Primary Roots Of Zea Mays Research Articles

Evaluation of Metaxylem Vessel Histogenesis and the Occurrence of Vessel Collapse during Early Development in Primary Roots of Zea mays ssp. mexicana: A Result of Premature Programmed Cell Death?

Root apical meristem histological organization in Zea mays has been carefully studied previously. Classical histology describes its system as having a “closed organization” and a development of xylem that conforms to predictable rules. Among the first cell types to begin differentiation are late-maturing metaxylem (LMX) vessels. As part of a larger study comparing domestic maize root development to a wild subspecies of Z. mays (teosinte), we encountered a metaxylem development abnormality in a small percentage of our specimens that begged further study, as it interrupted normal maturation of LMX. Primary root tips of young seedlings of Zea mays ssp. mexicana were fixed, embedded in appropriate resins, and sectioned for light and transmission electron microscopy. Longitudinal and serial transverse sections were analyzed using computer imaging to determine the position and timing of key xylem developmental events. We observed a severe abnormality of LMX development among 3.5% of the 227 mexicana seedlings we screened. All LMX vessel elements in these abnormal roots collapsed and probably became non-functional shortly after differentiation began. Cytoplasm and nucleoplasm in the abnormal LMX elements became condensed and subdivided into irregularly-shaped “macrovesicles” as their cell walls collapsed inward. We propose that these seedlings possibly suffered from a mutation that affected the timing of the programmed cell death (PCD) that is required to produce functional xylem vessels, such that autolysis of the cytoplasm was prematurely executed, i.e., prior to the development and lignification of secondary walls.

Growth is required for perception of water availability to pattern root branches in plants

Water availability is a potent regulator of plant development and induces root branching through a process termed hydropatterning. Hydropatterning enables roots to position lateral branches toward regions of high water availability, such as wet soil or agar media, while preventing their emergence where water is less available, such as in air. The mechanism by which roots perceive the spatial distribution of water during hydropatterning is unknown. Using primary roots of Zea mays (maize) we reveal that developmental competence for hydropatterning is limited to the growth zone of the root tip. Past work has shown that growth generates gradients in water potential across an organ when asymmetries exist in the distribution of available water. Using mathematical modeling, we predict that substantial growth-sustained water potential gradients are also generated in the hydropatterning competent zone and that such biophysical cues inform the patterning of lateral roots. Using diverse chemical and environmental treatments we experimentally demonstrate that growth is necessary for normal hydropatterning of lateral roots. Transcriptomic characterization of the local response of tissues to a moist surface or air revealed extensive regulation of signaling and physiological pathways, some of which we show are growth-dependent. Our work supports a "sense-by-growth" mechanism governing hydropatterning, by which water availability cues are rendered interpretable through growth-sustained water movement.

Abscisic Acid Response of Corn (Zea mays 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.

Spatial separation of light perception and growth response in maize root phototropism.

Although the effects of gravity on root growth are well known and interactions between light and gravity have been reported, details of root phototropic responses are less documented. We used high-resolution image analysis to study phototropism in primary roots of Zea mays L. Similar to the location of perception in gravitropism, the perception of light was localized in the root cap. Phototropic curvature away from the light, on the other hand, developed in the central elongation zone, more basal than the site of initiation of gravitropic curvature. The phototropic curvature saturated at approximately 10 micromoles m-2 s-1 blue light with a peak curvature of 29 +/- 4 degrees, in part due to induction of positive gravitropism following displacement of the root tip from vertical during negative phototropism. However, at higher fluence rates, development of phototropic curvature is arrested even if gravitropism is avoided by maintaining the root cap vertically using a rotating feedback system. Thus continuous illumination can cause adaptation in the signalling pathway of the phototropic response in roots.

