Root Growth and Water Uptake by Maize Plants in Drying Soil

Sharp, R. E and Da vies, W. J. 1985. Root growth and water uptakeby maize plants in drying soil.— J. exp. Bot. 36: 1441–1456. The influence of soil drying on maize (Zea mays L.) root distributionand use of soil water was examined using plants growing in thegreenhouse in soil columns. The roots of plants which were wateredwell throughout the 18 d experimental period penetrated thesoil profile to a depth of 60 cm while the greatest percentageof total root length was between 20–40 cm. High soil waterdepletion rates corresponded with these high root densities.Withholding water greatly restricted root proliferation in theupper part of the profile, but resulted in deeper penetrationand higher soil water depletion rates at depth, compared withthe well watered columns. The deep roots of the unwatered plantsexhibited very high soil water depletion rates per unit rootlength. Key words: Maize, roots, water deficit, soil water depletion     

Does ABA in the Xylem Control the Rate of Leaf Growth in Soil-Dried Maize and Sunflower Plants?

Maize (Zea mays L.) and sunflower (Helianthus annuus L.) plantswere grown in large volumes of soil and leaf growth rate wasmonitored on a daily basis. Half the plants were given a soildrying treatment and when they showed a significant restrictionof growth rate (compared to both their daily growth rate beforedrying and the average growth rate of well-watered plants onthe same day), leaf water relations were measured and xylemsap was extracted using several techniques. There was a significant negative log-linear relationship betweenthe rate of leaf growth and the concentration of ABA in thexylem for both species. There was no clear relationship betweenleaf growth rate and leaf water potential or turgor for eitherspecies. Assessment of different methods for sampling xylemsap suggests that exudates collected from stem stumps or samplescollected by pressurizing the whole root system are suitablefor estimating ABA concentration in xylem, at least with largeplants of maize or sunflower, provided the first few hundredcubic millimetres of collected sap are used for the assay. Centrifugationof sections of stems resulted in dilution of ABA in the xylemsap with sap squeezed from parenchyma tissue. This is because,at least in plants subjected to mild soil drying, the concentrationof the ABA in the xylem is far higher than that in the cellsap of stem tissue. Results support the proposal that ABA plays a major role asa chemical signal involved in the root-to-shoot communicationof the effects of soil drying. The non-hydraulic restrictionof leaf growth by a chemical signal can be explained by theextra root-sourced ABA in the xylem and may be an importantcomponent of the modification of growth and development whichresults from prolonged soil drought. Key words: Soil drying, ABA, leaf growth, Zea mays L., Helianthus annuus L.     

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     

Simulating Growth and Root-shoot Partitioning in Prairie Grasses Under Elevated Atmospheric CO2and Water Stress

We constructed a model simulating growth, shoot-root partitioning,plant nitrogen (N) concentration and total non-structural carbohydratesin perennial grasses. Carbon (C) allocation was based on theconcept of a functional balance between root and shoot growth,which responded to variable plant C and N supplies. Interactionsbetween the plant and environment were made explicit by wayof variables for soil water and soil inorganic N. The modelwas fitted to data on the growth of two species of perennialgrass subjected to elevated atmospheric CO₂and water stresstreatments. The model exhibited complex feedbacks between plantand environment, and the indirect effects of CO₂and water treatmentson soil water and soil inorganic N supplies were important ininterpreting observed plant responses. Growth was surprisinglyinsensitive to shoot-root partitioning in the model, apparentlybecause of the limited soil N supply, which weakened the expectedpositive relationship between root growth and total N uptake.Alternative models for the regulation of allocation betweenshoots and roots were objectively compared by using optimizationto find the least squares fit of each model to the data. Regulationby various combinations of C and N uptake rates, C and N substrateconcentrations, and shoot and root biomass gave nearly equivalentfits to the data, apparently because these variables were correlatedwith each other. A partitioning function that maximized growthpredicted too high a root to shoot ratio, suggesting that partitioningdid not serve to maximize growth under the conditions of theexperiment.Copyright 1998 Annals of Botany Company plant growth model, optimization, nitrogen, non-structural carbohydrates, carbon partitioning, elevated CO₂, water stress,Pascopyrum smithii,Bouteloua gracilis, photosynthetic pathway, maximal growth     

Comparison of Osmotic Adjustment Responses to Water and Temperature Stresses in Spring Wheat and Sudangrass

