Oxygen Distribution in Wetland Plant Roots and Permeability Barriers to Gas-exchange with the Rhizosphere: a Microelectrode and Modelling Study with Phragmites australis

Adventitious roots of intact Phragmites plantlets were securedhorizontally 2–3 mm below the surface of an oxygen-depletedfluid agar across which oxygen-free nitrogen was gently streamedto create a constant oxygen sink; the leafy shoot was fullyexposed to air. Radial oxygen profiles through rhizosphere androot at different distances from the apex were obtained polarographicallyusing Clark-type bevelled microelectrodes servo-driven in stepsof 10 µm (root) or 10–50 µm (rhizosphere).The pattern of radial oxygen loss (ROL) typical of wetland plants,viz. high at the apex and declining sharply sub-apically, wasrelated to synergism between ROL, and oxygen consumption andincreasing impedance to diffusion within the epidermal/hypodermalcylinder rather than to a surface resistance. The smallest oxygendeficit (2 kPa) to develop across the 80 µm thick epidermal/hypodermalcylinder was within the apical 10 mm and was consistent withtissue oxygen diffusivities similar to water. At 100 mm fromthe apex, consumption and impedance had increased the deficitto about 15 kPa and reduced ROL almost to zero. The developingimpedance within the epidermal/hypodermal cylinder was leastin cell layers immediately adjoining the cortex and increasedmost in the hypodermal cell layer abutting the epidermis. Thesub-apical decline in ROL appeared to coincide with the appearanceof aerenchyma in the cortex but thin walled ‘passage areas’(windows) in the hypodermal/epidermal cylinder persisted locallyand remained leaky to oxygen to some degree. It is through thesewindows that lateral roots emerge and the cortex in line withthe windows remains non-aerenchymatous. The radial and longitudinaloxygen profiles were consistent with modelling predictions.The shapes of the stelar oxygen profiles were consistent witha higher oxygen demand in the outer region (viz. pericycle,phloem, protoxylem and early metaxylem cylinder) than in theinner core (late metaxylem cylinder and medulla), but the deficitswere relatively small (     

Development of the Hypodermal Casparian Band in Corn and Onion Roots

A hypodermal Casparian band develops 40–50 mm from theroot tip in corn and 30–40 mm from the root tip in onion.In both plants, the endodermal Casparian band matures about20 mm closer to the root tip than the hypodermal Casparian band.Using the apoplastic fluorescent dye, Calcofluor white M2R (CFW),a permeability barrier could be distinguished in the radialwalls of the hypodermis 40–50 mm from the root tip incorn and onion. In progressively younger regions of the roots,CFW was first excluded from the outer tangential hypodermalwalls and the inner tangential epidermal walls, then the radialepidermal walls so that in very young regions only the outertangential epidermal walls were permeated. In contrast to CFW,the symplastic fluorescent dye, uranin, was translocated fromthe epidermis into the stele at all distances tested (5.0–50mm from the root tips). CFW and uranin at a concentration of0.01% proved nontoxic to corn and onion roots on the basis ofroot growth tests. Key words: Zea mays, Casparian band, Hypodermis, Allium cepa     

Scanning Electron Microscopy of Plant Roots

A glycol methacrylate infiltration and polymerization techniquewas used to prepare clover roots inoculated with Rhizobium forscanning reflection electron microscopy. Root hairs and epidermalcells were coated with many bacteria; some bacteria seemed tobe embedded in the wall surface. Root hair tips were often smoothbut some older root hair surfaces showed a fibrillar meshworkpattern. Small granules c. 0.18 µm diameter were presenton the root hair and epidermal cell walls. The root cap, someroot hairs, and some epidermal cells were covered by an amorphousfilm thought to be the mucigel.     

