corn (<Emphasis Type="Italic">Zea mays</Emphasis> L.) Roots and Protoplasts to Lanthanum

Lanthanum ions antagonize calcium and are used as a Ca²⁺ channel blocker but their direct effects are unknown. We investigated lanthanum effects on endogenous abscisic acid (ABA) levels in protoplasts and intact primary roots of Zea mays L. Application of 1 mM La³⁺ reduced primary root elongation, caused swelling of root tips, and essentially doubled the ABA content in intact roots but decreased ABA in root protoplasts in a concentration-dependent manner. Osmotic stress increased ABA level in protoplasts more than in intact roots. Temporal ABA changes in response to La³⁺ treatment indicate that La³⁺ affects root growth at least partially via ABA pathway.     

Variations in maturation strains and root shape in root systems of Maritime pine (Pinus pinaster Ait.)

 In order to determine if different types of wood were being laid down in the root system of Maritime pine (Pinus pinaster Ait), in response to wind loading, longitudinal residual maturation strains (LRMS), indicating the existence of mechanical stress in developing wood cells, were measured in the trunk and lateral roots. Two age groups of trees (5- and 13-year- old) were compared. LRMS were greater in the trunk and roots of 13-year-old trees than in 5-year-old trees. This phenomenon may be due to increased competition between older trees. LRMS in leeward roots of both age-groups were positive i.e. the wood cells had developed under compression, as also occurs in reaction wood of gymnosperms. As leeward roots are placed under compression during tree sway, an abnormal type of wood may form in the roots in order to counteract the increased stress. In other roots, the strains were negative i.e. the cells had developed under tension, as occurs in normal wood. In the roots of younger trees, LRMS were also positive nearer the stem, thus indicating that wood formation may also be influenced by bending stresses experienced in this zone. In addition to LRMS measurements, radial growth in roots was examined in order to determine the influence of mechanical loading on secondary growth. In older trees, there was a significant increase of 34% in woody growth below the biological centre, compared to that above. This eccentricity is unlike that found in most other tree species, where secondary growth is usually greater on the upper side of the root. However, Maritime pine has a tap root, which will alter the pattern of stress within the root system. Under wind loading, a concentration of mechanical stress will develop at the bases of the stem, lateral roots and tap root. Received: 7 July 1997 / Accepted: 11 December 1997     

Drought-induced Short Roots in Arabidopsis thaliana: Structural Characteristics

In Arabidopsis thaliana, as in other Brassicaceae species, a progressive drought stress induced changes in root morphogenesis: from a threshold plant water deficit, the new emerging roots remain short, hairless and often take a tuberized shape at their base while drought persists. The organization of these drought-induced roots was examined in light microscopy in Arabidopsis thaliana, Columbia wild-type ecotype, and compared to the normal, well-watered lateral roots. The main structural traits were the absence of elongation zone, the arrest of cell cap expansion, the lack of root hairs (despite epidermal differentiation in trichoblasts and atrichoblasts) and the radial enlargement of epidermal and cortical cells. The early differentiation, close to the short root apex, of large and highly lignified metaxylem elements, the absence of starch accumulation in hypertrophied cortical cells appeared to be characteristic of the species Arabidopsis, as compared to other Brassicaceae. These structural alterations are discussed in terms of drought-induced changes in gene expression with regard to similar modifications described in root morphogenesis and root hair-defective Arabidopsis mutants.     

Effect of Fluoride on Nucleic Acids and Growth in Germinating Corn Seedling Roots

Corn seeds were treated with 0.01 M sodium fluoride for various time periods. The treated seeds were germinated and grown until the seedling roots reached a standard size of 12±3 mm. Analyses were made for RNA and DNA contents of 3-mm seedling root tips. Determinations also were made for growth rate, rate of cell elongation, cell multiplication, and tissue maturity of 12-mm roots. RNA contents of 3-mm root tips were found to be directly proportional to the growth rates of the entire seedling root of corn seeds treated with sodium fluoride for various periods of time. The RNA content was reduced on a cell basis and was independent of the root tip cell number. The amount of DNA was not related to the growth rate of the intact seedling roots. Since fluoride reduced the number of mitotic figures, it was likely that fluoride inhibited DNA synthesis during the interphase of the mitotic cycle. Growth by cell multiplication was inhibited more than that by cell elongation in the sample treated with fluoride for a shorter period. The two types of growth, however, showed a similar level of growth reduction in the sample treated with fluoride for a longer period. Fluoride seemed to reduce the rates of cellular elongation and multiplication not more than about 40 per cent of the control value in these tissues under present experimental conditions. Fluoride also induced maturity in the seedling roots in proportion to the periods of fluoride treatment.