This study explores the mechanisms of osmotic adjustment bycomparing the growth of spring wheat and sudangrass, which exhibitdifferent degrees of osmotic adjustment, under soil water andtemperature stresses. Leaf water potential ( ₁), osmotic potential(), and rate of leaf area growth of spring wheat and sudangrassseedlings were measured at combinations of five soil water potentials,from -0·03 to -0·25 MPa, and six root temperatures,from 14 to 36°C. Spring wheat exhibit little osmotic adjustment.The leaf osmotic potential was not affected by either soil wateror root temperature stress. Osmotic potential of sudangrassdecreased in parallel with the decreasing leaf water potentialas a result of osmotic adjustment. As soil water potential decreasedfrom -0·03 to -0·25 MPa, the rates of growth andphotosynthesis of spring wheat both decreased by about 30%.For sudangrass with the same range of soil water potential,the photosynthesis rate decreased by only 10% while the leafarea growth rate decreased by 49%. We introduce a dimensionlessindex (R) to quantify the degree to which environmental stressesalter the balance between production of photosynthates and theiruse for growth. The index, R, is equal to 1 when stress reducesgrowth and photosynthesis by the same degree, i.e. the balancebetween production and consumption of photosynthate is not disturbed.R is smaller than 1 when growth is reduced more than photosynthesis.R was equal to 1 for spring wheat where there was no osmoticadjustment. For sudangrass, R decreased from 1 to 0·25as osmotic potential decreased from -1·10 to -1·63MPa. These findings lead to the hypothesis that osmotic adjustmentcould result from an imbalance between production, consumptionand translocation of photosynthates under stressed conditions.Copyright1993, 1999 Academic Press Osmotic adjustment, water stress, root temperature     

Dry Matter and Water Import Rates in the Tomato Fruit: a Model Incorporating the Changes in Sap Viscosity and Osmotic Potential with Temperature

The change in water import rate in tomato fruit was modelledby incorporating into a previously-published model the changesin sap viscosity and osmotic potential into fruit with temperature.An experimental relationship between water and dry matter importrates was used to compare the model to dry matter import ratesin fruit measured by Walker and Thornley (Annals of Botany 41:977-985, 1997) at different temperatures. The effect of temperatureon the water import rate, calculated from the model, was alsocompared with the effect of temperature on the fruit growthrate measured by Pearce, Grange and Hardwick (Journal of HorticulturalScience 68: 1-11 and 13-23, 1993). The model accounted for alarge part of these temperature effects. It was concluded that resistances in sap transfer pathways inthe tomato fruit could be due to viscosity. These results supported,on the one hand, the hypothesis that the progressive decreaseof water import rate during fruit growth could result partlyfrom the progressive increase in transfer pathway length, and,on the other hand, the hypothesis that the ratio between waterand dry matter import rates could depend on flow conditionsin transfer pathways. The equations of the model could be usedto simulate tomato fruit growth, mass and dry matter contentin relation to fruit size, to nutrient solution salinity andto fruit temperature.Copyright 1995, 1999 Academic Press Dry matter, fruit, growth, model, resistance, sap, temperature, tomato, transfer, viscosity, water     

Development of Golgi Apparatus in the Root Cap Cells of Maize (Zea mays L.) as Affected by Compacted Soil

Effects of soil mechanical impedance on the development of Golgiapparatus in the root cap cells of maize were studied undercontrolled soil-water conditions Heavily compacted soil (bulkdensity = 1.50 g cm^–2) had 3.3 to 3.4 times greater mechanicalimpedance than control soil (bulk density = 1.33 g cm^–3),but their oxygen diffusion rates were not significantly differentThe number of dictyosomes and the number and area of secretoryvesicles per unit area of tangentially sub-peripheral root capcells in the heavily compacted soil increased compared to thosein the control These results suggest that secretory activityof the root cap cells is promoted by soil mechanical impedance Dictyosome, Golgi apparatus, maize, mucilage, root cap, secretory activity, secretory vesicles, soil mechanical impedance, Zea mays L     

Effect of Soil pH on Growth and Cation Deposition in the Root Tip of <Emphasis Type="BoldItalic">Zea mays</Emphasis> L.

Abstract The effects of sandy soil pH on the distribution of growth velocities and on cation concentrations and deposition rates in root growth zones of Zea mays L. seedlings were investigated. The pH values of the rooting medium varied between 4.2 and 8.6 in sand culture (70% saturated) without external supply of nutrients. At all pH values, densities (in 7moles per g fresh weight) of potassium, magnesium, and calcium increased toward the root tip. Lower pH in the medium increased calcium tissue density fivefold and magnesium density 1.7-fold, whereas the density of potassium, the overall elongation rate, and the growth velocity distribution did not show any significant pH dependence. Throughout the growth zone the deposition rates of the divalent cations, as calculated on the basis of the continuity equation, increased with lower pH. The data are consistent with the hypothesis that the effects of pH on the cation deposition rates are due to the increase in the divalent cation concentration of the soil solution at low pH and that the abundant uronic acid residues of the young walls of the meristem provide a reservoir of storage capacity for Ca and Mg under conditions of low nutrient availability.     

