Effects of Drought on Partitioning of Nitrogen in Two Wheat Varieties Differing in Drought-tolerance

A model was constructed to describe the translocation and partitioningof nitrogen on the seventh day after anthesis for well-wateredand droughted plants of two wheat varieties (Triticum aestivumL. cv. Warigal and Condor). The glasshouse-grown plants weredetillered so that a simplified model could be derived for themain stem. A 9-d drought treatment was imposed just after anthesisand this coincided with the period of endosperm cell divisionin the grains. Warigal, which had a higher grain yield thanCondor under drought, absorbed up to 15-times more nitrogenand translocated 1.5-fold more nitrogen to the shoot via thexylem. In both varieties, nitrogen redistributed from vegetativeorgans accounted for more than 60 per cent in control and 70per cent in droughted plants of the nitrogen needed for eargrowth. The net loss of nitrogen increased by 4-3 per cent inthe leaves, but decreased by 60 per cent in the stem under drought.Stem and roots appeared to play an important role in the nitrogeneconomy of droughted plants: less nitrogen was translocateddirectly to the grains from the senescing leaves and 40–60per cent more nitrogen was translocated to the roots. Nearlyall the nitrogen reaching the roots in the phloem was reloadedinto the xylem stream and translocated back to the shoot. Thetransfer of nitrogen through the stem was reduced under droughtand this resulted in a constant C:N ratio of the grains whichmay be important in the regulation of endosperm cell division. Triticum aestivum L., wheat, drought, nitrogen, senescence, translocation     

Effect of Post-anthesis Drought on Cell Division and Starch Accumulation in Developing Wheat Grains

Wheat plants (Triticum aestivum L., cv. Warigal) were subjectedto 20 d of water deficit during the period of endosperm celldivision. Drought accentuated the differences in final grainweight between spikelets and between grains within spikelets.The distal grains of top spikelets were most affected by drought.The maximum number of endosperm cells was, respectively, 30and 40 per cent lower in basal grains and distal grains of draughtedplants. In basal grains of middle spikelets, the number of largestarch granules per cell was unaffected but the number of smallstarch granules per cell was 45 per cent lower in grains ofdraughted plants. The initiation of small starch granules wasmore affected than cell division because severe water deficitoccurred earlier during the former process than the latter.Final dry weight appeared to correlate well with the maximumnumber of endosperm cells, but depended also on the number ofstarch granules per cell. Consequently, the amount of dry matterper cell was not constant in both treatments. The concentration of sucrose per endosperm cell was lower onlyin the droughted distal grains of top spikelets. The supplyof sucrose to endosperm cells did not regulate the initiationof small starch granules. Triticum aestivum L., wheat, drought, grain growth, cell division, starch     

Genotypic Differences in the Production and Partitioning of Carbohydrates Between Roots and Shoots of Wheat Grown under Zinc or Manganese Deficiency

The partitioning of soluble carbohydrates and starch betweenroots and shoots was investigated in wheat genotypes differingin Zn or Mn efficiency. The plants were grown for 11 d in achelate-buffered nutrient solution with sufficient or deficientZn and Mn supply. The Zn-efficient cultivar Warigal had a greatershoot fresh weight under sufficient Zn compared with the Zn-inefficientcultivar Durati. When supplied with sufficient Zn, Warigal hada greater concentration and content of soluble carbohydratesin roots and shoots in comparison with Durati. Under deficientZn supply, Durati had a greater concentration and content ofstarch in roots and shoots compared with Warigal. In an experimentwith varying supply of Mn, the Mn-efficient genotype C8MM hada greater shoot fresh weight than the Mn-inefficient cultivarBayonet under sufficient or deficient Mn supply. The concentrationof soluble carbohydrates in roots and shoots was decreased bydeficient Mn supply in C8MM but not in Bayonet. Starch accumulatedin the roots of Bayonet under deficient Mn supply. The resultssuggest that synthesis of carbohydrates is decreased under Zndeficiency, while they are preferentially partitioned to theroots to increase growth and thus the surface area availablefor Zn uptake. In the case of Mn deficiency, carbohydrate productionwas limited, but partitioning between roots and shoots was notaltered.Copyright 1997 Annals of Botany Company Carbohydrate; deficiency; manganese; assimilate partitioning; starch; Triticum aestivum; zinc     

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     

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     

Uptake of zinc from chelate-buffered nutrient solutions by wheat genotypes differing in zinc efficiency

