Mechanisms of waterlogging tolerance in wheat – a review of root and shoot physiology

We review the detrimental effects of waterlogging on physiology, growth and yield of wheat. We highlight traits contributing to waterlogging tolerance and genetic diversity in wheat. Death of seminal roots and restriction of adventitious root length due to O₂ deficiency result in low root:shoot ratio. Genotypes differ in seminal root anoxia tolerance, but mechanisms remain to be established; ethanol production rates do not explain anoxia tolerance. Root tip survival is short‐term, and thereafter, seminal root re‐growth upon re‐aeration is limited. Genotypes differ in adventitious root numbers and in aerenchyma formation within these roots, resulting in varying waterlogging tolerances. Root extension is restricted by capacity for internal O₂ movement to the apex. Sub‐optimal O₂ restricts root N uptake and translocation to the shoots, with N deficiency causing reduced shoot growth and grain yield. Although photosynthesis declines, sugars typically accumulate in shoots of waterlogged plants. Mn or Fe toxicity might occur in shoots of wheat on strongly acidic soils, but probably not more widely. Future breeding for waterlogging tolerance should focus on root internal aeration and better N‐use efficiency; exploiting the genetic diversity in wheat for these and other traits should enable improvement of waterlogging tolerance.     

Morphological Changes in Wheat Seedlings (Triticum aestivum L.) following Root Anaerobiosis and Partial Pruning of the Root System

Morphological changes of roots and shoots following oxygen deficiencyin the root medium and after partial pruning of the root systemwere analyzed to obtain easily measurable parameters of theadaptive capacity of the root system against stress. Wheat seedlings(Triticum aestivum L. cv. Hatri) were cultivated on nutrientsolution which was either aerated or flushed with nitrogen,or were cultivated on flooded sand. On the third day after grainswelling in two pruning variants, roots 1–3 or 4–8were excised. Root anaerobiosis retarded longitudinal growth and biomass accumulationof the shoot and the seminal roots, and stimulated the developmentof adventitious roots. Partial removal caused a general compensativegrowth of the remaining roots under aerobic conditions. Root pruning plus anaerobiosis exceeded the compensatory capacityof the seedlings and thus caused a strong delay of elongationand biomass accumulation of both roots and shoots, includingdecrease of the root/shoot ratio. Roots became independent ofendosperm reserves on the seventh day under aerobic conditionsthough caryopses were not completely exhausted at this time.Additionally, oxygen deficiency delayed the reserve exhaustionprocess. Triticum aestivum L. cv Hatri, wheat, roots, growth analysis, morphology, anaerobiosis, strees, root pruning, compensatory capacity, caryopsis     

Differential Inhibition of Shootvs.Root Growth at Low Temperature and its Relationship with Carbohydrate Accumulation in Different Wheat Cultivars

Shoot and root growth rate, carbohydrate accumulation (includingfructan), reducing sugar content and dry matter percentage weremeasured in six wheat cultivars, ranging from winter to springtypes, grown at either 5 or 25 °C. At 5 °C (comparedwith 25 °C), the relative growth rate (RGR) of shoots wassimilarly reduced in all cultivars, but the RGR of shoots wasmore affected in winter wheats. This difference resulted insmaller root:shoot ratios than in spring wheats, which alsodeveloped more first-order lateral roots. A direct relationshipbetween carbohydrate accumulation at low temperatures and reductionin root growth was established. These results suggest that differentialshootvs.root growth inhibition at low temperature may play akey role in carbohydrate accumulation at chilling temperatures.This differential response might lead to improvements in survivalat temperatures below 0 °C, regrowth during spring, andwater and nutrient absorption at low temperatures.Copyright1997 Annals of Botany Company Wheat; Triticum aestivum; low temperatures; root growth; root: shoot ratio; sugar accumulation     

Physiology,gene expression,and metabolome of two wheat cultivars with contrasting submergence tolerance