Simulation of Growth and Hydrotropism of Maize Roots

A three-dimensional model simulating the formation of root system architecture of maize was designed using object oriented programming (OOP) techniques. The model has been used to simulate the growth of roots in contrasting water profiles with or without gravitropism, and the mechanism of hydrotropism of root system and its relationship with gravitropism has been studied. In this model, the frontier of root system was treated as a population of root tips, each member of which responded individually to its local environment, and only a few of them could branch. The results of simulation showed that hydrotropism of maize roots could arise through the control of the elongation rate of single root by its local soil water potential. The difference in growth rate caused by the gradient of water potential along the soil profile alone could cause the root system as a whole to grow predominantly downwards, resulting in a shift of root distribution towards deeper layers. Gravitropism enhanced the downward predominance of the growth of root system, but the mechanism was different from that of hydrotropism.     

A Study of the Nitrogen and Water Factors in Competition with Agropyron repens (L.) Beauv.

From a consideration of the possible relationships between thelevel of a nutrient in the soil, the depletion due to a competitor,and the response of a test plant, it is argued that a validcriterion for the implication of a nutrient factor in root competitionis that some growth response of the test plant to competitionshould be less at higher levels of supply of that nutrient thanat lower levels. Young test plants of Impatiens parviflora were transplantedinto pans of soil in which stands of Agropyron repens were growing.Two levels of nitgrogen and gtwo rates of watering were appliedin a multifactorial design, and from half of the test plantsroot competition was excluded, so that their responses to rootcompetition could be measured under the various nitrogen andwatering treatments. A preliminary experiment hadf shown, and the competition experimentconfirmed, that the optimum nitrogen level for root growth andNet Assimilation Rate was less than the higher level employed,whereas this level was not sufficient for maximum leaf-growth.In agreement with this, the depressions of relative root growth-rateand Net Assimilation Rate caused by competition at the lowernitrogen level were significantly reduced at the higher level,but for relative leaf-growth this reduction did not reach significance.The effects of competition were reduced by the higher rate ofwstering, and for relative leaf growth-rate this interactionwas highly significant. It is deduced that competition from A. repensfinvolves bothnitrogen and water, and that water is probably the more importantfactor under normal conditions. There is no need to postulkateany toxic root product to account for the experimentalo results.Compared with competition above ground, it appeared that rootcompetition could have an effect on the relative growth-rateof I. parviflora about as great as a reduction of light intensityfrom full to 5–10 per cent. of summer daylight.     

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.     

Dryland Soil Hydrological Processes and Their Impacts on the Nitrogen Balance in a Soil-Maize System of a Freeze-Thawing Agricultural Area

Understanding the fates of soil hydrological processes and nitrogen (N) is essential for optimizing the water and N in a dryland crop system with the goal of obtaining a maximum yield. Few investigations have addressed the dynamics of dryland N and its association with the soil hydrological process in a freeze-thawing agricultural area. With the daily monitoring of soil water content and acquisition rates at 15, 30, 60 and 90 cm depths, the soil hydrological process with the influence of rainfall was identified. The temporal-vertical soil water storage analysis indicated the local albic soil texture provided a stable soil water condition for maize growth with the rainfall as the only water source. Soil storage water averages at 0–20, 20–40 and 40–60 cm were observed to be 490.2, 593.8, and 358 m³ ha⁻¹, respectively, during the growing season. The evapo-transpiration (ET), rainfall, and water loss analysis demonstrated that these factors increased in same temporal pattern and provided necessary water conditions for maize growth in a short period. The dry weight and N concentration of maize organs (root, leaf, stem, tassel, and grain) demonstrated the N accumulation increased to a peak in the maturity period and that grain had the most N. The maximum N accumulative rate reached about 500 mg m⁻²d⁻¹ in leaves and grain. Over the entire growing season, the soil nitrate N decreased by amounts ranging from 48.9 kg N ha⁻¹ to 65.3 kg N ha⁻¹ over the 90 cm profile and the loss of ammonia-N ranged from 9.79 to 12.69 kg N ha⁻¹. With soil water loss and N balance calculation, the N usage efficiency (NUE) over the 0–90 cm soil profile was 43%. The soil hydrological process due to special soil texture and the temporal features of rainfall determined the maize growth in the freeze-thawing agricultural area.     

