Root Systems and Fractals: How Reliable are Calculations of Fractal Dimensions?

Fractal dimensions of the root systems of Betula populifoliaand Betula alleghaniensis grown at two levels of soil nutrientswere calculated from tracings of root systems (Fitter and Stickland,1992a) and from roots removed from the soil using a pin-board(Tatsumi, Yamauchi and Kono, 1989). The fractal dimensions obtainedfrom these two methods showed similar qualitative patterns betweenspecies and nutrient treatments, but were not significantlycorrelated with one another. These observations suggest thatcurrent methods used for estimating the fractal dimensions ofplant root systems are not yet reliable. It is suggested thatthe calculation of fractal dimensions needs to be carried outin three dimensions in order to appropriately quantify the fractaldimensions of plant root systems.Copyright 1994, 1999 AcademicPress Betula populifolia Marsh (gray birch), Betula alleghaniensis Britton (yellow birch), fractal dimension, root system     

The Rate of Growth and Partitioning of Assimilates in Young Grass Plants: A Mathematical Model

A model describing the increase in weight with time of younggrass plants is formulated. The parameters are the relativegrowth rates of the root and shoot systems; k, the ratio ofthe relative growth rate of the root system relative to thatof the shoot system; b, the weight of the root system when thatof the shoot system is unity, and u the rate of increase inweight of the whole plant per unit of shoot system per unitof time, k and b are the constants in the allometric formula,r = bs^k where r and are the weights of the root and shoot systems.The model enables the effect of changes in the distributionof assimilates between the root and shoot systems upon the rateof growth of the plant to be assessed. Data from a number ofexperiments are analysed in this manner and the significanceof the results discussed.     

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     

Genotypic, Developmental and Drought-Induced Differences in Root Hydraulic Conductance of Contrasting Sugarcane Cultivars

Hydraulic properties of entire root systems and isolated rootsof three contrasting sugarcane clones were evaluated using transpiration-induceddifferences in hydrostatic pressure across intact root systems,root pressure-generated xylem sap exudation, and pressure-fluxrelationships. Regardless of the measurement technique employed,the clones were ranked in the same order on the basis of theirleaf area–specific total root system hydraulic conductance(C_root). All methods employed detected large developmental changesin G_rootroot with maximum values occurring in plants with approximately02 m₂ total leaf area. Genotypic ranking according to G_root,was reflected as a similar ranking according to root length-specifichydraulic conductance (L) of individual excised roots. Genotypicdifferences in G_root and L were consistent with anatomical characteristicsobserved in individual roots. Patterns of G_root, during soildrying and following re-irrigation suggested that the declinein G_root, observed during soil drying occurred within the rootsrather than at the soil–root interface and may have beencaused in part by xylem cavitation in the roots. Key words: Root hydraulic conductance, Saccharum spp, transpiration, root pressure, pressure-flux     

Oxygen Transport in the Salt Marsh Genus Puccinellia with Particular Reference to the Diffusive Resistance of the Root--Shoot Junction and the use of Paraffin Oil as a Diffusive Barrier in Plant Studies

Results are presented which contradict previous reports thatin the graminaceous genus Puccinellia the root-shoot junctionhas a high diffusive resistance which severely restricts oxygentransfer from shoot to root. No such restriction was evidentin P. maritima, P. festuciformis var.festuciformis, and P. festucfformisvar.intermedia(= P. peisonis) and the plants produced well-developedroot systems in permanently flooded soils. The evidence obtainedsupports the view that, whether the root systems are whollyor only partially submerged by soil flooding the shoot is themajor source or entry point for oxygen required by the roots;also, that oxygen passes more readily from shoot to root thanit does through the wall cells of the root base. It is pointed out that paraffin oil is an undesirable productto employ as a diffusion barrier in studies of root aeration. Key words: Puccinellia, Oxygen transport, Diffusive resistance, Polarography     

New Gene Crt(Curly Roots) Controlling Pea (Pisum sativum L.) Root Development

Using ethyl methane sulfonate (EMS) treatment of the seeds ofline SGE, a new mutant of pea (Pisum sativum L.) with alterationsin root development was obtained. The mutant phenotype dependson the density of the growth substrate: on sand (a high densitysubstrate) the mutant forms a small compact curly root systemwhereas on vermiculite (a low density substrate) differencesbetween the root systems of the mutant and wild type plantsare less pronounced. Genetic analysis revealed that the mutantcarries a mutation in a new pea gene designedcrt (curly roots).Gene crt has been localized in pea linkage group V. The mutantline named SGEcrt showed increased sensitivity to exogenousauxin and an increased concentration of endogenous indole-3-aceticacid (IAA) in comparison with the wild type line SGE. Copyright2000 Annals of Botany Company Pisum sativum L., root development, garden pea mutant, curly roots, auxin, environmental stimulus response     