Responses of wheat (Triticum aestivum) to complete submergence are not well understood as research has focused on waterlogging (soil flooding). The aim of this study was to characterize the responses of 2 wheat cultivars differing vastly in submergence tolerance to test if submergence tolerance was linked to shoot carbohydrate consumption as seen in rice. Eighteen‐day‐old wheat cultivars Frument (intolerant) and Jackson (tolerant) grown in soil were completely submerged for up to 19 days while assessing responses in physiology, gene expression, and shoot metabolome. Results revealed 50% mortality after 9.3 and 15.9 days of submergence in intolerant Frument and tolerant Jackson, respectively, and significantly higher growth in Jackson during recovery. Frument displayed faster leaf degradation as evident from leaf tissue porosity, chlorophyll_a, and metabolomic fingerprinting. Surprisingly, shoot soluble carbohydrates, starch, and individual sugars declined to similarly low levels in both cultivars by day 5, showing that cultivar Jackson tolerated longer periods of low shoot carbohydrate levels than Frument. Moreover, intolerant Frument showed higher levels of phytol and the lipid peroxidation marker malondialdehyde relative to tolerant Jackson. Consequently, we propose to further investigate the role of ethylene sensitivity and deprivation of reactive O₂ species in submerged wheat.     

Effects of NO2 and O3 Alone or in Combination on Kidney Bean Plants (Phaseolus vulgaris L.): Growth, Partitioning of Assimilates and Root Activities

Ten-day old kidney bean plants (Phaseolus vulgaris L. cv. Shin-edogawa)were exposed to 2.0 and 4–0 parts 10^–6 NO2, and0.1, 0.2, and 0.4 parts 10^–6 O3 alone or in combinationfor 2, 4, and 7 d. The effects of these air pollutants wereexamined with respect to the growth, partitioning of assimilates,nitrogen uptake, soluble sugar content, and root respiration. Decreased dry matter production was significant with all treatmentsexcept 2.0 parts 10^–6 NO₂ and 0.1 parts 10^–6 O₃.Exposure to mixtures of the gases produced more severe suppressionof growth than exposure to the single gases. Root/shoot ratiowas significantly lowered at 7 d by the gas treatments otherthan 2.0 parts 10^–6 NO₂ and 0.1 parts 10^–6 O₃. Thetotal nitrogen content of plants was increased by all treatments;the higher percent of nitrogen found with O₃ exposure will resultfrom the growth retardation which increases the concentrationof nitrogen in the plants because the absorption of nitrogenby roots was unaffected. The combination of O₃ with NO₂ significantlydecreased the assimilation of NO₂ by the plants. The concentration of soluble sugars in roots was decreased bythe gas treatments. There was a strong positive correlationbetween soluble sugar content and dry weight of the roots harvestedat 7 d. Root respiration was relatively unchanged until 5 dand then decreased significantly at 7 d by 2.0 parts 10^–6NO₂ and 0–2 parts 10^–6 O₃. Retarded growth of theroots and the decreased root respiration may be due to diminishedtranslocation of sugars from leaves to roots caused by exposureto air pollutants. The uptake of soil nitrogen was not closelyrelated with root respiration in the case of O₃ exposure. Key words: NO₂, O₃, Phaseolus vulgaris, Growth, Sugars, Root respiration     

Waterlogging Tolerance Among a Diverse Range of Trifolium Accessions is Related to Root Porosity, Lateral Root Formation and 'Aerotropic Rooting'

Waterlogging tolerance, root porosity and root anatomy wereevaluated for 20 Trifolium accessions (species and sub-species,all annuals) selected from the eight Sections of the genus.Nine accessions were sensitive [relative growth rate (RGR) reducedby up to 80%] to waterlogging, nine accessions were tolerant(RGR not reduced), and in two accessions RGR increased (up to1.9-fold), when compared to drained controls. Growth of themain (i.e. tap) root axis was severely reduced in all accessionswhen waterlogged. Lateral roots formed the bulk of the rootsystem of tolerant accessions when grown in waterlogged soil.Lengths of the longest lateral roots were up to three-timeslonger than the main root axis. Root porosity varied from 0.7–12%among accessions when grown in aerated solution and from 1.1–15.5%in plants grown in hypoxic (0.031–0.045 mol O₂m^-3) solution.In some accessions aerenchyma formed by cell lysigeny; in othersit formed by schizogenous cell separation, or a combinationof both processes. O₂consumption rates of expanded lateral roottissues varied by up to 1.7-fold (on a mass basis) among thesix accessions tested and was reduced by an average of 24% forroots of plants grown in hypoxic solution prior to measurements.Accessions with the highest root porosity tended to have longerroots when grown in waterlogged soil. Three accessions formed‘aerotropic roots’ and the lateral root lengthsof these plants exceeded those of all other accessions, suggestingenhanced O₂movement to the submerged lateral root axis via theaerotropic roots. Waterlogging-tolerant accessions were identifiedin seven of the eight Sections in Trifolium, and the tolerantaccessions tended to be those with extensive lateral root systemsof relatively high porosity. Copyright 2001 Annals of BotanyCompany Waterlogging, Trifolium, aerenchyma, hypoxia, flooding, root respiration, clover, root anatomy, root porosity, pasture, aerotropic roots     