Water Retention Capacity in Root Segments Differing in the Degree of Exodermis Development

Water loss from roots back into drying soil is a problem ofpractical importance in plants growing under conditions of verylow substrate water potential, such as dry or saline areas.Root exodermis is relatively impermeable and has been suggestedto play a protective role against water loss. The relative waterretention ability was compared in root segments from exodermal(maize, onion, sunflower, Rhodes grass and sorghum) and non-exodermalspecies (Pisum sativum,Vicia fabaand wheat). Apical and basalsegments from exodermal roots, with different degrees of exodermisdevelopment, were also compared, as were segments from sorghumroots in which the exodermis thickness had been modified bysubjecting the plants to a 30 d water stress treatment. Waterretention was significantly higher in segments from exodermalroots. In each root, water loss was higher in apical than inbasal segments, regardless of the presence of exodermis. Insorghum, prolonged drought treatment increased exodermis thickeningin nodal roots, however, no differences in rates of water losswere observed in segments obtained from control and droughtedplants. Soil sheaths formed around roots of Rhodes grass growingin very dry soil with the epidermis adhering tightly to thesheath. In plants growing in the field, soil sheaths may bemore effective than the exodermis in preventing root water loss.Copyright1999 Annals of Botany Company. Root, exodermis, rhizosheaths, water loss.     

玉米根系生长及向水性的模拟

用面向对象的程序设计（ＯＯＰ）技术组建了玉米（Ｚｅａ ｍ ａｙｓ）根系生长的三维模型,并用该模型模拟了根系在不同土壤水分剖面以及有、无向地性响应时的生长过程,探讨了根系向水性产生的机制以及向水性与向地性之间的关系．模型中将根系前沿看作由根尖构成的群体,每个根尖都对其周围环境独自作出响应,其中只有少数能继续分支． 模拟结果显示,玉米根系各单根受局部水势的影响而以不同速率伸长,即可造成总体的向水性． 如果土壤剖面上的水势由下到上递减,由此引起的单根伸长速率的不均匀分布将使整个根系在总体上表现出收拢和一致向下的生长趋势,并导致下层的根量相对增加．根系的向地性虽能使上述趋势增强,但它与向水性的机制是完全不同的     

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     

Uptake and Partitioning of Nitrogen by Maize Infected withStriga hermonthica

The uptake and partitioning of nitrogen (N) by maize infectedwith the parasitic angiosperm,Striga hermonthicawas investigatedin sand culture in a glasshouse. The purpose was to determinethe effect ofStrigaon N uptake and partitioning in maize. Maizewas grown at 22, 66 and 133 mg N per plant and sampled fivetimes. There was no significantStrigaxN interaction in any measuredresponse. Leaf dry matter ofStriga-infected maize, averagedover all N treatments, was 92% that of uninfected maize at thefour-leaf stage but by the 18-leaf stage it had decreased to58%. Similarly, stem dry matter of infected maize which was91% that of uninfected maize at the four-leaf stage was only42% at the 18-leaf stage. Root dry matter was similar for infectedand uninfected maize. N concentration in the leaf, stem androot declined asymptotically from the first to the last samplingdate for both infected and uninfected maize. The asymptoticvalue of N concentration inStriga-infected maize was 16% greaterin the leaf, 55% in the stem, and 21% in the root than in uninfectedmaize. The concentration of N inStrigawas higher than in maizeat the 16- and 18-leaf stages. Uptake of N was similar for infectedand uninfected plants at the four–eight leaf stage butat the eight–12 leaf stage, N uptake by infected maizewas 52% that of uninfected maize. However, the proportion oftotal plant nitrogen partitioned to the root was greater (P<0.001)forStriga-infected maize. These results showed that the extentto whichS. hermonthicareduced maize growth and N uptake, butincreased the proportion of N partitioned to the roots, didnot depend on the rate of N fertilizer applied.Copyright 1998Annals of Botany Company Maize; nitrogen; partitioning;Striga hermonthica; uptake.     

Evaluation in field conditions of a three-dimensional architectural model of the maize root system: Comparison of simulated and observed horizontal root maps

Most existing water and nutrient uptake models are based on the assumption that roots are evenly distributed in the soil volume. This assumption is not realistic for field conditions, and significantly alters water or nutrient uptake calculations. Therefore, development of models of root system growth that account for the spatial distribution of roots is necessary.The objective of this work was to test a three dimensional architectural model of the maize root system by comparing simulated horizontal root maps with observed root maps obtained from the field. The model was built using the current knowledge on maize root system morphogenesis and parameters obtained under field conditions. Simulated root maps (0.45 × 0.75 m) of horizontal cross sections at 3 depths and 3 dates were obtained by using the model for a plant population. Actual root maps were obtained in a deep, barrier-free clay-loamy soil by digging pits, preparing selected horizontal planes and recording root contacts on plastic sheets.Results showed that both the number of cross-sections of axile roots, and their spatial distribution characterized with the R-index value of Clark and Evans (1954), were correctly accounted for by the model at all dates and depths. The number of cross-sections of laterals was also correctly predicted. However, laterals were more clustered around axile roots on simulated root maps than on observed root maps. Although slight discrepancies appeared between simulated and observed root maps in this respect, it was concluded that the model correctly accounted for the general colonization pattern of the soil volume by roots under a maize crop.