A Comparison of Dry Matter Partitioning in Seminal and Adventitious Plants of Trifolium repens L

Trifolium repens has two types of root, one derived from theseed and the adventitious roots derived from the stolon nodes.It has been suggested that these two systems have differentpotentials for supporting growth. This paper presents a comparisonof plants grown on single seedling or adventitious roots anddemonstrates that although the shoot: root ratios for the twotypes differ this may be explained by differing shoot morphologies.Comparison of the lamina: root ratios for the two types of plantproduced no statistically significant differences and it isproposed that the two types of root system do not differ intheir relationship with leaf growth. A mechanism for large diameter‘tap’ root formation is suggested. white clover, Trifolium repens L., adventitious roots, seedling roots, shoot: root ratio     

Root system growth and nodule establishment on pea (Pisum sativum L.)

Development of the root system, appearance of nodules, and relationshipsbetween these two processes were studied on pea (Pisum sativumL., cv. Solara). Plants were grown in growth cabinets for 4weeks on a nitrogen—free nutrient solution inoculatedwith Rhizobium leguminosarum. Plant stages, primary root length,distance from the primary root base to the most distal first-orderlateral root, and distance from the root base to the most distalnodule, were recorded daily. Distribution of nodules along theprimary root and distribution of laterals were recorded by samplingroot systems at two plant stages. Primary root elongation ratewas variable, and declined roughly in conjunction with the exhaustionof seed reserves. First-order laterals appeared acropetallyon the primary root. A linear relationship was found betweenthe length of the apical unbranched zone and root elongationrate, supporting the hypothesis of a constant time lag betweenthe differentiation of first-order lateral's primordia and theiremergence. Decline of the primary root elongation rate was precededby a reduction in density and length of first-order laterals.Nodules appeared not strictly but roughly acropetally on theprimary root. A linear relationship was found between the lengthof the apical zone without nodule and root elongation rate,supporting the hypothesis of a constant time lag between infectionand appearance of a visible nodule. A relationship was foundbetween the presence/absence of nodules on a root segment andthe root elongation rate between infection and appearance ofnodules on the considered root segment. Regulation of both processesby carbohydrate availability, as a causal mechanism, is proposed. Key words: Pisum sativum L, root system, nodules     

Hydraulic and osmotic properties of oak roots

Hydraulic and osmotic properties of root systems of 2.5–8-months-oldoak seedlings (Quercus robur and Q. petraea) were measured usingthe root pressure probe. Root pressures of excised roots rangedbetween 0.05 and 0.15 MPa which was similar to values obtainedfor herbaceous species. Root hydraulic conductivity (Lp_r; perunit of root surface area) was much larger in the presence ofhydrostatic than in the presence of osmotic pressure differencesdriving water flow across the roots. Differences were as largeas a factor of 20 to 470. Roots of the young seedlings of Q.robur grew more rapidly than those of Q. petraea and had a hydraulicconductivity which was substantially higher. Nitrogen nutritionaffected root growth of Q. robur more than that of Q. petraea,but did not affect root Lp_r of either species. For Q. robur,Lp_r decreased with root age (size) which is interpreted by aneffect of suberization during the development of fine roots.Root hydraulic conductance remained constant for both species.For Q. robur, this was due to the fact that the overall decreasein Lp_r was compensated for by an increase in root surface area.Root reflection coefficients (_sr) were low and ranged between_sr=0.1 and 0.5 for solutes for which cell membranes exhibitreflection coefficients of virtually unity (salts, sugars etc.).Solute permeability was small and was usually not measurablewith the technique. When root systems were attached to the rootpressure probe for longer periods of time (up to 10d), solutepermeability increased due to ageing effects which, however,did not cause a general leakiness of the roots as Lp_r decreased.Hence, values were only used from measurements taken duringthe first day. Transport properties of oak roots are comparedwith those recently obtained for spruce (Rdinger et al., 1994).They are discussed in terms of a composite transport model ofthe root which explains low root _sr at low solute permeabilityand reasonable rootLp_r The model predicts differences betweenosmotic and hydraulic water flow and differences in the transportproperties of roots of herbs and trees as found. Key words: Composite transport, hydraulic conductivity, nitrogen nutrition, Quercus, reflection coefficient, root transport, water relations     