Root Flooding of Muskmelon (Cucumis melo L.) Affects Fruit Sugar Concentration But Not Leaf Carbon Exchange Rate

Flooding the roots of greenhouse-grown muskmelon (Cucumis meloL. cv. Noy Yizreel) plants for 4 days reduced sucrose accumulation36% in the inner mesocarp and 88% in the outer mesocarp of developingfruit. Concentration of the translocated sugars raffinose andstachyose were also lower in fruit on flooded plants than inthose from nonflooded plants. In contrast, fruit hexose concentrationwas similar in both flooded and nonflooded plants. There wasno alteration in activities of enzymes associated with sucrosemetabolism in the fruit which could explain the decreased sucroseconcentration. Four days of root flooding caused no reductionin leaf carbon exchange rate or assimilate export rate, indicatingthat the reduction in fruit sucrose accumulation was not dueto source limitation. Root respiration, measured as CO₂ evolution,was approximately 30% lower in anaerobic roots than in aerobicroots. When viewed as carbohydrate consumed, a doubling of glycolyticactivity occurred in the anaerobic root mass. Increased demandfor carbohydrates by anaerobic roots may lead to a reductionin translocated carbohydrates available for sucrose biosynthesisin the developing fruit. (Received August 29, 1990; Accepted February 21, 1991)     

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

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

Influences of Water Status, Temperature, and Root Age on Daily Patterns of Root Respiration for Two Cactus Species

Daily patterns of root respiration measured as CO₂, efflux werestudied at various soil water potentials, temperatures, androot ages for individual, attached roots of the barrel cactusFerocactus acanthodes and the platyopuntia Opuntia ficus-indica.The daily patterns of root respiration for both establishedroots and rain roots followed the daily patterns of root temperature.Root respiration increased when root temperature was raisedfrom 5 °C to 50 °C for F. acanthodes and from 5 °Cto 55 °C for O. ficus-indica; at 60 °C root respirationdecreased 50° from the maximum for F. acanthodes and decreased25° for O. ficus-indica. Root respiration per unit d. wtdecreased with root age for both species, especially for rainroots. Root respiration rates for rain roots were reduced tozero at a soil water potential (     

Responses of shoot and root gas exchange, leaf blade expansion and biomass production to pulses of elevated C02 in hydroponic wheat

Short-term effects of elevated CO₂ during the early life phaseof plants may have long lasting consequences for growth andbiomass in later periods. We exposed hydroponically grown wheatseedlings to 5 d pulses of elevated CO₂ while leaf expansiongrowth as well as shoot and root gas exchange were measuredsimultaneously and continuously. Shoot photosynthesis, night-timeshoot respiration and below-ground respiration (largely by roots)roughly doubled when atmospheric CO₂ concentration was doubled.An interruption of CO₂ enrichment caused CO₂ assimilation andrespiration to return to control levels. However, while theresponse of photosynthesis was immediate, that of respirationshowed a hysteresis of about 3 d. Since shoot biomass increasedat elevated CO₂ (with no change in allocation pattern) equalfluxes per shoot or root system after a return to control CO₂concentrations indicate substantial downward adjustment of thecapacity for CO₂ fixation and release in high-CO₂ grown plants.Leaf expansion growth was completely unaffected by CO₂ enrichment,whereas tiller initiation was significantly increased (doubledin 18 d). We conclude that leaf growth in these wheat plantswas already carbon-saturated at ambient CO₂ concentration atoptimum mineral nutrient supply. The stimulation of growth ofwhole plants was exclusively due to enhanced tillering duringthis very early part of the life of these wheat plants. Key words: Allocation, atmospheric carbon dioxide enrichment, growth, photosynthesis, respiration, tillering, Triticum aestivum     

Relationship between mineral coating on roots and yield performance of wheat under waterlogging stress