Progressive Cortical Senescence and Formation of Lysigenous Gas Space (Aerenchyma) Distinguished by Nuclear Staining in Adventitious Roots of Zea mays

The pattern of loss of nuclear integrity in the epidermis andcortex of maize adventitious roots was examined during (1) non-pathogeniccortical senescence associated with root ageing, and (2) lysigenousaerenchyma formation, to determine whether these phenomena arerelated. Nuclear integrity was estimated by counting the percentageof cells with nuclei detectable by acridine orange fluorescence. In roots of both soil-grown (90 d) and solution-grown (19 d)plants, nuclei were lost progressively, from the epidermis andfrom successively deeper cortical cell layers, with increasingdistance behind the root tips; this occurred irrespective ofthe degree of aeration in solution culture, and independentlyof aerenchyma formation. Aerenchyma developed in soil-grownplants and in sub-ambient oxygen concentrations (<5 kPa partialpressure) in solution culture. It started to form in the middlecortex and coincided with a marked loss of nuclear stainingin the inner cortex, especially in the innermost cortical celllayer next to the endodermis, but not in the remaining cellsof the middle cortex. Two distinct patterns of nuclear deletionfrom the cortex were thus demonstrated; they occurred independentlybut simultaneously in some conditions. These findings are discussed in relation to mechanisms of celldeath, and the metabolic status of root cortical cells participatingin ion transport to the xylem. Zea mays L., maize, roots, aerenchyma, cell death, nuclei     

Exploring the Radial and Longitudinal Aeration of Primary Maize Roots by Means of Clark-Type Oxygen Microelectrodes

Clark-type oxygen microelectrodes were used to measure the radial and longitudinal oxygen distribution in aerenchymatous and nonaerenchymatous primary roots of intact maize seedlings. A radial intake of oxygen from the rooting medium was restricted by embedding the roots in 1% agar causing aeration to be largely dependent upon longitudinal internal transport from the shoot. In both root types, oxygen concentrations declined with distance from the base, and were lower in the stele than in the cortex. Also, the bulk of the oxygen demand was met internally by transport from the shoots, but a little oxygen was received by radial inward diffusion from the surrounding agar, and in some positions the hypodermal layers received oxygen from both the agar and the cortex. Near to the base, the oxygen partial pressure difference between the cortex and the center of the stele could be as much as 6–8 kPa. Nearer to the tip, the differences were smaller but equally significant. In the nonaerenchymatous roots, cortical oxygen partial pressures near the apex were becoming very low (< 1 kPa) as root lengths approached 100 mm, and towards the center of the stele values reached 0.1 kPa or lower. However, the data indicated that respiratory activity did not decline until the cortical oxygen pressure was less than 2 kPa. Mathematical modeling based on Michaelis–Menten kinetics supported this and suggested that the respiratory decline would be mostly restricted to the stele until cortical oxygen pressures approached very low values. At a cortical oxygen pressure of 0.75 kPa, it was shown that respiratory activity in the pericycle and phloem might remain as high as 80–100% of maximum even though in the center of the stele it could be less than 1% of maximum. Aerenchyma production resulted in increases in oxygen concentration throughout the roots with cortical partial pressures of ca. 5–6 kPa and stelar values of ca. 3–4 kPa near the tips of 100 mm long roots. In aerenchymatous roots, there was some evidence of a decline in the oxygen permeability of the epidermal–hypodermal cylinder close to the apex; a decline in stelar oxygen permeability near the base was indicated for both root types. There was some evidence that the mesocotyl and coleoptile represented a very significant resistance to oxygen transport to the root.     

Increased Synthesis of ABA in Partially Dehydrated Root Tips and ABA Transport from Roots to Leaves

Zhang, J. and Davies, W. J. 1987. Increased synthesis of ABAin partially dehydrated root tips and ABA transport from rootsto leaves.—J. exp. Bot. 38: 2015–2023. Isolated root tips of pea (Pisum sativum L. cv. Feltham First)and Commelina communis L. were air-dried until they lost between10% and 40% of their fresh weight, followed by a period of incubationat these reduced water contents. These treatments resulted inincreased ABA production, suggesting that root tips of bothspecies have the capacity to synthesize ABA in increased amountswhen water deficits develop in the root. The ABA concentrationin pea roots increased linearly as turgors fell below about0·15 M Pa and relative water contents (R WC) fell below90%. Commelina roots produced more ABA when RWC fell below asimilar value but the threshold turgor for increased ABA productionin Commelina roots was around 0·30 MPa. Roots of intact plants loaded with ABA as a result of incubationin solutions of varying concentrations provided ABA to leaveswhich resulted in increased ABA concentrations in the leaveswhen these were assayed several hours later. This occurred whenthese roots were not contributing substantially to transpirationalflux. Leaves on shoots that were enclosed and darkened and thereforenot transpiring, did not accumulate ABA from ‘loaded’roots. A role for root-sourced ABA in root-to-shoot communication ofthe effects of soil drying is discussed. Key words: ABA, roots, water relations     

High pH Causes Disintegration of the Root Surface in Lupinus angustifolius L.