The Vascular Pattern of a Rhizomatous Ginger (Alpinia speciosa L. Zingiberaceae). 2. The Rhizome

The vascular system in the underground rhizome of Alpinia speciosaL. (Zingiberaceae) is seen to be arranged in three distinctzones. (1) An inner system of ‘scattered’ vascularbundles which serial cinematography reveals to have an axialpattern conforming to the basic ‘palm’ configuration(a system of upwardly branching leaf traces with interconnections).(2) An intermediate zone comprising a thin perforated cylinderof anastomosing vascular strands having direct contact withboth roots and inner system bundles. (3) An outer system offreely-anastomosing vascular bundles. Connexion of outer andinner system occurs in the form of extensive bridging from innersystem leaf traces as they depart obliquely between the outersystem network. The interrelation of the three systems, plus root and branchinsertion, is illustrated by means of diagrammatic three-dimensionalreconstructions. The intermediate zone is intimately associatedwith root insertions and with the inner system, and is shownto obliviate potential bottlenecks at the point of lateral branchinsertion in this sympodial rhizome system. A comparison ismade with other monocotyle-donous vascular systems. Alpinia speciosa L., shell ginger, rhizome, vascular anatomy     

The Influence of Pine on the Form of Sitka Spruce Fine Roots

The form of fine roots of Sitka spruce (Picea sitchensis Bong.Carr.) when grown in immediate proximity to roots of Scots pine(Pinus sylvestris L.) and when grown separately was comparedusing split-root systems. When spruce roots were intimatelymixed with pine roots the mean length of individual spruce lateralswas significantly greater, while the length: weight ratio andnumber of root tips: weight ratio were smaller than when grownalone. A similar alteration in growth strategy was achievedby direct addition of mineral nitrogen. Key words: Fine root form, pine-spruce interactions     

A method for measuring xylem hydraulic conductance and embolism in entire root and shoot systems

Current methods for determining the influence of xylem cavitationon hydraulic conductance are limited to unbranched stem or rootsegments with hydraulic conductances above c. 2 mmol s^–1MPa^–1. Lower conductances and/or highly branched systemsare encountered in seedlings, arid-land shrubs, herbs, and distalportions of shoot and root systems of trees. In order to quantifythe hydraulic impact of cavitation in such systems, existingtechniques have been modified. Branched shoot or root systemswere prepared for measurement by removal of leaves, or roottips, respectively. The shoot or root system was enclosed ina vacuum chamber with the proximal end protruding and suppliedwith perfusing solution. Flow through the xylem was inducedby chamber vacuum. Hydraulic conductance was determined fromthe slope of the flow rate versus pressure relationship. Xylemembolism was quantified from the increase in hydraulic conductancefollowing high pressure (100 kPa) perfusion of solution throughthe plant. Examples are provided of the application of the methodto cavitation studies in the cold desert shrub Artemisia tridentata. Key words: Hydraulic conductance, xylem cavitation, embolism, whole root/shoot system     

Responses of young trees to wind and shading: effects on root architecture

Two wind tunnels were designed to detect influences of windon the development of the root systems of young trees: Piceasitchensis and Larix decidua were grown in the first and L.decidua only in the second. In the second experiment, the taproot of each L. decidua seedling was removed in order to mimicthe formation of a shallow root-plate. Responses of shoot growthto wind stress were small, although uneven irradiance levelsresulted in asymmetric growth of both shoots and roots; thedistribution of root biomass around the tree was related spatiallyto that of shoot growth In both experiments there was an increase in the number of largewindward and leeward roots in both species. In the first experiment,the sum of the crosssectional area (     

