Effects of Temperature on the Development of Metaxylem in Primary Wheat Roots and Its Hydraulic Consequence

The effects of different temperatures on the development ofmetaxylem were studied in the primary seminal root of winterwheat (Triticum aestivum L.) seedlings. Xylem development wasstudied microscopically at different distances behind the rootapex after safranin staining to reveal lignification. Diameter of the central late metaxylem (LMX) and its proportionto the stele cross-sectional area increased in the acropetaldirection. Diameter of the LMX and stele decreased with an increasein growing temperature. Numbers of early metaxylem (EMX) wereseven, seven and six at 10, 20 and 30 C, respectively. EMXwas lignified much more rapidly than the LMX along the seminalroot axes. Lignification of xylem elements commenced furthertowards the root apex at the higher temperatures. The LMX vesselsof the roots grown at the higher temperature had thicker secondarywalls. The relative conductivity of seminal roots, calculated fromPoiseuille's equation, decreased as growing temperature increased.In a drought-prone environment where wheat plants rely heavilyon stored soil water, a lowered axial conductivity in the rootswould be advantageous. The plants would tend to conserve waterduring vegetative growth for use during the critical periodsof flowering and grain-filling. Breeders selecting wheat plants for altered LMX diameters shouldcontrol temperatures during primary root development, sectionthe roots at the same distance from the tip and be aware thatcross walls may exist in the LMX for up to 30 cm from the tip. Wheat, Triticum aestivum L., roots, xylem development, hydraulic conductivity, temperature     

Late Maturation of Large Metaxylem Vessels in Soybean Roots: Significance for Water and Nutrient Supply to the Shoot

The large, late metaxylem (LMX) in the roots of soybean beginsdevelopment in the centre of the stele after lignification ofthe early metaxylem poles. Subsequent maturation of the firstappearing LMX elements is gradual. They were never mature inthe 8-d-old seedlings examined. In 10 to 15-d-old plants thefirst LMX matured to open vessels at a mean of 17 cm proximalto the root tip. Additional LMX vessels developed in more proximalregions of the roots and these also matured gradually. Based on calculations from relative vessel diameters, the potentialflow of xylem sap in a single central LMX vessel is 50 timesthat in the total of all the early metaxylem (EMX) vessels ofa typical primary root of soybean. There was a marked dependence of relative leaf area on the lengthof primary root with open LMX vessels. This may result fromthe predicted increased water and nutrient flow to the shoot,facilitated by the opening of the large vessels. It is suggestedthat, as in maize, the living LMX elements may function in ionaccumulation. Dicotyledonous roots, soybean, Glycine max, xylem vessels, xylem maturation, water conduction     

Root metaxylem and architecture phenotypes integrate to regulate water use under drought stress

At the genus and species level, variation in root anatomy and architecture may interact to affect strategies of drought avoidance. To investigate this idea, root anatomy and architecture of the drought‐sensitive common bean (Phaseolus vulgaris) and drought‐adapted tepary bean (Phaseolus acutifolius) were analyzed in relation to water use under terminal drought. Intraspecific variation for metaxylem anatomy and axial conductance was found in the roots of both species. Genotypes with high‐conductance root metaxylem phenotypes acquired and transpired more water per unit leaf area, shoot mass, and root mass than genotypes with low‐conductance metaxylem phenotypes. Interspecific variation in root architecture and root depth was observed where P. acutifolius has a deeper distribution of root length than P. vulgaris. In the deeper‐rooted P. acutifolius, genotypes with high root conductance were better able to exploit deep soil water than genotypes with low root axial conductance. Contrastingly, in the shallower‐rooted P. vulgaris, genotypes with low root axial conductance had improved water status through conservation of soil moisture for sustained water capture later in the season. These results indicate that metaxylem morphology interacts with root system depth to determine a strategy of drought avoidance and illustrate synergism among architectural and anatomical phenotypes for root function.     