Waterlogging stress limits the productivity of winter wheat(Triticum aestivum L. emend. Thell.) in many parts of the world.When wheat is grown under waterlogged conditions, a reddish-brownmineral coating can form on the epidermal surface of the roots.In wetland plants such as rice, the amount of mineral coatingformed on root surfaces is positively related to yield. Thisstudy was conducted to determine whether mineral coating onthe roots of wheat is related to yield potential under waterloggedconditions. Root mineral coatings formed under waterlogged conditionswere studied in 12 cultivars and two breeding lines over threeyears of greenhouse pot studies. Soil redox potential in thewaterlogged treatment ranged between - 46 and 171 mV, and grainyield was suppressed by 28–49% compared to well-drainedcontrols. Mineral coating formed on the roots from the waterloggedtreatment was determined to be composed primarily of iron, basedon ICP elemental analysis, iron-specific staining, and ion-mappingby scanning electron microscopy using an X-ray detector. Of11 elements quantified by ICP spectroscopy, six were significantlyaffected by waterlogging treatment, and three of these, Fe,Mn and P, were well-correlated negatively to yield. Aerenchymaformation in the heavily coated waterlogged roots appeared todisrupt the internal root structure, and exceeded 40% of cross-sectionalarea in one cultivar. Unlike rice, which shows a positive relationshipbetween oxygen release from roots, grain yield and mineral coating,in winter wheat, the amount of mineral coating is negativelyrelated to grain yield under waterlogged conditions. Key words: Wheat, mineral coating, waterlogging, aerenchyma, hypoxia     

Root Development and Aerenchyma Formation in Two Wheat Cultivars and One Triticale Cultivar Grown in Stagnant Agar and Aerated Nutrient Solution

Stagnant nutrient solution containing 0.1% agar and with anextremely low oxygen level (‘stagnant agar solution’)was used to simulate the gaseous composition and slow gas diffusionof waterlogged soils. Comparisons were made between the growthof two wheat cultivars(Triticum aestivum,cvs. Gamenya and Kite)and one triticale cultivar(Triticosecale,cv. Muir) grown instagnant relative to aerated solution. For all genotypes tested,immersion of roots in stagnant agar solution resulted in thedeath of the entire seminal root system and led to profuse branchingof the laterals of the nodal roots. In the stagnant agar solutionaerenchyma, as a percentage of the total cross sectional areaof nodal roots, was 18% for Muir, 14% for Kite and 12% for Gamenya;the roots of species with more aerenchyma also attained a longermaximum root length as predicted by the model of Armstrong (in:Woolhouse HW, ed.Advances in botanical research, vol. 7. London:Academic Press, 1979). Muir also had a nodal root/shoot freshweight ratio of 0.5 compared with 0.2–0.3 in Kite andGamenya. The greater number and length of nodal roots of Muirdid not lead to better shoot growth than in the other genotypes;one possible reason for this lack of improvement is a low efficiencyof aerenchymatous roots in wheat.Copyright 1998 Annals of BotanyCompany Root development; aerenchyma; stagnant agar;Triticum aestivumcv. Gamenya;Triticum aestivumcv. Kite;Triticosecalecv. Muir.     

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis I. Anatomy and Respiration in Triticum aestivum L.

Anatomical changes in roots of wheat seedlings (Triticum aestivumL. cv. Hatri) following oxygen deficiency in the rooting mediumwere investigated. The response of the plant to stress was testedat a very early developmental stage when the first adventitiousroots had just emerged. In order to analyze the adaptation ofdifferent roots, respiration rates of the roots 1–3 and4–n were compared with the respiration rates of the totalroot system. Oxygen deficiency was induced either by flushingnutrient solution with nitrogen or flooding of sand. In contrast to plants grown in well aerated media, both stressvariants led to a significant increase of the intercellularspace of the root cortex in seminal and first adventitious roots.Radial cell enlargement of cortical cells near the root tip,cell wall thickenings in flooded sand cultures and an increasein phloroglucinol-stainable substances were found to be furtherindicators of low oxygen supply. The roots 4–n which were promoted in growth under hypoxiashowed higher respiration rates; hence the total root respirationwas not restricted. Triticum aestivum L. cv. Hatri, wheat, roots, anatomy, anaerobiosis, stress, root respiration, intercellular space     

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     

Root Respiration for Agave deserti: Influence of Temperature, Water Status and Root Age on Daily Patterns