Zinc-efficient Triticum aestivum (cv. Warigal) and Zn-inefficientTriticum turgidum conv. durum (cv. Durati) were grown in chelate-buffered,complete nutrient solutions providing either deficient or sufficientZn supply. When transferred to fresh chelatebuffered nutrientsolutions containing a wide range of Zn supplies (0–1.28µmol m^–3 Zn²⁺ activity) for 24–48 h, bothgenotypes increased net Zn uptake linearly with an increasein solution Zn²⁺ activities. Zincefficient Warigal accumulatedZn at a greater rate than Zn-inefficient Durati. The greaterrate of net Zn uptake was observed by plants of both genotypeswhen pretreated at deficient Zn supply. Net loss of Zn to thesolution was higher in plants pretreated with sufficient Znand was inversely related to Zn²⁺ activity in the external solution.When continuously supplied with 40 nmol m^–3 Zn²⁺, netZn uptake by Zn-efficient Warigal was significantly greaterthan that of Zn-inefficient Durati, but the difference diminishedwith plant age. Shoot concentrations of Fe, Mn and Cu were higherwhen plants were grown at deficient than at sufficient Zn supply.The Zn-efficient genotype transported less Zn and Fe to shootsand had higher Fe concentrations in roots than the Zn-inefficientgenotype, supporting the hypothesis that Zn efficiency may beconnected with inefficient transport of Fe from roots to shootsand thus initiation of the Fe-deficiency response resultingin increased release of Zn- and Fe-binding phytosiderophores.It is concluded that differential Zn efficiency of wheat genotypesis at least partly due to a greater ability of efficient genotypesto accumulate Zn. Key words: Chelate-buffering, genotypes, micronutrients, Triticum spp., uptake, zinc efficiency     

Thermal and Water Relations of Roots of Desert Succulents

Two succulent perennials from the Sonoran Desert, Agave desertiEngelm. and Ferocactus acanthodes (Lem.) Britton and Rose, loselittle water through their roots during drought, yet respondrapidly to light rainfall. Their roots tend to be shallow, althoughabsent from the upper 20 mm or so of the soil. During 12–15d after a rainfall, new root production increased total rootlength by 47 per cent to 740 m for A. deserti and by 27 percent to 230 m for F. acanthodes; root dry weight then averagedonly 15 per cent of shoot dry weight. The annual carbon allocatedto dry weight of new roots required 11 per cent of shoot carbondioxide uptake for A. deserti and 19 per cent for F. acanthodes.Elongation of new roots was greatest near a soil temperatureof 30°C, and lethal temperature extremes (causing a 50 percent decrease in root parenchyma cells taking up stain) were56°C and -7°C. Soil temperatures annually exceeded themeasured tolerance to high temperature at depths less than 20mm, probably explaining the lack of roots in this zone. Attached roots immersed in solutions with osmotic potentialsabove -2·6 MPa could produce new lateral roots, with50 per cent of maximum elongation occurring near -1·4MPa for both species. Non-droughted roots lost water when immersedin solutions with osmotic potentials below -0·8 MPa,and root hydraulic conductance decreased markedly below about-1·2 MPa. Pressure-volume curves indicated that, fora given change in water potential, non-droughted roots lostthree to five times more water than droughted roots, non-droughtedleaves, or non-droughted stems. Hence, such roots, which couldbe produced in response to a rainfall, will lose the most tissuewater with the onset of drought, the resulting shrinkage beingaccompanied by reduced root hydraulic conductance, less contactwith drying soil, and less water loss from the plant to thesoil. Agave deserti, Ferocactus acanthodes, roots, soil, temperature, water stress, drought, Crassulacean acid metabolism, succulents     

The Effects of Copper Supply and Shading on Retranslocation of Copper from Mature Wheat Leaves

Plants of Gamenya wheat (Triticum aestivum L.) were grown inpots of a Cu-deficient sand at two levels of Cu (deficient andsufficient), and harvested on days 13, 22, 28 and 38. In 50per cent of the pots in each Cu treatment, the oldest leaf andleaf 2 of the main stem were shaded when they reached full expansion. The Cu content of the oldest leaf of Cu-sufficient, unshadedplants was high at day 13 and declined rapidly to day 38. Thatof Cu-deficient, unshaded plants was initially relatively lowand declined much more slowly, so that at day 38 it resembledthat of Cu-sufficient plants. Shading the oldest leaf acceleratedthe loss of its Cu in both Cu-deficient and Cu-sufficient plants.The effects of shading and of Cu supply on the loss of Cu fromthe oldest leaf paralleled their effects on the loss of N andchlorophyll. The results suggest that most of the Cu in theoldest leaf does not move out until the leaf senesces. In Cu-deficient plants retention of Cu by old green leaves accentuatedCu deficiency. The release of Cu, resulting from shading theold leaves of Cu-deficient plants, stimulated the growth ofnew leaves. In Cu-sufficient plants, shading depressed growth. copper, shading, retranslocation, wheat, Triticum aestivum L.     