Pathways and processes of water and nutrient movement in roots

Recent work in our laboratory provides evidence for a revised view of the functioning of roots of maize, and probably of all the grasses. The development of coherent soil sheaths on the distal 30-cm of these roots, and the loss of the sheaths further back, led us to investigate the differences in surface structure, anatomy, carbon exudation and microflora of the sheathed and bare zones. The significant differences are summarized. But the fact which underlies all these differences is the maturation of the late metaxylem (LMX). In the sheathed zones the LMX elements are still alive and non-conducting; only the early metaxylem (EMX) and protoxylem are open. In the bare zones they are open vessels. This leads directly to the dryness of bare zones and the wetness of sheathed zones, and indirectly to the other differences noted. Branch root junctions are shown to be structures of great significance. Besides connecting the branches to the axile systems, they serve also to connect the EMX and LMX vessels, and contain a tracheid barrier which prevents air embolisms entering the main vessels. These discoveries force us to revise the traditional view of water uptake by the root hair zone, and to suggest that much water must also enter bare roots, possibly via the laterals. There is some published evidence for this. The living LMX elements of the sheathed zone accumulate large concentrations of potassium which must joint the transpiration water at the transition to the bare zone. Calculations suggest that this may be only a tenth of the requirement of a mature plant, and that the balance may enter the bare zones with the transpiration water.     

Effects of Root Zone Restriction on Amino Acid Status and Bean Plant Growth

The possibility that the suppression of shoot growth in restrictedroot zone plants (RRZP) is caused by a deficiency in N-aminocompounds (NAC) in the shoot, possibly due to an insufficientsupply from the roots, was studied in bean (Phaseolus vulgarisL.). Root zone restriction to 10 cm³ in an aerated nutrientsolution resulted in suppressed plant growth, as compared withcontrol plants grown in a non-limiting root zone volume. Rootxylem exudation of solution and N-amino compunds (NAC) followingdecapitation was much greater in the control, as compared withRRZP, both per plant and per unit root fresh weight (FWT). Inboth treatments, asparagine comprised more than 52% of the NACfraction in the root xylem exudate (RE). Its reduced exudationin the RRZP was of a proportion similar to the combined fractionof NAC left over in both treatments. Asparagine accumulationin leaves of the control plants was very high, comprising 73%of the total NAC pool, while in RRZP, it was much smaller anddid not exceed 25%. The total NAC amount per unit of leaf FWTwas 3·3 times smaller for the RRZP, as compared withthe control, resulting mainly from the dramatic drop in asparagineaccumulation. In the roots, RRZP accumulated more NAC per unitroot FWT than the control. Raising both treatments in distilledwater reduced considerably the accumulation of NAC, includingasparagine, in their leaves. RRZP was relatively more suppressedby the absence of nutrients than control plants. This phenomenondid occur, despite the fact that NAC and asparagine concentrationsin the root and shoot of RRZP were greater than in the controlwhen grown in distilled water; Therefore, it was concluded thatroot zone restriction might affect the accumulation of NAC andasparagine in the leaves, but that deficiency in these compoundsis not the primary or the major cause of growth suppressionin RRZP. Key words: Root zone restriction, asparagine, amino-acids, Phaseolus vulgaris     

Vascular tissue-specific gene expression of xylem sap glycine-rich proteins in root and their localization in the walls of metaxylem vessels in cucumber