The effect of high pH on the morphology and anatomy of the rootsof lupin (Lupinus angustifolius L. cv. Yandee) and pea (Pisumsativum L. cv. Dundale) was examined in buffered solution. Themorphology and anatomy of lupin roots were markedly altered,and root growth was reduced by increasing solution pH from 5·2to 7·5, whereas pea roots were unaffected. In lupin roots,pH 7·5 caused disintegration of the root surface andimpaired root hair formation. Lupin roots grown at pH 7·5also had decreased cell lengths but increased cell diameterin both the epidermis and the cortex in comparison to rootsgrown at pH 5·2. High pH reduced cell volume greatlyin the epidermis, to a lesser extent in the outer cortex andnot at all in the inner cortex. It appears that in lupins, theprimary detrimental effects of growth at pH 7·5 is reducedlongitudinal growth of cells near the root surface with a consequentreduction in elongation of the cells in inner cortex.Copyright1993, 1999 Academic Press Lupinus angustifolius L., Pisum sativum L., high pH, root morphology, root anatomy     

Ethylene Production by Fe-deficient Roots and its Involvement in the Regulation of Fe-deficiency Stress Responses by Strategy I Plants

Species that showed marked morphological and physiological responsesby their roots to Fe-deficiency (Strategy I plants) were comparedwith others that do not exhibit these responses (Strategy IIplants). Roots from Fe-deficient cucumber (Cucumis sativusL.‘Ashley’), tomato (Lycopersicon esculentumMill.T3238FER) and pea (Pisum sativumL. ‘Sparkle’) plantsproduced more ethylene than those of Fe-sufficient plants. Thehigher production of ethylene in Fe-deficient cucumber and peaplants occurred before Fe-deficient plants showed chlorosissymptoms and was parallel to the occurrence of Fe-deficiencystress responses. The addition of either the ethylene precursorACC, 1-aminocyclopropane-1-carboxylic acid, or the ethylenereleasing substance, Ethephon, to several Fe-sufficient StrategyI plants [cucumber, tomato, pea, sugar beet (Beta vulgarisL.),Arabidopsis(Arabidopsis thaliana(L.) Heynh ‘Columbia’), plantago(Plantago lanceolataL.)] promoted some of their Fe-deficiencystress responses: enhanced root ferric-reducing capacity andswollen root tips. By contrast, Fe-deficient roots from severalStrategy II plants [maize (Zea maysL. ‘Funo’), wheat(Triticum aestivumL. ‘Yécora’), barley (HordeumvulgareL. ‘Barbarrosa’)] did not produce more ethylenethan the Fe-sufficient ones. Furthermore, ACC had no effecton the reducing capacity of these Strategy II plants and, exceptin barley, did not promote swelling of root tips. In conclusion,results suggest that ethylene is involved in the regulationof Fe-deficiency stress responses by Strategy I plants.Copyright1999 Annals of Botany Company. Arabidopsis (Arabidopsis thaliana(L.) Heynch), barley (Hordeum vulgareL.), cucumber (Cucumis sativusL.), ethylene, iron deficiency, maize (Zea maysL.), pea (Pisum sativumL.), plantago (Plantago lanceolataL.), ferric-reducing capacity, sugar beet (Beta vulgarisL.), tomato (Lycopersicon esculentumMill.), wheat (Triticum aestivumL.).     

Redistribution of Phosphorus, Sulfur, Potassium and Calcium in Relation to Light-Induced Gravitropic Curvature in Zea Roots

Phosphorus, sulfur, potassium and calcium were measured by particle-inducedX-ray emission (PIXE) in horizontally oriented, light-irradiatedor non-irradiated primary roots of Zea mays L., cv. Golden CrossBantam 70 which exhibit gravitropic response only after exposureto light. The content of the four elements increased in thelower half of horizontally oriented roots which had been brieflyexposed to white or red light, while there were no marked differencesin distribution between the upper and lower halves of non-irradiatedroots. The increase of each element in the lower half was observed15–30 min after irradiation in root caps and 30–60min after irradiation in the elongation zones. The effect ofred light was not reversed by far-red light given immediatelyafter the red irradiation. Ethylene glycol-bis(ß-aminoethylether)-N, N, N',N'-tetraacetic acid (EGTA) treatment of roottips inhibited the gravitropic curvature of roots, and the additionof Ca reduced this inhibition. The meaning of Ca redistributionin root caps and elongation zones during light-induced gravitropiccurvature of maize roots is discussed. (Received December 4, 1985; Accepted March 22, 1986)     