Depletion of Oxygen in Roots of Wheat and Maize by Respiration Following Encasement in Olive Oil

The extent to which isolated root systems of 14-d-old seedlingsof wheat (Triticum aestivum) and maize (Zea mays) were depletedof oxygen by respiration was measured after immersing them inoutgassed olive oil to exclude oxygen entry from the air. Atintervals over 45 min, gas from the roots was removed underpartial vacuum and oxygen partial pressures measured by gaschromatography. Contrary to earlier findings (Erdmann and Wiedenroth,1988), roots were able to utilize almost all of their oxygenwithin 20 min at 25°C, including that dissolved in the thinwater covering interposed between roots and oil. The rate ofaerobic respiration could be estimated readily from the timecourse of oxygen depletion.Copyright 1993, 1999 Academic Press Triticum aestivum, Zea mays, roots, anaerobiosis, respiration, oxygen, methods     

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     

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     

Changes in the Root System of Wheat Seedlings Following Root Anaerobiosis: III. Oxygen Concentration in the Roots

The oxygen status in roots of wheat seedlings (Triticum aestivum)was determined by a volumetric micro-absorption method. Plantsgrew in nutrient solution (aerated or nitrogen-flushed) or onflooded sand up to the 10th day. The roots were then exposedto aerated or hypoxic conditions for several hours before gaswas extracted by reducing the pressure within a concentratedsalt solution or by physical crushing. The oxygen content ofthe extracted gas bubbles was measured with pyrogallol. Comparativeexperiments with the helophytes Phalaris arundinacea and Carexacutiformis yielded similar oxygen concentrations to those alreadydescribed in literature. The concentrations of oxygen (13–16%)in young wheat roots were surprisingly high when exposed tonutrient solution flushed with nitrogen gas. Removal of the shoots decreased the oxygen concentration inthe roots, indicating some internal oxygen transport from shootsto roots. Detached, submerged roots of wheat still contained6% oxygen following 20 h of submergence in nitrogen-flushedsolution. A linear relationship was found between the oxygenconcentration in roots of Triticum aestivum, Zea mays and thetwo helophytes and the volume of extractable gas per volumeof root. This ratio corresponded to the extent of aerenchymaformation. Hence, a certain amount of oxygen may have been adsorbedonto the inner surfaces of the lacunae of the roots. However, the large amount of oxygen in the roots of intact wheatplants suggest that some parts of the root system are unlikelyto suffer from the oxygen shortage imposed by oxygen-deficientexternal conditions such as flooded soil. Triticum aestivum L. cv. Hatri, wheat, helophytes, roots, micro-absorption method, oxygen concentration, hypoxia, intercellular space     

Examination of a Model and Data Describing the Effect of Temperature on the Respiration Rate of Crop Plants

Some of the published evidence used in the synthesis and maintenancemodel of plant respiration is discussed in relation to the effectof temperature. Recalculations from the data of de Vries (1975b) give different results from those claimed by him. The modelis considered in terms of the use of substrate in the dark andits production in the light. It is suggested that starvationestimates of maintenance are not valid. The most reliable methodof observing synthesis respiration in whole plants appears tobe by following a discrete pool of substrate, as is possiblewith labelled carbon. Triticum aestivum L., Zea mays L., Helianthus annuus L., Vicia faba L., carbon dioxide, respiration, temperature, substrate content     

Growth of Near-isogenic Wheat Lines Differing in Development--Plants in a Simulated Canopy

An understanding of the principal factors regulating the growthof temperate cereals will identify opportunities to manipulatecrop growth. In an accompanying paper (Gomez-Macpherson, Richardsand Masle,Annals of Botany82: 315–322, 1998), growth aroundthe start of floral initiation was studied in isogenic wheat(Triticum aestivumL.) lines grown as spaced plants. In thispaper, two of the same near-isogenic wheat lines were grownas mini-canopies in a growth chamber. The objective was to determinewhether results obtained using spaced plants also apply to plantsgrown in a simulated canopy as a first step to emulate fieldconditions. Biomass of plant organs, leaf area and leaf andtiller appearance were determined from sowing to ear emergenceof the early developing line. Contrary to results obtained usingspaced plants, lines differed in their above-ground biomassaccumulation, although total plant biomass accumulation wassimilar. After the early line reached terminal spikelet stage(TS), biomass partitioning to the roots and leaves decreased,whereas partitioning to the stem and ear increased. This resultedin a lower root:shoot ratio in the early flowering line thanin the late line which remained vegetative. Tiller senescencealso began after TS in the early line whereas no tiller senescencewas observed in the late line during the experiment. Furthermore,after TS, net assimilation rate was greater and leaf area ratiowas lower in the early line. It is suggested that, after reachingTS, plants grown in a canopy become source limited comparedto widely spaced plants, or compared to plants that have notreached TS, and this results in less root growth.Copyright 1998Annals of Botany Company Development, growth, partitioning,Triticum aestivumL., wheat.     