Root-specific cDNAs of glycine-rich protein (cucumber root glycine rich protein-1 and -2; CRGRP-1 and CRGRP-2) were cloned previously by use of an antiserum raised against whole xylem sap of Cucumis sativus. The accumulation of the corresponding mRNA at high levels was detected in the root-hair zone of cucumber tap root [Sakuta et al. (1998) Plant Cell Physiol. 39: 1330]. The RNA gel blot analysis with the CRGRP-1- and -2-specific probes revealed that the CRGRP genes expressed only in root but not at all in aboveground organs. When the localization of these mRNAs were examined by in situ hybridization, CRGRP mRNAs were found only in the parenchyma cells in the central cylinder of young lateral roots and it was most abundant in the cells that surrounded xylem vessels in the root-hair zone of the tap root. In immunoblotting of xylem sap collected from cucumber stem with an antiserum raised against CRGRP-1 that had been produced in an E. coli expression system, the antibodies, which did not cross-react with GRP1.8 of kidney bean, reacted with two proteins, whose mobilities corresponded to those of proteins deduced from the CRGRP-1 and -2 cDNAs. Immunohistochemical staining revealed that the CRGRPs accumulated specifically in the lignified walls of metaxylem vessels in the root, stem and leaf and in the lignified cell walls of perivascular fibers in cucumber stems. Immunostaining was also detected in the walls of metaxylem vessels and in the cell walls of adjacent sclerenchyma in the hypocotyl of kidney bean. These data clearly indicate that the novel glycine-rich proteins were produced in the vascular tissue of the root, transported systemically over a long distance via the xylem sap and immobilized in the walls of metaxylem vessels and sclerechyma cells in aboveground organs.     

Sodium Recirculation and Loss from Phaseolus vulgaris L.

In a split-root experiment, ²²Na was supplied to Phaseolus vulgarisL. roots emerging from the stem, 2.5 cm above the main roots.Sodium exported from these upper roots was translocated a shortdistance upward in the stem and downward to the main roots.Most of the ²²Na arriving in the main roots was lost to themedium. Sodium loss from P. vulgaris roots into KCI or NaCl was similarand was not affected by oligomycin. The results confirm a previous hypothesis regarding the mechanismof sodium exclusion from the tops of sodium non-accumulatorplants. Phaseolus vulgaris L., bean, sodium transport     

Zinc Toxicity and Xylem Vessel Wall Alterations in White Beans

When white beans are exposed to excess zinc, reddish brown patchesappear along the leaf veins. Ultrastructural observations ofthe xylem vessels in the discoloured zones show several modificationsof the vessel walls including gelation of the pit membranes,coating of the lumen surface with an abnormal layer and depositionof electron-dense material in the secondary vessel walls. Histochemicalstudies indicate that the altered pit membranes and the coatinglayer stain positively for lipid, while the secondary wall depositsstain positively for phenolic compounds. Phaseolus vulgaris L., white bean, xylem vessels, zinc toxicity     

Inhibition of phosphorus and water passage across intact roots by polyethylene glycol and phenylmercuric acetate

Application of polyethylene glycol or phenylmercuric acetate to intact bean (Phaseolus vulgaris L., cv. Red Wade) roots inhibited passage of phosphorus across the roots to the xylem. The same results occurred for foliar application of phenylmercuric acetate when time was allowed for absorption and distribution of the chemical in the plant. For both chemicals the inhibition of phosphorus was proportional to or greater than any accompanying restriction on water flow across the root.     

Root Gravitropism and Below-ground Competition among Neighbouring Plants: A Modelling Approach

Competition for nutrients among neighbouring roots occurs whentheir individual depletion volumes overlap, causing a reductionin nutrient uptake. By exploring different spatial niches, plantswith contrasting root architecture may reduce the extent ofcompetition among neighbouring root systems. The main objectivesof this study were: (1) to evaluate the impact of root architectureon competition for phosphorus among neighbouring plants; and(2) to compare the magnitude of competition among roots of thesame plant vs. roots of neighbouring plants. SimRoot, a dynamicgeometric model, was used to simulate common bean root growthand to compare the overlap of depletion volumes. By varyingthe gravitropism of basal roots, we simulated three distinctroot architectures: shallow, intermediate and deep, correspondingto observed genetic variation for root architecture in thisspecies. Combinations of roots having the same architectureresulted in more intense inter-plant competition. Among them,the deep-deep combination had the most intense competition.Competition between deep root systems and shallow root systemswas only half that of deep root systems competing with otherdeep root systems. Inter-plant root competition increased assoil diffusivity increased and the distance among plants decreased.In heterogeneous soils, co-localization of soil resources androots was more important in determining resource uptake thaninter-plant root competition. Competition among roots of thesame plant was three- to five-times greater than competitionamong roots of neighbouring plants. Genetic variation for rootarchitecture in common bean may be related to adaptation todiverse competitive environments. Copyright 2001 Annals of BotanyCompany Root architecture, phosphorus, competition, common bean, Phaseolus vulgaris L. nutrient uptake, gravitropism     

