Growth and Development of Young Roots of Barley (Hordeum vulgare L.) Genotypes

Barley (Hordeum vulgare L.) genotypes from countries with aMediterranean climate grown in temperature-controlled glasshousein nutrient solution to determine whether the co-ordinationof root branching and growth found by other workers appliedto a wider of up to 14 genotypes. There was substantial variationin the number of seminal axes produced by the genotypes, rangingfrom about seven for Hoshimasari and Swanneck to about fourfor Gerbel 'B'. The number of nodal axes was linearly relatedto the number of leaves and typically between one and two mainstemleaves were required before nodal axes appeared. There weresmall genotypic differences in the number of axes produced perleaf with values ranging from 1·5 to 2·3. Theproduction and growth of lateral roots were coordinated so thatthe mean length of laterals generally increased with time. Landraces(Arabic abiad and Arabic aswad) produced more lateral rootswith a faster rate extension compared with other genotypes.The length and number of primary and secondary lateral rootswere related linearly, but no genotypic differences in thisrelation were evident. Length of primary lateral roots increasedmore rapidly than that of secondary lateral roots throughoutthe three to five leaf stage. The ratio of root weight to totalplant weight decreased with time but there were only small differenceswithin this range of genotypes.Copyright 1995, 1999 AcademicPress Barley, seminal axes, nodal axes, primary lateral roots, relative extension rates, relative multiplication rates     

Germination and Early Growth of Plants Studied Using Neutron Radiography

Seed swelling, germination, root extension, lateral root initiationand shoot growth were studied in soils of different water contents,using non-destructive, serial neutron radiography. Seeds fromthree varieties of soya beans (Glycine max L.) and one varietyeach of maize (Zea mays L.) and vetch (Vicia sativa L.) wereused. The seeds germinated when they had increased in size bya certain amount, if germination is taken as the time when theradicle first appears. The rate at which roots and shoots extendalso depend on soil water content. Glycine max L., Vicia sativa L., Zea mays L., Soya bean vetch, maize, seed germination, root extension, lateral root initiation, neutron radiography     

Effect of Localized Anoxia on Phaseolus vulgaris L. Root Growth

Heterogeneity within the root environment results in differentialgrowth within root systems. The response of five Phaseolus vulgarisL. cultivars to non-uniform root aeration was evaluated. Threetreatments were applied to a split root system for a periodof 72 h. Treatments consisted of an aerated control, a non-aeratedcontrol (both halves non-aerated, using N₂). and localized anoxia(one-half the root system aerated and the remaining half subjectedto N₂). Shoot and root growth were reduced in the anoxic controlbut not in the aerated control or localized anoxia treatment. Root growth was greatest in the aerated portion of the localizedanoxia treatment for all genotypes. Contributions of the rootcomponents to the compensatory responses differed dependingon the plant cultivar examined. The growth of branched and lateralroots present before the treatment period increased by 65% inline 31908. A 50% increase in the growth of lateral roots whichemerged during the treatment period occurred in another line(Swan Valley). Other genotypes responded in an intermediatemanner. These observations indicate differences in cultivarresponses to localized soil stress. Key words: Phaseolus vulgaris L., Anoxia, Root growth     

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     

Effects of the Temperature of the Rooting Zone on the Growth and Development of Roots of Potato (Solanum tuberosum)

In solution culture the effect of optimal (20 °C) and supra-optimal(30 °C) root-zone temperatures on root growth and root morphologyof six potato clones (Solanum tuberosum L spp) was studied Growthwas compared with sweet potato (Ipomoea batatas L) and cassava(Manihot esculenta Crantz) Significant genotypical differencesin the responses of potato roots to supra-optimal temperatureswere observed, indicating the potential for selecting heat tolerantpotato clones In both heat tolerant and heat sensitive clones,the size of the root system was reduced by supra-optimal root-zonetemperature This was principally a result of decreased numberand length of lateral roots The first symptom of heat damagewas a reduction in the rate of cell division, followed by cessationof root elongation Bending of the apical root-zone togetherwith the formation of root hairs on the inner (concave) andlateral roots on the outer (convex) side were other symptoms,these coincided with the loss of starch granules in the root-cap Potato, genotypical differences, root morphology, root growth, temperature, Solanum tuberosum L     