Aluminium and the Hydraulic Conductivity of Quercus rubra L. Root Systems

Two hydroponic experiments were conducted to determine the effectsof brief and prolonged AI³⁺ exposures on the hydraulic conductivity(Lp) of northern red oak (Quercus rubra L.) root systems. RootLp was determined using the pressure chamber method of Fiscus(1977). In the first experiment, 28- to 40-d-old seedlings weretreated for 4 d with complete nutrient solutions containingone of three Al concentrations (0.04, 1.85 or 3.71 mol m^–3)and either 0 or 50 mmol m^–3 P. Neither Lp nor daily transpirationwas affected by treatment. In Experiment II, seedlings were grown for 48–63 d incomplete solutions containing one of three Al concentrations(0, 0.75 or 2.00 mol m^–3) and either 10 or 250 mmol m^–3Ca. Lp and leaf area to root length ratio (LA/RL) were reducedwhen (AI³⁺/ Ca²⁺), the solution activity ratio, was 2.9 andhigher. Lp and LA/RL were also negatively correlated with Alconcentration and Al/Ca concentration ratio in the roots. Lpwas positively correlated with LA/RL in both experiments. Itis unclear whether Lp in the second experiment was reduced directlyby solution and root chemistry or whether Lp changed in responseto altered leaf/root balance. Key words: Al phytotoxicity, Al x Ca interaction, Quercus rubra, root hydraulic conductivity     

Morphological, Anatomical and Physiological Responses Related to Differential Shoot vs. Root Growth Inhibition at Low Temperature in Spring and Winter Wheat

Several morphological, anatomical and physiological changesand their relationship with differential root vs. shoot growthinhibition at low temperature (5°C) were studied in springand winter wheat cultivars. Root:shoot ratios, expressed eitheras a function of root and shoot fresh weight or as a functionof root and leaf areas, increased at low temperature and thisincrement was more pronounced in spring cultivars than in winterones. Although winter cultivars developed relatively smallerroot systems at 5°C, this characteristic was counterbalancedby a lower stomatal frequency and increased thickness of epidermalcell walls in leaves unfolded at this temperature, relativeto spring cultivars. Likewise, at 5°C a decrease in theosmotic potential of shoots and roots was observed in parallelwith sugar accumulation; this decrease was more marked in wintercultivars. These results indicate a differential morpho-anatomicaland physiological plasticity of winter and spring cultivarsduring development at low temperature. The possible associationbetween these changes and plant water economy at low temperaturesis discussed. Copyright 2001 Annals of Botany Company Spring wheat, winter wheat, Triticum aestivum, low temperature, root:shoot ratio, root surface area, stomatal frequency, osmotic potential     

Comparisons of the Respiratory Effluxes of Nodules and Roots in Six Temperate Legumes

The specific respiration rates of nodulated root systems, ofnodules and of roots were determined during active nitrogenfixation in soya bean, navy bean, pea, lucerne, red clover andwhite clover, by measurements on whole plants before and afterthe removal of nodule populations. Similar measurements weremade on comparable populations of the six legumes, lacking nodulesbut receiving abundant nitrate-nitrogen, to determine the specificrespiration of their roots. All plants were grown in a controlled-environmentclimate which fostered rapid growth. The specific respiration rates of nodulated root systems ofthe three grain and three forage legumes during a 7–14-dayperiod of vegetative growth varied between 10 and 17 mg CO₂g^–1 (dry weight) h^–1. This mean value consistedof two components: a specific root respiration rate of 6–9mg CO₂ g^–1 h^–1 and a specific nodule respirationrate of 22–46 mg CO₂ g^–1 h^–1. Nodule respirationaccounted for 42–70 per cent of nodulated root respiration;nodule weight accounted for 12–40 per cent of nodulatedroot weight. The specific respiration rates of roots lackingnodules and utilizing nitrate nitrogen were generally 20–30per cent greater than the equivalent rates of roots from nodulatedplants. The measured respiratory effluxes are discussed in thecontext of nitrogen nitrogen fixation, nitrate assimilation. Glycine max, Phaseolus vulgaris, Pisum sativum, Medicago sativa, Trifolium pratense, Trifolium repens, soya bean, navy bean, pea, lucerne, red clover, white clover, nodule respiration, root respiration, fixation, nitrate assimilation     

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.     

Episodic Growth and Relative Shoot: Root Balance in Loblolly Pine Seedlings

Leaf, root and stem systems of loblolly pine seedlings are characterizedby a seasonal periodicity in growth, during which they alternatein spurts of activity. Despite this periodicity, the allometriccoefficient describing the ratio of the relative growth ratesof leaf to root remains constant for at least the first twoyears of development. In part, constancy results from the inabilityof variation in the allocation of growth increment during briefperiods to change a pre-existing structure accumulated overthe life of the seedling. In addition, alternating periods ofleaf, root and stem growth may represent the action of feedbackmechanisms which operate to maintain an adaptive balance betweenorgan systems. Pinus taeda L., loblolly pine, allometric coefficient, homeostatic control of growth