Root respiration, measured as CO₂ efflux, was studied for asucculent perennial from the Sonoran Desert, Agave deserti,with a new technique using individual, attached roots. The dailypatterns of root respiration closely followed the daily patternsof root temperature for both established roots and rain roots,with higher rates during the day when root temperature averaged27?C and lower rates at night when root temperature averaged17?C. When root temperature was raised from 5?C to 40?C, rootrespiration increased about 7-fold; from 45 ?C to 55 ?C, rootrespiration decreased about 2-fold, except for old establishedroots. Root respiration per unit dry weight for both root typesdecreased with age, the initial decrease being greater for rainroots than for established roots. Root respiration rates forrain roots were reduced to zero at a soil water potential (_soil)of –0.9 MPa and did not recover upon rewatering. Upondrying, root respiration rates for established roots were maintainedat about 12% of maximum, even when _soil fell to –1.6 MPa,and fully recovered 1.5 d after rewatering the soil. Such responsesof rain and established roots must be taken into account whenassessing the carbon costs for the root system. Key words: Agave deserti, CO₂ exchange, root respiration, temperature, soil water potential     

Site, Scale and Time-course for Adjustments in Lateral Root Initiation in Wheat Following Changes in C and N Supply

The pattern of lateral root initiation in seminal roots of wheat(Triticum aestivumL. cv. Alexandria) and the location, scaleand time-course for adjustments in initiation were studied afterchanges in C and N supply. Macroscopically visible primordiaappeared in a non-acropetal sequence with the frequency (numberper unit length) increasing with distance behind the main rootapex to a maximum at 40–50 mm behind the root tip. Pruningthe root system to a single seminal axis increased the primordiafrequency by 23% within 15 h. After longer periods, the effectof root-pruning was greater. The enhanced primordia frequencywas first observed in tissue located 0–10 mm behind theapex at the start of treatment. Feeding glucose (50 mM) alsoincreased primordia frequency within 15 h, but to a greaterextent, and here additional primordia were initiated in tissuelocated 0–10and10–20 mm behind the apex at the startof treatment. Withdrawing NO₃^-from one part of a split-rootsystem, whilst maintaining the supply to the other, reducedprimordia frequency in the non-fed roots and, in some cases,a compensatory increase in the NO₃^--fed roots was observed.The location and scale of the adjustments were similar to thosefound with root-pruning and glucose-feeding, but were slightlyslower to appear. In spite of some differences in detail, therewas a broad similarity in site, scale and time-course for adjustmentsin lateral root initiation with these treatments, which is consistentwith the operation of a common mechanism. Whenever an increasein primordia frequency was observed, it was associated withan increase in the ethanol-soluble sugar content of the tissue.However, the reduction in frequency in NO₃^--deprived roots wasalso accompanied by an increase in sugar content. There wasno consistent relationship between total N content of the tissueand primordia frequency, but there was between primordia frequencyand the rate of net NO₃^-uptake. The possible mechanisms controllinglateral root initiation are discussed. Compensatory growth; correlative growth; glucose; initiation; lateral root; nitrate; primordium; split-root; Triticum aestivum; wheat     

The influence of soil drought and partial waterlogging on water relations of Gmelina arborea seedlings

Summary Stomatal conductance of unstrossed, soil drought, and previously drought (predrought) Gmelina arborea seedlings increased in the morning and decreased before or immediately after midday. In the unstressed and predrought seedlings, leaf water potential decreased with increases in transpiration. In soil drought seedlings, there was some evidence of decreased hydraulic conductivity from soil to the plant, as indicated by the shape in the slope of the water potential/transpiration relationship. Root growth of drought plants was greater than in their unstressed counterparts at the lowest soil segment of a pot. The partial recovery of predrought seedlings was attributed to this subtantial root growth in the lowest soil segment.In the second experiment, Gmelina arborea seedlings were partially waterlogged, by flooding the polyethylene bag to half its length, for a period of 23 days. Waterlogging induced stomatal closure and reduction in leaf water potential but there was some evidence of tolerance to waterlogging towards the end of treatment. Root growth, shoot and root dry weights were slightly reduced below those of controls. After 9 days of waterlogging, adventitious roots began to form which correlated with depletion of soluble sugars in the shoot but with an increase in the roots.It is suggested that the tolerance of Gmelina plants to either soil drought or waterlogging may partly be due to partitioning of the soluble sugars from shoot to roots for production of roots and formation of adventitious roots respectively which are likely to enhance the flow of water from the soils to the plant. Therefore the plant response is very similar under conditions of increased deficits and surplus of soil water.     