Simultaneous measurement of root force and elongation for seedling pea roots

A new method was developed to measure simultaneously, continuously,and non-destructively the elongation rate and the force exertedby the roots of seedlings grown in moist air. A pea (Pisum sativumL. cv. Helka) seedling was suspended inside a modified sampletube on one side of a pulley, with the tip of the radicle pushingon to a force transducer through a hole in the tube. The forceon the root tip was monitored by the force transducer and couldbe adjusted by adding or removing mass from the counterweighton the other side of the pulley. As the root grew, the sampletube was raised and the elongation of the root was monitoredusing a linear variable differential transformer (LVDT) attachedto the thread connecting the sample tube and counterweight.The changes in elongation rate were recorded which occurredin response to increases and decreases in the applied force.Forces of up to 125 mN were exerted on the root, correspondingto forces per unit final cross-sectional area (i.e. root growthpressures) of up to 0.1 MPa. As soon as the force on the root was changed there was a rapidreversible compression or extension of the root. Superimposedon this elastic/viscoelastic deformation, the root elongationrate slowed by more than 50% within 30 min of increasing theforce applied to the root by 100 mN. A similarly fast but smallerincrease in growth rate occurred when the force was removed.Both of these ‘fast’ responses were followed bya longer period of more gradual change in the root elongationrate over a period of 20 h or longer. Both ‘fast’and ‘slow’ responses may be explained in terms ofa modified Lockhart model of growth. The initial ‘fast’response of the root is probably due to the immediate changein the effective pressure (i.e. the turgor pressure minus theyield stress and external resisting pressure) available to drivecell elongation. The reason for the second slower adjustmentof the elongation rate is not known, but is probably due tosome combination of a decrease in the rate of cell productionand/or a stiffening of the cell walls in the longitudinal directionwith increasing mechanical resistance. The increase in rootdiameter in response to mechanical impedance decreased the rootgrowth pressure that the root exerted, but was associated witha slower root elongation rate. Key words: Compaction, mechanical impedance, penetration resistance, root diameter, soil strength     

Root growth and nitrate uptake by wheat (Triticum aestivum L.) following wetting of dry surface soil

The nitrate uptake capacity of surface roots of spring wheat(Triticum aestivum L. cv. Kulin) was investigated followingwetting of dry surface soil. Plants were grown to stem elongationstage with adequate watering at depth while the surface soilwas allowed to dry. Eight weeks after sowing, water or a ¹⁵N-nitratesolution was added to the surface soil to simulate rainfall.Root growth and nitrate uptake were measured up to 4 d afterwetting on plants with unconfined nodal root growth and on plantswith the majority of nodal roots confined within small vials.Prior to wetting, plants from both nodal treatments had seminalroots with collapsed cortices along the upper 10 cm and manyshort, viable lateral roots. Nodal roots, where present, wereonly a few cm long and unbranched. Only plants with unconfinednodal roots were able to take up nitrate within the 24 h beforeany new root growth. By 2 d after wetting there was significantgrowth of the seminal lateral roots, and rapid growth and branchingof nodal roots. From 2 d after wetting, plants with confinednodal roots also took up nitrate, presumably due to the growthof the seminal lateral roots. Hence it appears as though thenodal roots in the unconfined treatment could immediately takeup nitrate, but the seminal roots required new lateral rootgrowth to become active in nitrate uptake. The plants with confinednodal roots had a lower nitrate uptake than those with unconfinednodal roots 4 d after wetting, indicating that the seminal rootsystem was not able to compensate for lack of nodal roots. Insufficientnitrate was taken up after 4 d, by plants from either nodalroot treatment, to increase the shoot N concentration significantly. Key words: Triticum aestivum, nitrate uptake, drought, seminal roots, nodal roots     

Biomechanics of nodal, seminal and lateral roots of barley: effects of diameter, waterlogging and mechanical impedance

Background and aims

Biomechanical properties of cereal root systems largely control both resistance to root lodging and their ability to stabilise soil. Abiotic stresses can greatly modify root system growth and form. In this paper the effect of waterlogging and moderate mechanical impedance on root biomechanics is studied for both lateral roots and the main axes of barley.

Methods

Barley (Hordeum vulgare) plants were subjected to transient water-logging and moderate mechanical impedance in repacked soil columns. Roots were excavated, separated into types (nodal, seminal or lateral) and tested in tension to measure strength and elastic modulus.

Results

Water-logging and mechanical impedance substantially changed root system growth whilst root biomechanical properties were affected by waterlogging. Root strength was generally greater in thin roots and depended on root type. For example, seminal roots 0.4–0.6 mm in diameter were approximately seven times stronger and five times stiffer than lateral roots of the same diameter when mechanically impeded. Root sample populations typically exhibited negative power-law relationships between root strength and diameter for all root types. Mechanical impedance slowed seminal root elongation by approximately 50 % and resulted in a 15 % and 11 % increase in the diameter of in nodal and seminal roots respectively. Power-law relationships between root diameter and root biomechanical properties corresponded to the different root types. Coefficients for between root diameter, strength and elastic modulus improved when separated by root type, with R² values increasing in some roots from 0.05 to 0.71 for root strength and 0.08 to 0.74 for elastic modulus.