Ultrastructure of xylem parenchyma cells of barley roots in relation to ion transport to the xylem

Summary The structure of xylem parenchyma cells is examined in relation to transport of ions through the root. Measurement of uptake of ⁸⁶Rb⁺ and its transport through the root at different distances from the apex show that this is a general activity along the length of the root and not confined to a limited region. Thus transport through the root is not stopped by removal of that part of the root tip containing metaxylem vessels with living contents. The structure of xylem parenchyma appears to be suitable for involvement in ion transport from the stele to the xylem. At 1 cm behind the tip, where metaxylem vessels have no living contents but ion uptake and transport are going on at high rates, xylem parenchyma cells are rich in cytoplasm with extensive rough endoplasmic reticulum and well-developed mitochondria. Their cell walls contain numerous plasmodesmata, establishing the possibility of a symplastic pathway across the stele up to the vessels. The results are discussed in relation to regulation of ion transport to the xylem vessels in roots.Dedicated to Professor O. Stocker on the occasion of his 85th birthday.     

Cell Wall Thickening of the Xylem Tracheary Elements in Different Zones of the Adventitious Root of Allium cepa L.

Cell wall thickness of the xylem tracheary elements was measuredin the proto- and metaxylem of the Allium cepa L. adventitiousroot. Measurements were taken in root fragments of known age(1, 3, 5, 7 and 9 d) located in either the basal or medio-apicalzone. Tracheary elements in the protoxylem matured within ashorter period of time than those in the metaxylem. Final cellwall thickness was greater in metaxylem than in protoxylem components.The cell wall thickening in the tracheary elements in both proto-and metaxylem was more rapid in the basal zone of the root thanin the medio-apical zone. Additionally, cell walls of the maturetracheary elements were thicker in the basal zone than in areasfurther from the bulb. Allium cepa, onion, root, cell wall, xylem maturation     

Solute concentrations in xylem sap along vessels of maize primary roots at high root pressure

The elemental composition of xylem sap has been determined by cryo-analytical microscopy in situ along vessels in the roots of maize plants frozen intact while root pressure was high. The only chemical element (including carbon) present in significant concentrations in the vessels was potassium at 11 mM and 15 mM in the late (LMX) and early (EMX) metaxylem, respectively. There was no gradient of [K] along the vessels, which each run the length of the mature proximal end of the roots. At the distal end of each vessel, in the oldest still living vessel, which each run the length of the mature proximal end of the roots. At the distal end of each vessel, in the oldest still living vessel elements, there was sharp rise in [K] to 110 mM and 130 mM in the LMX and EMX, respectively.     

Anatomical root variations in response to water deficit: wild and domesticated common bean (Phaseolus vulgaris L)

Root anatomical responses to water deficit are diverse and regulation of water uptake strongly depends on plant anatomy. The ancestors of common bean (Phaseolus vulgaris L.) cultivars are the wild common beans. Because wild beans adapt and survive well in the natural environment, it is hypothesized that wild common bean roots are less affected than those of domesticated beans at low substrate water potential (ψW). A wild common bean accession from Chihuahua Mexico and cv. Bayomex were studied. Seedlings with a mean root length between 3 and 4 cm were maintained for 24 h in vermiculite at ψW of -0.03 (well hydrated), -0.65, -1.48 and -2.35 MPa (partially dry). Ten anatomical characteristics of differentiation and cell division in root regions were evaluated. Thickness of epidermis and protoderm diminished similarly in wild and domesticated beans growing at low substrate ψW (between -0.65 and -2.35 MPa). At the same time, parenchymatic cell area diminished by 71 % in the domesticated variety, but by only 32 % in the wild bean at -2.35 MPa. The number of cells in the cortex and the thickness of the xylem wall increased in both wild and domesticated beans at low substrate ψW; nevertheless, the effect was significantly lower in the wild bean. The number of xylem vessels increased in the cultivar (up to 40 %) while in the wild bean it decreased (up to 33 %). The diameter of xylem vessels and transverse root area diminished (15 and 57 %, respectively) in the cultivar, but in the wild common bean were not affected. Anatomical root characteristics and their modifications in both differentiation and cell division in root regions demonstrated that the wild bean reacted quite differently to substrate ψW than the domesticated common bean.     