Cellular Dimorphism in the Maize Root Cortex: Involvement of Microtubules, Ethylene and Gibberellin in the Differentiation of Cellular Behaviour in Postmitotic Growth Zones

AbstractDetailed morphometric analysis of cell shapes and an immunofluorescent study of microtubules were carried out on primary roots of Zea mays L. Two types of cells were found to be formed within the postmitotic isodiametric growth (PIG) region of the root cortex that were differentially responsive to low level of exogenous ethylene. The innermost and central cell rows of the cortex were sensitive to ethylene treatment and showed a disturbed distribution of cortical microtubules (CMTs) as well as changed polarity of cell growth, whereas the 2–3 outermost cell rows were less sensitive in this respect. This suggests that post‐mitotic cells of the inner cortex are specific targets for ethylene action. These properties of the inner cortex are compatible with its cells being involved in the formation of aerenchyma; they may also favour root growth in compacted soil. By contrast, the specific properties of the outer cortex indicate that this tissue domain is necessary for the gaseous impermeability and the mechanical strengthening of subjacent aerenchymatous cortex, especially in the mature region of the root.Ethylene affected neither the pattern of cortical cell expansion in the meristem nor the position of the PIG region with respect to the root tip. This contrasts with gibberellin‐deficiency which affected these parameters in both parts of the cortex. These observations indicate a fundamental difference between the role of these two phytohormones in the morphogenesis and development of maize roots.

Gravitropism of Primary Roots of Zea mays L. at Different Displacement Angles

Primary roots of Zea mays were oriented at various angles from the vertical ranging from 99° to 1° and their subsequent bending analysed from filmed records. The maximum rate of bending and the time before bending commenced both varied two-fold, but showed no correlation with the initial angle of tip displacement. Roots orientated to small initial angles (< 40°) often overshot the vertical and proceeded to oscillate around this orientation, whereas roots oriented to large initial angles (> 60°) often failed to achieve the vertical. Roots in this latter group resumed bending after an indeterminate time, or did so immediately after a second displacement of their tip, showing that they were not intrinsically unable to bend. The apparently spontaneous resumption of bending after a temporary plagiogravitropic phase is suggested as being due to noise in the graviperception system in the root cap. The tips of roots growing vertically downwards showed oscillatory bending movements up to 10° either side of vertical. This angle corresponds to the minimum angle of displacement which induces gravitropic bending. Only when roots were oriented 10-20° from vertical did they begin unequivocally to show a gravitropism since at such angles the deflection of their tips exceeded that due to their natural oscillation.

Distribution of IAA and ABA in gravistimulated primary roots ofZea mays. L.

The transport of14C-IAA and14C-ABA applied exogenously to root cap toward the elongation zone was investigated in gravi- and light-stimulated primary roots ofZea mays L. cv. Golden Cross Bantam 70. No significant difference of either IAA or ABA in radioactivities was observed between upper and lower halves of elongation zones during the latent period (0–60 min after the stimulation) of gravitropic response. When quantitative analysis of endogenous IAA and ABA by an internal standard method was carried out 60 min after gravi- and/or light-stimulation, no asymmetric redistribution of either IAA or ABA was observed between upper and lower halves of elongation zones. Light irradiation increased by 20% the contents of ABA in elongation zones. These results suggest that although both IAA and ABA are basipetally transportable and can transmit their information to the elongation zone during a latent period we cannot explain the gravitropic curvature by their redistributions between the two (upper and lower) halves of primary roots ofZea. On the basis of results from the present work and previous papers, the distribution of IAA and ABA in gravistimulatedZea roots is discussed.

Graviresponsiveness of surgically altered primary roots of Zea mays.