Conclusions

Moderate mechanical impedance did not influence the tensile strength of roots, but, waterlogging diminished the relationship between root strength and diameter. Separation of root type improved predictions of root strength and elastic modulus using power-law regressions.     

Relationship Between Lateral Root Primordia in Different Ranks

Comparisons with theoretical random distributions indicate thatthe longitudinal positional relationships between lateral rootsin different ranks are random in roots of Musa acuminata, Pistiastratiotes and i, species in which lateral root primordia arisein the root tip. In Potentilla palustris, where lateral rootprimordia arise further back in the root, there is a deficiencyof close spacings, which indicates that there is some interactionbetween ranks. Musa acuminata, Pistia stratiotes, Pontederia cordat, Potentilla palustris, roots, lateral roots, pattern     

Chlorsulfuron Reduces Extension of Wheat Root Tips in Low-zinc Solution Culture

Root tip extension was used as a measure of wheat root responseto exposure to the sulfonylurea herbicide chlorsulfuron. Plantspre-grown in low-zinc (0.2 µMZnHEDTA) solutions were placedin a perspex chamber with nutrient solution on both sides ofa partition separating the root tip from the rest of the plant.The root tip was exposed to different concentrations of chlorsulfuronand observations were made during 22 h. Increasing the concentrationof zinc in the solution around the root tip to 4 µMZnHEDTAdid not alter root tip extension in the absence of chlorsulfuron.Significant decreases in root growth after 22 h were obtainedwith concentrations of 120 µg chlorsulfuron l^-1and greater.Increasing the Zn concentration from 0.2 to 20 µMZnHEDTAin the nutrient solution around the root tip decreased controlroot growth but stimulated chlorsulfuron-treated roots to extendat the same rate as chlorsulfuron-free control plants. AddingZn and chlorsulfuron to the more mature root parts above theroot tip partition did not significantly influence root tipextension. It is concluded that chlorsulfuron inhibits wheatroot growth and that increased Zn concentrations can alleviateor prevent the deleterious effects of chlorsulfuron.Copyright1998 Annals of Botany Company Wheat,Triticum aestivumL., chlorsulfuron, root growth, zinc.     

Water deficit accelerates determinate developmental program of the primary root and does not affect lateral root initiation in a Sonoran Desert cactus (Pachycereus pringlei, Cactaceae)

Determinate root growth is an important adaptation feature for seedling establishment in some Cactaceae. We show that seedlings of Pachycereus pringlei have primary roots with a stable determinate developmental program. How water stress affects determinate root growth and lateral root development has not been studied. Here we address this question. Root growth was analyzed in plants growing in vitro under well-watered and water-deficient (created by polyethylene glycol) growth conditions. Under severe water stress roots terminated their growth earlier and the rate of growth was significantly decreased as a result of inhibition of both cell elongation and cell production. Under severe water stress the number of lateral roots and primordia per millimeter of primary root was 1.5-1.7 times greater than under well-watered conditions; however, the total number of lateral roots and primordia was the same under all conditions. Lateral roots resembled root spurs found in some Opuntioideae. Analysis of the dynamics of meristem exhaustion indicated that initial-cell activities are required for the maintenance of proliferation before meristem exhaustion. We conclude that lateral root formation is a stable developmental process resistant to severe water stress and that water stress accelerates the determinate developmental program of the primary root. Both of these features appear to be important for successful seedling establishment in a desert.     

Effects of soil mechanical impedance on root and shoot growth of Lolium perenne L., Agrostis capillaris and Trifolium repens L.