Root Respiration and Carbohydrate Status of Two Wheat Genotypes in Response to Hypoxia

To investigate root respiration and carbohydrate status in relationto waterlogging or hypoxia tolerance, root respiration rateand concentrations of soluble sugars in leaves and roots weredetermined for two wheat (Triticum aestivum L.) genotypes differingin waterlogging-tolerance under hypoxia (5% O₂) and subsequentresumption of full aeration. Root and shoot growth were reducedby hypoxia to a larger extent for waterlogging-sensitive Coker9835. Root respiration or oxygen consumption rate declined withhypoxia, but recovered after 7 d of resumption of aeration.Respiration rate was greater for sensitive Coker 9835 than fortolerant Jackson within 8 d after hypoxia. The concentrationsof sucrose, glucose and fructose decreased in leaves for bothgenotypes under hypoxia. The concentration of these sugars inroots, however, increased under hypoxia, to a greater degreefor Jackson. An increase in the ratio of root sugar concentrationto shoot sugar concentration was found for Jackson under hypoxicconditions, suggesting that a large amount of carbohydrate waspartitioned to roots under hypoxia. The results indicated thatroot carbohydrate supply was not a limiting factor for rootgrowth and respiration under hypoxia. Plant tolerance to waterloggingof hypoxia appeared to be associated with low root respirationor oxygen consumption rate and high sugar accumulation underhypoxic conditions.Copyright 1995, 1999 Academic Press Oxygen consumption rate, sugar accumulation, Triticum aestivum L., waterlogging tolerance     

Changes in the acquisition and partitioning of carbon and nitrogen in the gibberellin-deficient mutants A70 and W335 of tomato (Solanum lycopersicum L.)

Even though the growth‐promoting effects of gibberellins (GAs) in plants are well established, little is known about GA action on carbon metabolism and the available reports seem contradictory. We studied the effects of GA deficiency in mutants of tomato (Solanum lycopersicum L.) on rates of carbon acquisition and the allocation of acquired carbon to growth and respiration of leaves, stems and roots. Carbon budgets were calculated from 24 h measurements of photosynthesis and respiration. The partitioning of nitrogen compounds to leaves, stems and roots, which strongly influences carbon budgets, was also studied. The GA‐deficient mutants acquired less carbon per unit plant mass per day than did the wild type and used a larger fraction of it for root growth and root respiration. To find out to what extent these changes were just consequences of restriction of growth, the experiment was repeated at a low exponential nitrate addition rate, which forced all genotypes to grow at the same rate. Under these conditions, the low‐GA mutants still photosynthesized and respired faster and partitioned more carbon to root growth than the wild type did. The reasons for the observed differences in carbon economies between the wild type and the low‐GA mutants are discussed.     

Growth Habits of Water-stressed Wheat Seedlings as Selection Criteria for Drought Resistance

A technique is described to induce slowly increasing water deficitsin wheat (Triticum aestivum L.) seedlings by rooting them indamp vermiculite and growing them in a humid atmosphere. Aftereight weeks, drought-resistant genotypes can be identified eitherby drought avoidance or by drought tolerance responses. Genotypeswith lesser drought resistance become dormant or produce sterilespikes. The test roughly predicts the relative yield performanceof cultivars in dry field conditions. Triticum aestivum, wheat, matric water deficits, drought resistance, yield selection     

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     

Elevated CO2 enhances plant growth in droughted N2-fixing alfalfa without improving water status

The long-term interaction between elevated CO₂ and soil water deficit was analysed in N₂-fixing alfalfa plants in order to assess the possible drought tolerance effect of CO₂. Elevated CO₂ could delay the onset of drought stress by decreasing transpiration rates, but this effect was avoided by subjecting plants to the same soil water content. Nodulated alfalfa plants subjected to ambient (400 μmol mol^?1) or elevated (700 μmol mol^?1) CO₂ were either well watered or partially watered by restricting water to obtain 30% of the water content at field capacity (ampproximately 0.55 g water cm^?3). The negative effects of soil water deficit on plant growth were counterbalanced by elevated CO₂. In droughted plants, elevated CO₂ stimulated carbon fixation and, as a result, biomass production was even greater than in well-watered plants grown in ambient CO₂. Below-ground production was preferentially stimulated by elevated CO₂ in droughted plants, increasing nodule biomass production and the availability of photosynthates to the nodules. As a result, total nitrogen content in droughted plants was higher than in well-watered plants grown in ambient CO₂. The beneficial effect of elevated CO₂ was not correlated with a better plant water status. It is concluded that elevated CO₂ enhances growth of droughted plants by stimulating carbon fixation, preferentially increasing the availability of photosynthates to below-ground production (roots and nodules) without improving water status. This means that elevated CO₂ enhances the ability to produce more biomass in N₂-fixing alfalfa under given soil water stress, improving drought tolerance.