We examined the gravitropic responses of surgically altered primary roots of Zea mays to determine the route by which gravitropic inhibitors move from the root tip to the elongating zone. Horizontally oriented roots, from which a 1-mm-wide girdle of epidermis plus 2-10 layers of cortex were removed from the apex of the elongating zone, curve downward. However, curvature occurred only apical to the girdle. Filling the girdle with mucilage-like material transmits curvature beyond the girdle. Vertically oriented roots with a half-girdle' (i.e. the epidermis and 2-10 layers of the cortex removed from half of the circumference of the apex of the elongating zone) curve away from the girdle. Inserting the half-girdle at the base of the elongating zone induces curvature towards the girdle. Filling the half-circumference girdles with mucilage-like material reduced curvature significantly. Stripping the epidermis and outer 2-5 layers of cortex from the terminal 1.5 cm of one side of a primary root induces curvature towards the cut, irrespective of the root's orientation to gravity. This effect is not due to desiccation since treated roots submerged in water also curved towards their cut surface. Coating a root's cut surface with a mucilage-like substance minimizes curvature. These results suggest that the outer cell-layers of the root, especially the epidermis, play an important role in root gravicurvature, and the gravitropic signals emanating from the root tip can move apoplastically through mucilage.

Abscisic acid is not necessary for gravitropism in primary roots of Zea mays.

Primary roots of Zea mays L. cv. Tx 5855 treated with fluridone are strongly graviresponsive, but have undetectable levels of abscisic acid (ABA). Primary roots of the carotenoid-deficient w-3, vp-5, and vp-7 mutants of Z. mays are also graviresponsive despite having undetectable amounts of ABA. Graviresponsive roots of untreated and wild-type seedlings contain 286 to 317 ng ABA g-1 f. wt, respectively. These results indicate that ABA is not necessary for root gravicurvature.

Morphometric Analysis of Epidermal Differentiation in Primary Roots of Zea mays

Epidermal differentiation in primary roots of Zea mays was divided into six cell types based on cellular shape and cytoplasmic appearance. These six cell types are: 1) apical protoderm, located at the tip of the root pole and characterized by periclinally flattened cells; 2) cuboidal protoderm, located approximately 230 microns from the root pole and characterized by cuboidal cells; 3) tabular epidermis, located approximately 450 microns from the root pole and characterized by anticlinally flattened cells; 4) cuboidal epidermis, located approximately 900 microns from the root pole and characterized by cuboidal cells having numerous small vacuoles; 5) vacuolate cuboidal epidermis, located approximately 1,500 microns from the root pole and characterized by cuboidal cells containing several large vacuoles; and 6) columnar epidermis, located approximately 2,200 microns from the root pole (i.e., at the beginning of the zone of elongation) and characterized by elongated cells. We also used stereology to quantify the cellular changes associated with epidermal differentiation. The quiescent center and the apical protoderm have significantly different ultrastructures. The relative volume of dictyosomes increases dramatically during the early stages of epidermal differentiation. This increase correlates inversely with the amount of coverage provided by the root cap and mucilage.

MORPHOMETRIC ANALYSIS OF EPIDERMAL DIFFERENTIATION IN PRIMARY ROOTS OF ZEA MAYS

Epidermal differentiation in primary roots of Zea mays was divided into six cell types based on cellular shape and cytoplasmic appearance. These six cell types are: 1) apical protoderm, located at the tip of the root pole and characterized by periclinally flattened cells; 2) cuboidal protoderm, located approximately 230 μm from the root pole and characterized by cuboidal cells; 3) tabular epidermis, located approximately 450 μm from the root pole and characterized by anticlinally flattened cells; 4) cuboidal epidermis, located approximately 900 μm from the root pole and characterized by cuboidal cells having numerous small vacuoles; 5) vacuolate cuboidal epidermis, located approximately 1,500 μm from the root pole and characterized by cuboidal cells containing several large vacuoles; and 6) columnar epidermis, located approximately 2,200 μm from the root pole (i.e., at the beginning of the zone of elongation) and characterized by elongated cells. We also used stereology to quantify the cellular changes associated with epidermal differentiation. The quiescent center and the apical protoderm have significantly different ultrastructures. The relative volume of dictyosomes increases dramatically during the early stages of epidermal differentiation. This increase correlates inversely with the amount of coverage provided by the root cap and mucilage.