A method was used for applying a uniform mechanical impedanceto plant roots using sand packed at different bulk densitieswith depth within each growth cylinder. In a growth experiment,replicate cylinders were packed with sand to give the followingmechanical impedances: 0.25 MPa penetration resistance (negligibleimpedance), 1.40 MPa (moderate impedance) and 2.30 MPa (severeimpedance). Seedlings of Lolium perenne L., Trifolium repensL. and Agrostis capillaris were grown for 23 d in each impedancetreatment and effects on both roots and shoots were studied.Severe mechanical impedance affected both root and shoot growthrates for all three species resulting in smaller leaves andshorter roots. For the grasses, the root-to-shoot ratio at harvestwas the same for all the treatments, but a delay in the initiationof both shoots and roots was observed in the severe mechanicalimpedance treatment. The results are discussed in relation tothe possibility that roots penetrating the impeding treatmentsmay have caused signalling that kept shoot and root growth insynchrony. Differences observed in the response to mechanicalimpedance of T. repens compared to the other two species maybe a function of differences in the physiology between mono-and dicotyledonous species. Key words: Mechanical impedance, Lolium perenne, Trifolium repens, roots, shoots     

Seminal Root Growth in Sorghum (Sorghum bicolor) Under Allelopathic Influences from Residues of Taro (Colocasia esculenta)

The length of the seminal root (SR) axis and the number andlength of lateral roots (LRs) of sorghum (Sorghum bicolor Moench)were markedly inhibited by taro [Colocasia esculenta (L.) Schott]residues incorporated into a sand growing medium. The sand profilewas divided equally into zones with and without residues. Productionand elongation of the first-order LRs of the SR axis facingthe zone containing taro residues were severely suppressed.On the side facing the zone that was free of residues, productionand elongation of LRs was not inhibited. SR and LR growth wasdrastically impaired and many plants were killed when taro residueswere incorporated in large amounts into the uppermost 2 cm ofthe growing medium. The activity of the allelopathic substancesin the root zone appeared to be location-specific. Sorghum bicolor, seminal root, lateral root, Colocasia esculenta, taro, taro residues, allelopathic substances, root growth     

The Effect of Light Intensity and Relative Humidity on Growth Rate and Root Respiration of Plantago lanceolata and Zea mays

Plants of Plantago lanceolata L. and Zea mays L., cv. ‘Campo’were grown at two levels of light intensity. Especially in theroots, the rate of dry matter accumulation decreased at lowlight intensity. The carbohydrate content of both roots andshoots of P. lanceolata was not affected by light intensity.The relative contribution of SHAM¹-sensitive respiration, thealternative chain, to total root respiration of both P. lanceolataand Z. mays, was not affected by light intensity during thedaytime. The alternative pathway was somewhat decreased at theend of the dark period, but not in the root tips (0–5mm) where it still contributed 56% in respiration. It was, therefore,concluded that photosynthesis is not a major factor in regulationof root growth in the species investigated. To see whether the effect of light intensity on root growthrate was via transpiration, plants of Z. mays were grown atdifferent air humidities. Both high humidity and low light intensityaffected the root morphology in such a way that the distancebetween the apex and the first laterals on the primary rootaxis increased. It is suggested that this effect on root morphologyis due to transpiration and the subsequent removal of root-producedinhibitors of lateral root growth; although light intensityalso affected the rate of dry matter accumulation of roots andthe rate was not affected by the humidity of the air. It is,therefore, concluded that the effect of light intensity on therate of dry matter accumulation of roots of Z. mays is not viaan effect on transpiration.     

Symplastic communication between primary and developing lateral roots of Arabidopsis thaliana

The functional symplastic connections between primary and developinglateral roots of Arabidopsis were studied non-invasively usingconfocal laser scanning microscopy (CLSM), following ester-loadingof the phloem with carboxyfluorescein (CF). Prior to the formationof lateral primordia in the pericycle, the phloem of the primaryroot behaved as an isolated conducting domain. However, thedifferentiation of phloem connector elements within the dividingpericycle allowed the rapid establishment of intercellular communicationbetween the phloem and the cells of the lateral primordium.This communication was often established prior to the completeemergence of the lateral root from the parent root. Shortlyafter its emergence, functional conducting phloem became differentiatedwithin the developing lateral root. A progressive isolationbetween the phloem and surrounding cells at the base of thelateral root was observed as the lateral continued to grow;the new phloem conducting CF to the elongation zone where itwas unloaded symplastically from the protophloem into surroundingcells of the cortex and stele, a feature mirroring the patternfound near the apex of growing primary roots. Anomalous patternsof intercellular communication were found which indicated thatpreviously functional symplastic pathways may have become sealedoff following the emergence of some of the lateral roots. Key words: Arabidopsis, carboxyfluorescein, confocal laser scanning microscopy (CLSM), intercellular transport, lateral roots, phloem (unloading), symplast     

Early growth of Brazilian pine (Araucaria angustifolia[Bertol.] Kuntze) in response to soil compaction and drought