Anatomy of Seedling Roots of Tropical Maize (Zea mays L.) Cultivars at Low Water Supply

Establishment of maize seedlings can be difficult at low soilmoisture content. Anatomy of root metaxylem vessels may influencethe capacity for water transport and respective genotypic differencesmight be useful for selection purposes. To test this, six tropicalmaize (Zea mays L.) cultivars were grown in large PVC tubescontaining a sandy substrate at 5% (M5) and 10% (M10) moisturecontents for 2 weeks. The percentage changes in root diametersdue to M5 was similar for most cultivars but differed for mainroot types. Root diameters were not consistently related tometaxylem structure, but in a few cases, thin roots had smallerdiameter metaxylem vessels. The M5 treatment reduced the numberof late metaxylem vessels of primary roots by about 0 to 20%,while effects on nodal roots were slight. Generally, the ratioof cross-sectional areas between late and early metaxylem vesselsincreased from primary to seminal and nodal roots. Within thecultivar Tuxpefio this ratio was much reduced by M5. A few cultivarsmaintained the combined cross-sectional areas of metaxylem vesselsat M5 in some main root types, but only one cultivar could achievethis for the total of cross-sectional areas of metaxylem vessels,calculated over all root axes, by increasing the number of seminaland nodal roots. These anatomical traits seemed to be mostlyconstitutive with limited response to an actual environment,but they could be decisive for the suitability of a cultivarto an environment with frequent water shortages during seedlingestablishment. Key words: Metaxylem vessels, water stress, tropical maize     

Hydraulic propagation of pressure along immature and mature xylem vessels of roots of Zea mays measured by pressure-probe techniques

Turgor pressure was measured in cortical cells and in xylem elements of excised roots and roots of intact plants of Zea mays L. by means of a cell pressure probe. Turgor of living and hence not fully differentiated late metaxylem (range 0.6–0.8 MPa) was consistently higher than turgor of cortical cells (range 0.4–0.6 MPa) at positions between 40 and 180 mm behind the root tip. Closer to the tip, no turgor difference between the cortex and the stele was measured. The turgor difference indicated that late-metaxylem elements may function as nutrient-storage compartments within the stele. Excised roots were attached to the root pressure probe to precisely manipulate the xylem water potential. Root excision did not affect turgor of cortical cells for at least 8 h. Using the cell pressure probe, the propagation of a hydrostatic pressure change effected by the root pressure probe was recorded in mature and immature xylem elements at various positions along the root. Within seconds, the pressure change propagated along both early and late metaxylems. The half-times of the kinetics, however, were about five times smaller for the early metaxylem, indicating they are likely the major pathway of longitudinal water flow. The hydraulic signal dissipated from the source of the pressure application (cut end of the root) to the tip of the root, presumably because of radial water movement along the root axis. The results demonstrate that the water status of the growth zone and other positions apical to 20 mm is mainly uncoupled from changes of the xylem water potential in the rest of the plant.Abbreviations and Symbols CPP cell pressure probe - EMX early metaxylem - LMX Late metaxylem - P_c cell turgor - P_r root pressure - RPP root pressure probe - t_1/2,c half-time of water exchange across a single cell - t_1/2 half-time of water exchange across multiple cells We thank Antony Matista for his expert assistance in the construction and modification of instruments. The work was supported by grant DCB8802033 from the National Science Foundation and grant 91-37100-6671 from USDA, and by the award of a Feodor Lynen-Fellowship from the Alexander von Humboldt-Foundation (Germany) to J.F.     