Influence of Electrical Fields and Asymmetric Application of Mucilage on Curvature of Primary Roots of Zea mays

Primary roots of Zea mays cv. Yellow Dent growing in an electric field curve towards the anode. Roots treated with EDTA and growing in electric field do not curve. When root cap mucilage is applied asymmetrically to tips of vertically-oriented roots, the roots curve toward the mucilage. Roots treated with EDTA curve toward the side receiving mucilage and toward blocks containing 10 mM CaCl2, but not toward "empty" agar blocks or the cut surfaces of severed root tips. These results suggest that 1) free calcium (Ca) is necessary for root electrotropism, 2) mucilage contains effector(s) that induce gravitropiclike curvature, and 3) mucilage can replace gravitropic effectors chelated by EDTA. These results are consistent with the hypothesis that the downward movement of gravitropic effectors to the lower sides of tips of horizontally-oriented roots occurs at least partially in the apoplast.

Effects of a Very Low Magnetic Field on the Gravitropic Curvature of &lt;italic&gt;Zea&lt;/italic&gt; Roots

Caryopses with horizontally oriented primary roots of Zea mays L. exposed to light, were incubated in magnetic fields. The gravitropic curvature of roots incubated in a field of 50 μ gauss was larger than the control value, while the control roots grew more rapidly

Effects of abscisic acid and xanthoxin on elongation and gravitropism in primary roots of zea mays

We examined the involvement of abscisic acid (ABA) and xanthoxin (Xan) in maize root gravitropism by (1) testing the ability of ABA to allow positive gravitropism in dark-grown seedlings of the maize cultivar LG11, a cultivar known to require light for positive gravitropism of the primary root, (2) comparing curvature in roots in which half of the cap had been excised and replaced with agar containing either ABA or indole-3-acetic acid (IAA), (3) measuring gravitropism in roots of seedlings submerged in oxygenated solutions of ABA or IAA and (4) testing the effect of Xan on root elongation. Using a variety of methods of applying ABA to the root, we found that ABA did not cause horizontally-oriented primary roots of dark-grown seedlings to become positively gravitropic. Replacing half of the root cap of vertically oriented roots with an agar block containing ABA had little or no effect on curvature relative to that of controls in which the half cap was replaced by a plain agar block. Replacement of the removed half cap with IAA either canceled or reversed the curvature displayed by controls. When light-grown seedlings were submerged in ABA they responded strongly to gravistimulation while those in IAA did not. Xan (up to 0.1 mM) did not affect root elongation. The results indicate that ABA is not a likely mediator of root gravitropism and that the putative ABA precursor, Xan, lacks the appropriate growth-inhibiting properties to serve as a mediator of root gravitropism.

Characterizing pathways by which gravitropic effectors could move from the root cap to the root of primary roots of Zea mays.

Plasmodesmata linking the root cap and root in primary roots Zea mays are restricted to approx. 400 protodermal cells bordering approx. 110000 microns2 of the calyptrogen of the root cap. This area is less than 10% of the cross-sectional area of the root-tip at the cap junction. Therefore, gravitropic effectors moving from the root cap to the root can move symplastically only through a relatively small area in the centre of the root. Decapped roots are non-responsive to gravity. However, decapped roots whose caps are replaced immediately after decapping are strongly graviresponsive. Thus, gravicurvature occurs only when the root cap contacts the root, and symplastic continuity between the cap and root is not required for gravicurvature. Completely removing mucilage from the root tip renders the root non-responsive to gravity. Taken together, these data suggest that gravitropic effectors move apoplastically through mucilage from the cap to the root.

Anatomical disturbances in primary roots of Zea mays following periods of cool temperature

Primary roots of maize ( Zea mays L.) were maintained at 5°C for various periods and then placed at 20°C to allow their regrowth. The growth of both roots and the first leaves in the recovery period decreased in proportion to the period spent in the cold. The anatomy of the portion of root exposed to 5°C and of the subsequently formed root was studied. The cold-treated portion contained aborted lateral root primordia and incompletely developed metaxylem elements. The regenerated portion of root often showed an abnormal vascular system, particularly where the original root had been exposed to the cold conditions for 8 days or more. The abnormalities included the loss of xylem elements and were accompanied by a thinning of the root. In other regenerated roots the vascular patterning was quite irregular and seemed to be triggered by the death, or reduction in diameter, or metaxylem elements in the more proximal, cold-treated portion of root.