Effects of shoot excision on in situ soil and root respiration of wheat and soybean under drought stress

Little information is available about the variability of root-derived respiration rate in relation to biotic factors such as photosynthesis and substrate availability in roots. Here we examine the role of decreased carbohydrates availability on root-derived respiration through removal of above ground biomass. Spring wheat (Triticum aestivum L. cv. Longchun 8139) and soybean (Glycina max L. cv. Tianchan 2) were grown in the field under a moveable rain shelter, and subjected to three different water regimes: (1) well-watered control; (2) moderate drought stress, and (3) severe drought stress. Root-derived respiration before and after shoot clipping, and the concentration of total nonstructural carbohydrate, malic and citric acid were measured for spring wheat and soybean. Root-derived CO₂ flux and total nonstructural carbohydrate concentration of clipped wheat decreased by 38% and 31%, respectively. However, for soybean the root- derived CO₂ flux and total nonstructural carbohydrate concentrations were only 58% and 62% of control, respectively, indicating the root respiration rate was controlled by the availability of carbon in the root. A significant positive correlation between total nonstructural carbohydrate concentration of the root and soil water content was observed in unclipped plants. Total nonstructural carbohydrate contributed 93% of the variance in root-derived respiration. Our results clearly show, that in the field, the availability of carbon substrate in roots determines root-derived respiration and plays a key link between soil moisture and root-derived respiration. A period of time is needed for root respiration to return to “steady-state” after shoot removal and this period needed is strongly dependent on species and soil water content.     

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis II. Morphology and Anatomy of Evolution Forms

Morphological and anatomical parameters which are variable underroot anaerobiosis in Triticum aestivum were checked on fivetaxa of primitive and modern wheats (and the related genus Aegilops).The plants were grown in nutrient solution which was eitheraerated or flushed with nitrogen. When the plants were flushedwith nitrogen a general retardation in longitudinal root growthoccurred in all of them, but only Triticum aestivum showed aclear promotion of growth of later appearing roots enablingit to maintain the same root/shoot ratio even under stress conditions.There was an increase in the volume of intercellular space inthe root cortex of nearly all the plants investigated. The diameterand the lignin content of the roots and the form of their corticalcells also varied. All these changes were expressed in the primitivewheats to a lesser extent than in the advanced Triticum aestivumindicating that there is a clear increase in the adaptive responsein the latter. Triticum species, Aegilops species, wheat, roots, anatomy, anaerobiosis, stress, intercellular space, selection     

The Influence of NO-3 and NH+4 Nutrition on the Gas Exchange Characteristics of the Roots of Wheat (Triticum aestivum) and Maize (Zea mays) Plants

Respiratory oxygen consumption by roots was 1·4- and1·6-fold larger in NH⁺₄-fed than in NO^-₃-fed wheat (Triticumaestivum L.) and maize (Zea mays L.) plants respectively. Higherroot oxygen consumption in NH⁺₄-fed plants than in NO^-₃-fedplants was associated with higher total nitrogen contents inNH⁺4-fed plants. Root oxygen consumption was, however, not correlatedwith growth rates or shoot:root ratios. Carbon dioxide releasewas 1·4- and 1·2-fold larger in NO⁺3-fed thanin NH⁺₄-fed wheat and maize plants respectively. Differencesin oxygen and carbon dioxide gas exchange rates resulted inthe gas exchange quotients of NH^-₄-fed plants (wheat, 0·5;maize, 0·6) being greatly reduced compared with thoseof NO^-₃-fed plants (wheat, 1·0; maize, 1·1). Measuredrates of HCO^-₃ assimilation by PEPc in roots were considerablylarger in 4 mM NH⁺₄-fed than in 4 NO^-₃ plants (wheat, 2·6-fold;maize, 8·3-fold). These differences were, however, insufficientto account for the observed differences in root carbon dioxideflux and it is probable that HCO^-₃ uptake is also importantin determining carbon dioxide fluxes. Thus reduced root extension in NH⁺₄-fed compared with NO^-₃-fedwheat plants could not be ascribed to differences in carbondioxide losses from roots.Copyright 1993, 1999 Academic Press Triticum aestivum, wheat, Zea mays, maize assimilation, ammonium assimilation, root respiration