Root Growth and Carbohydrate Metabolism at Low Temperatures

A study of carbohydrate metabolism in the roots of pea and maizeshows that the differing ability of these two species to growat low temperatures is associated with the maintenance of adequatesugar supplies to the root tip. In the maize root reducing theambient temperature to 2 °C causes a sharp and continuingfall in the soluble sugar content of the growing tip. A similartreatment with pea roots causes only a temporary reduction insugar content lasting no more than 24 h. The fall in root sugarsin maize is accompanied by a reduction in respiration rate andthe cessation of growth. During the periods of sugar shortagecaused by low temperatures both respiration and growth can bestimulated in root tips by a supply of exogenous glucose. Pearoots also show an additional ability to adapt to low temperaturesby lowering the K^m value for invertase after pretreatment ata low temperature. This effect is not seen in maize.     

Rectifier-like Activities of Roots of Two Desert Succulents

Axial and radial water flows for roots in response to appliedhydrostatic pressure drops, water loss from roots after variousperiods of drying, and development of new roots after rewettingdroughted plants were examined for two sympatric desert succulents.Agave deserti Engelm. and Ferocactus acanthodes (Lemaire) Brittonand Rose. For a 40 kPa hydrostatic pressure drop applied to20 mm long root pieces, radial water flows from the epidermisto the root xylem were 2- to 5-fold greater at the tip thanat midlength and were much less than axial flows along the xylem.Upon drying detached roots in air at 20 °C and a water vapoursaturation deficit of 1.2 kPa (50% relative humidity), radialwater flow decreased more than 10-fold in 3–6 h, and couldrecover to the original level 6 h after rewetting. The rateof water loss from attached roots of plants dried in air at20 °C and a 1.2 kPa saturation deficit decreased about 200-foldin 72 h, which would greatly limit water loss from the plantto a drying soil. At 96 h after rewetting roots of A. desertithat had been exposed to air at 20 °C and a 1.2 kPa saturationdeficit for 120 h, rehydration of existing roots and developmentof new roots contributed about equally to water uptake by thewhole plant. In summary, roots of these desert succulents canreadily take up water from a wet soil but do not lose much waterto a dry soil, thus effectively acting like rectifiers withrespect to plant-soil water movement. Key words: Agave, Cactus, Drought, Root, Water flow, Xylem     

Cell division activation in the quiescent center of excised maize root tip

The phenomenon of activating cell proliferation in the quiescent center of excised maize roots is described. The root tips were grown on wet filter paper in Petri dishes. This phenomenon was observed in 8 to 14 maize cultivars and was absent in excised Arabidopsis root tips. The distribution of mitoses in meristems greatly varied in individual seedlings roots from the same seed lot and seedlings of different cultivars. Meristem opening was observed after the removal of small root tips not longer than 3 mm and intact seminal roots. Sucrose (2%) and 10⁻⁶–10⁻⁸ M indole-3-acetic acid did not prevent meristem opening. These findings indicate that the state of quiescent center is maintained by a system of intercellular and interorgan relations, which are to be clarified.     

The Possible Involvement of Root-cap Mucilage in Gravitropism and Calcium Movement Across Root Tips of Allium cepa L

Roots of Allium cepa L. grown in aerated water elongate rapidly,but are not graviresponsive. These roots (1) possess extensivecolumella tissues comprised of cells containing numerous sedimentedamyloplasts, (2) lack mucilage on their tips, and (3) are characterizedby a weakly polar movement of calcium (Ca) across their tips.Placing roots in humid air correlates positively with the (1)onset of gravicurvature, (2) appearance of mucilage on tipsof the roots, and (3) onset of the ability to transport Ca polarlyto the lower side of the root tip. Gravicurvature of roots previouslysubmerged in aerated water is more rapid when roots are orientedvertically for 1–2 h in humid air prior to being orientedhorizontally. The more rapid gravicurvature of these roots correlatespositively with the accumulation of mucilage at the tips ofroots during the time the roots are oriented vertically. Therefore,the onset of gravicurvature and the ability of roots to transportCa to the lower sides of their tips correlate positively withthe presence of mucilage at their tips. These results suggestthat mucilage may be important for the transport of Ca acrossroot caps. Allium cepa, root gravitropism, root mucilage, calcium, onion     