The development of structural changes in epidermal cells of Maize Roots During Water Stress

Ultrastructural changes following increasing periods of water stress induced by means of polyethylene glycol 4000 (from 5 min to 18 h) were investigated in young epidermal cells of the primary roots ofZea mays. The sensitivity of the individual cell components to water stress was considered according to the time sequence in which their alterations appeared.Endoplasmic reticulum (ER) and mitochondria proved to be the most sensitive cell components, their structure changing after 5 min of water stress. By 8 h of stress, the condensation of nuclear chromatin in some cells was apparent, preceding polyribosome degradation but not all the other changes of the stressed cells. Similar types of structural alterations appeared slightly earlier in the more differentiated epidermal cells. The rapid changes in the structure of ER and mitochondria coincided with the rapid decrease of water potential of the root tips and the immediate cessation of the root growth.

Collection of gravitropic effectors from mucilage of electrotropically-stimulated roots of Zea mays L.

We placed agar blocks adjacent to tips of electrotropically stimulated primary roots of Zea mays. Blocks placed adjacent to the anode-side of the roots for 3 h induced significant curvature when subsequently placed asymmetrically on tips of vertically-oriented roots. Curvature was always toward the side of the root unto which the agar block was placed. Agar blocks not contacting roots and blocks placed adjacent to the cathode-side of electrotropically stimulated roots did not induce significant curvature when placed asymmetrically on tips of vertically-oriented roots. Atomic absorption spectrophotometry indicated that blocks adjacent to the anode-side of electrotropically-stimulated roots contained significantly more calcium than (1) blocks not contacting roots, and (2) blocks contacting the cathode-side of roots. These results demonstrate the presence of a gradient of endogenous Ca in mucilage of electrotropically-stimulated roots (i.e. roots undergoing gravitropic-like curvature).

The effects of aluminium on root cap function and root development in Zea mays L.

Aluminium concentrations ⩽1.25 mg dm −3 promoted a positive growth response in the primary root of Zea mays and this coincided with an increase in mean cap volume. Cap volume correlated with cap cell numbers. Significant alterations in mean cell volume were limited to the cap periphery where Al caused a decrease in cell volume. This implied that the action of Al was directed at differentiation of peripheral cap cells and that the observed changes in cap volume arose through interference by Al in mechanisms which control cell loss from the cap. This observation was supported by a reduction in the number of detached, mature cap cells surrounding Al-treated roots. Ultrastructural investigation revealed that disruption of Golgi apparatus activity in the peripheral cap cells occurred at the lowest Al concentration (0.5 mg dm −3). A decrease in amyloplast numbers coincided with diminished Golgi apparatus activity in the peripheral cap and these changes preceded reductions in mitotic activity, indicated by decreases in cap volume and root length which were only evident at Al concentrations in excess of 1.25 mg dm −3. Al treatment also resulted in a redistribution of amyloplasts to the proximal halves of central cap cells as well as alterations to the linear arrangement of these cells. The presence of Al was associated with an efflux of H +. Some of the more adverse responses to Al, including inhibition of cell division, were observed to be initiated at Al concentrations where the molar ratio of charge between Ca 2+ and Al 3+ favoured Al. This may attach relevance to the electrical properties of the Al ion in the initiation of an electro-physiological plant response to Al.

Importance of the cap in maize root growth

A large population of primary roots of Zea mays (cv. LG 11) was selected for uniform length at zero time. Their individual growth rates were measured over an 8-h period in the vertical position (in humid air, darkness). Three groups of these roots with significantly different growth rates were then chosen and their cap length was measured. It was found that slowly growing roots had long caps whereas rapidly growing roots had short caps. The production by the cap cells of basipetally transported growth inhibitors was tested (biologically by the curvature of half-decapped roots) and found to be significantly higher for longer root caps than that for shorter ones.