Soil compaction leads to changes in soil physical properties such as density, penetration resistance and porosity, and, by consequence, affects root and plant growth. The initial growth of Brazilian pine is considered as being more affected by soil physical than chemical conditions, and the presence of a well-developed tap root system has been associated with this fact. A greenhouse experiment was conducted in order to evaluate the impact of soil compaction on the growth of Brazilian pine seedlings and on their susceptibility to a simulated drought period. In the first phase of the experiment, the effects of three levels of soil compaction on root morphology and plant growth were examined. Soil cylinders were artificially compacted in PVC tubes. Pre-germinated seeds were planted, and 147 days later 10 plants from each treatment were harvested for analysis. Higher values of soil density were associated with a shorter and thicker tap root. Growth of lateral roots and shoots remained unaffected at this stage. In the second phase, half of the plants (12) in each compaction treatment were drought-stressed by withholding water for a period of 77 days. Increased soil compaction again resulted in reduced length and increased diameter of the main tap root. This time, the effects were also extended to the lateral roots. Shoot extension growth and overall plant mass, however, increased with soil compaction. This greater mass accumulation in plants growing under increased soil compaction may be attributed to a more intimate contact between roots and soil particles. Drought stress reduced both root and shoot growth, but root mass was more negatively affected by drought stress in plants growing under high levels of soil compaction. Future investigations on the effects of soil compaction on the initial growth of Brazilian pine should include a wider range of compaction levels to better establish the relationship between soil physical parameters and plant growth.     

Interaction between rhizobia and potato tissues10

Several cultivars of Solanum tuberosum L., the potato, weregrown on tissue culture media and their roots inoculated withstrains of rhizobia known to infect legumes at root junctionsor between epidermal cells. Infection incidence and severityshowed considerable cultivar/bacterial strain interaction. Bacteriaspread through intercellular spaces and invaded cells in a non-structuredway: some infections penetrated to the root xylem. There wasno evidence that potato root cells produced nod-inducing factorsand nitrogenase activity was not detected. In some host/rhizobialcombinations outgrowths were formed on roots. These varied fromloose infected callus tissue at the junctions of lateral roots,to modified lateral roots of limited growth which showed somecolonization by rhizobia. Key words: Solarium tuberosum L, potato, glycoprotein, immunogold labelling, rhizobia, flavonoids     

Root and Shoot Growth of Plants Treated with Abscisic Acid

Young seedlings of Capsicum annum L., Commelina communis L.and maize (Zea mays L.) were subjected to a mild water-stressingtreatment and/or treated with abscisic acid (ABA). Plants rootedin soil received a soil-drying treatment and their leaves weresprayed with a 10–⁴ M solution of ABA. Plants grown insolution culture were stressed by the addition of polyethyleneglycol (PEG) to the rooting medium and ABA was also added tothe rooting medium, either with or without PEG. The effectsof both treatments on the growth of roots and shoots and theultimate root: shoot dry weight ratio were very similar. Shootgrowth was limited both by water stress and by ABA application;while there was some evidence that mild water stress and/orABA application may have resulted in a stimulation of root growth.More severe water stress reduced the growth of roots but theoverall effect of stress was to increase the ratio of rootsto shoots. Capsicum annum L., Commelina communis L., Zea mays L., water stress, abscisic acid     

A comparative study of the response of the roots and shoots of sunflower and maize to mechanical stimulation

Despite numerous studies of the effects of mechanical stimulationon plant shoots, the response of roots to mechanical stimulationhas largely been neglected. In this study the effects of shootflexure on the morphology and mechanics of two contrasting speciesof herbaceous angiosperm, growing in a glasshouse were compared:maize (Zea mays), a monocot; and sunflower (Helianthus annuusL.) a dicot. Mechanical stimulation affected the root more than the shootcomponents. Root systems of mechanicallystressed sunflowershad a greater angle of spread and increased root number. Aswell as large morphological and weight effects, with increasesover the control of 33% in the length of rigid root and 38%in the dry weight of lateral roots, in sunflowers, there werealso mechanical effects. In both species roots of flexed plantswere more rigid, stronger and composed of stiffer material andtheir root systems also provided greater anchorage strength.In contrast, there was only a small reduction in shoot weightand shoot height in flexed plants and no effects on mechanicalproperties. There were differences in behaviour between species; maize rootmorphology responded less than that of sunflowers to mechanicalstimulation. The basal diameter of roots increased by only 8%compared with 16% in sunflowers, though the roots of both speciesshowed similar increases in material stiffness. This differenceis related to the lack of secondary thickening in the monocotscompared with the dicot sunflowers. Key words: Thigmomorphogenesis, Helianthus annuus L., Zea mays, anchorage, lodging