Effects of Root Pruning on Translocation of Photosynthates in Phaseolus vulgaris L

Root pruning increased the level of ethanol soluble sugars inred kidney bean plants (Phaseolus vulgaris L. ) grown in aeratednutrient solution. However, the concentration gradient of thesesugars down the stem and its translocation velocity remainedunchanged. Removal of 50% of the roots had no effect on thetotal photosynthates exported from source leaves but the finaldistribution pattern of photosynthates was altered; less movingtoward the upper plant parts, and accumulation occurring inthe lower stems. Translocation velocity of photosynthates towardthe upper plant parts was drastically reduced by root pruning. Key words: Phaseolus vulgaris, Photosynthate translocation, Root pruning     

Interspecific Grafts Demonstrate Root System Control of Leaf Water Status in Water-Stressed Phaseolus

We examined the importance and the mechanisms of the root systems'effect on leaf water status in two bean species: Phaseolus vulgarisL. cv. Redcloud (Pv) and P. acutifolius Gray MN cultivatedaccession 258/78 (Pa). Pa maintains a higher leaf water potential(₁) than Pv. We used reciprocal grafts between the two species.We grew four plants (one of each graft combination) in one potso they experienced the same soil water potential. Shoot genotypedetermined ₁ of well-watered plants. Root genotype determined₁ of the most stressed plants. Stressed Pa root systems increased₁ of Pv shoots by 0·1 MPa over Pv shoots on Pv roots.Pa roots did not maintain by affecting stomatal conductancenor by simply having more dry weight. Pa roots may have greaterhydraulic conductivity than Pv roots. Key words: Phaseolus acutifolius, Phaseolus vulgaris, leaf water potential, root-shoot communication     

Hydraulic properties of living late metaxylem and interactions between transpiration and xylem pressure in maize

A new approach to study dynamic interactions between transpiration and xylem pressure in intact plants is presented. Pressure probe measurements were preformed in living (immature) late metaxylem of maize roots rather than in adjacent mature xylem. This eliminated technical limitations related to the measurement of negative pressures. Water relations of single cells showed that turgor and volumetric elastic modulus were significantly larger in living metaxylem than in cortical cells; hydraulic conductivity was similar in both types of root cells. Increasing transpiration induced an immediate decrease of xylem pressure, and vice versa. Turgor in the living metaxylem could be continuously recorded for more than 1 h. The relationship between xylem pressure and transpiration yielded a root hydraulic resistance of 1.3 x 10⁹ MPa s m^-3. Control experiments indicated that the response of living xylem in the positive pressure range essentially paralleled that of mature root xylem in the negative range. In mature xylem, pressures as low as -0.55 MPa were recorded for short periods (several minutes). Several tests verified that the pressure probe was in contact with mature xylem during the measurements of tensions. The results demonstrate convincingly that transpiration generates an effective driving force for water uptake in roots, a central feature of the cohesion theory.Key words: Hydraulic conductivity, negative pressure, root development, turgor, water transport, Zea mays.      

Development of water conducting capacity in the root systems of young plants of corn and some other c4 grasses

Development of the primary and early nodal roots was studied in Zea mays L., Zea mexicana (Schrad.) Reeves & Mangelsd., Sorghum bicolor (L.) Moench., and Sorghum sudanese (Piper) Stapf. in relation to shoot development. In all the types studied all roots reached lengths of about 30 centimeters before the late metaxylem (LMX) was open, and young plants with total root lengths of around 100 centimeters had almost no open LMX. On average, corn seedlings with up to 36 square centimeters of leaf had no open LMX. The name “immature apices” is suggested for such long but not fully functional roots. In plants up to 50 days old a fairly constant proportion of less than half the total root length had open LMX. A pilot study of stomatal resistance on days of high evaporative demand suggested that young seedlings may show higher resistance than older plants in the afternoon. Estimates of longitudinal permeability of corn roots with only early metaxylem vessels open indicate very steep gradients of water potential would develop under such conditions.