Effect of Root and Shoot Meristem Temperature on Shoot to Root Dry Matter Partitioning and the Internal Concentrations of Nitrogen and Carbohydrates in Maize and Wheat

Maize (Zea mays L.) and spring wheat (Triticum aestivum L.)were grown in nutrient solution at uniformly high air temperature(20 °C), but different root zone temperatures (RZT 20, 16,12 °C). To manipulate the ratio of shoot activity to rootactivity, the plants were grown with their shoot base includingthe apical meristem either above (i.e. at 20 °C) or withinthe nutrient solution (i.e. at 20, 16 or 12 °C). In wheat, the ratio of shoot:root dry matter partitioning decreasedat low RZT, whereas the opposite was true for maize. In bothspecies, dry matter partitioning to the shoot was one-sidedlyincreased when the shoot base temperature, and thus shoot activity,were increased at low RZT. The concentrations of non-structuralcarbohydrates (NSC) in the shoots and roots were higher at lowin comparison to high RZT in both species, irrespective of theshoot base temperature. The concentrations of nitrogen (N) inthe shoot and root fresh matter also increased at low RZT withthe exception of maize grown at 12 °C RZT and 20 °Cshoot base temperature. The ratio of NSC:N was increased inboth species at low RZT. However this ratio was negatively correlatedwith the ratio of shoot:root dry matter partitioning in wheat,but positively correlated in maize. It is suggested that dry matter partitioning between shoot androots at low RZT is not causally related to the internal nitrogenor carbohydrate status of the plants. Furthermore, balancedactivity between shoot and roots is maintained by adaptationsin specific shoot and root activity, rather than by an alteredratio of biomass allocation between shoot and roots.Copyright1994, 1999 Academic Press Wheat, Triticum aestivum, maize, Zea mays, root temperature, shoot meristem temperature, biomass allocation, shoot:root ratio, carbohydrate status, nitrogen status, functional equilibrium     

The Effects of Mechanical Impedance to Growth on the Levels of ABA and IAA in Root Tips of Zea mays L.

Extraction and analytical methods have been refined and newones devised to allow precise determinations by GC-EC of thelevels of abscisic acid (ABA) and indol-3ylacetic acid (IAA)in samples of maize root tips as small as 1.0 g fr. wt. Seminalroots of 5-d-old maize seedlings grown in normal (bulk density1200 kg m^–3) and compacted (bulk density 1600 kg m^–3)sand/garden loam mixtures have been examined. Seminal rootsfrom compacted soil had an average length of about 40% of thatof control roots and were much thicker. The ABA levels in 10mm tips of impeded roots (c. 25–35 ng g^–1 fr.wt.)did not differ significantly from those of normal root tipson both a fresh and dry weight basis. The levels in 0–1mm tips were approximately double those in the remaining 1–10mm zones. IAA levels were increased by about 3 times in impededroots (176.3 as compared with 52.4 ng g^–1 fr.wt) and itis concluded that this response is likely to be the main causeof the morphological and growth changes brought about by soilcompaction.     

Depletion of Oxygen in Roots of Wheat and Maize by Respiration Following Encasement in Olive Oil

The extent to which isolated root systems of 14-d-old seedlingsof wheat (Triticum aestivum) and maize (Zea mays) were depletedof oxygen by respiration was measured after immersing them inoutgassed olive oil to exclude oxygen entry from the air. Atintervals over 45 min, gas from the roots was removed underpartial vacuum and oxygen partial pressures measured by gaschromatography. Contrary to earlier findings (Erdmann and Wiedenroth,1988), roots were able to utilize almost all of their oxygenwithin 20 min at 25°C, including that dissolved in the thinwater covering interposed between roots and oil. The rate ofaerobic respiration could be estimated readily from the timecourse of oxygen depletion.Copyright 1993, 1999 Academic Press Triticum aestivum, Zea mays, roots, anaerobiosis, respiration, oxygen, methods     

Modelling of the Hydraulic Architecture of Root Systems: An Integrated Approach to Water Absorption--Distribution of Axial and Radial Conductances in Maize

The ‘Hydraulic Tree Model’ of the root system simulateswater uptake through root systems by coupling a root architecturemodel with laws for water flow into and along roots (Doussan,Pagès and Vercambre,Annals of Botany81: 213–223,1998). A detailed picture of water absorption in all roots comprisingthe root system is thus provided. Moreover, the influence ofdifferent distributions of radial and axial hydraulic conductancesin the root system on the patterns of water uptake can be analysed.Use of the model with Varney and Canny's data (1993) for flowalong maize roots demonstrated that a constant conductance inthe root system cannot reproduce the observed water flux profiles.Taking into account the existing data on hydraulic conductancesin maize roots, we fitted the distribution of conductances inthe root system to the observed flux data. The result is that,during root tissue maturation, the radial conductivity decreasesby one order of magnitude while the axial conductance increasesby about three orders of magnitude. Both types of conductanceexhibit abrupt changes in their evolution. Due to the conductancedistribution in the root system, appreciable water potentialgradients may develop in the roots, in both the branch rootsand main axes. An important point is that the conductance distributionin the branch roots described by the model should be relatedto the age of the tissue (and not the distance from the branchroot tip) and is therefore closely related to the developmentprocess. Thus for branch roots, which represent about 90% ofthe calculated total water uptake in 43-d-old maize, water absorptionwill depend on the opening of the metaxylem in the axes, andon the time dependent variation of the conductances in the branchroots.Copyright 1998 Annals of Botany Company Water; absorption; root system; architecture; model; hydraulic conductance;Zea maysL.     

Ion Transport to the Xylem in Aerenchymatous Roots of Zea mays L.

Aerenchyma was induced in the developing nodal (adventitious)roots of maize (Zea mays L. cv. LG11) in solution culture bytreatment with an atmosphere of 5% (v/v) oxygen (oxygen partialpressure 5.0 k Pa). Ion uptake by aerenchymatous and ordinary(non-aerenchymatous) roots was then compared to determine theeffect of degeneration of much of the cortex in aerenchymatousroots on radial ion movement to the xylem. Rates of uptake andtransport of phosphate, potassium and chloride were followedby labelling 4–12mm length segments of intact roots usingradioactive tracers. In some experiments, enhanced rates ofphosphate transport were induced by means of a divided rootsystem and nutrient deprivation. For all three nutrient ions,and over a range of concentration for phosphate (10–100mmol m^–3 and potassium (0·25–5·0 molm^–3 and with enhanced rates of transport, aerenchymatousroots were at least as effective as ordinary roots (per mm³of root) and sometimes more effective (per mm root length).These findings are discussed in relation to hypotheses concerningthe pathways for radial ion movement across the cortex in ordinaryand aerenchymatous roots. Correspondence to: Long Ashton Research Station, Long Ashton,Bristol BS18 9AF, U.K. Key words: Aerenchyma, ion transport, mineral nutrition, oxygen deficiency, radio-isotopic tracers, roots, structure, Zea mays     

Experimental Modification of Cell Division Patterns in the Root Meristem of Zea mays

In the meristem of the young primary root of maize seedlingsthe first transverse division in the cortex 250 µm fromthe root apex results in two daughter cells of distinctly unequalsize. This division could be rendered equal by raising the seedlingsin up to 7.5% methanol. The pattern of the subsequent two orthree transverse divisions in the cortex, as revealed by thearrangement of the newly divided cells in the resultant cellularpackets, was acropetal in the methanol-treated roots but basipetalin the control roots. The sequence of division within a cellularpacket tended to follow the distribution of cell sizes - largercells divided earlier than smaller cells. A temporary arrestof cell division by exposing roots to cold (5 °C) conditionshad no effect on the sequence of divisions that followed whenthe roots were allowed to recover at 20 °C. The resultssuggest that the normally asymmetric position of the cell wallformed at cytokinesis is subject to active regulation and thatmethanol interferes with this process. The cytoplasm of certaincells in the root meristem was also found to be unequally distributed,as judged by Azure B staining, between the two ends of the cell.Cytoplasmic asymmetry was not directly correlated with inequalityof division, although it too was affected by methanol. Cell polarity, root meristem, unequal division, Zea mays