An Isotopic Study of the Fixation of Nitrogen associated with Nodulated Plants of Alnus, Myrica, and Hippophae

Nodulated plants of Alnus glutinosa, Myrica gale, and Hippophaërhamnoides, the root systems of which had been exposed to excessfree ¹⁵N, showed substantial enrichment in fixed ¹⁵N contentin all parts of the plant, but particularly in the root nodules.The data resemble closely those obtained by the present andprevious authors with legumes under comparable conditions, andsupport the conclusion already drawn from experiments of traditionaltype that the nodules of these non-legumes are similar to thoseof legumes in their function and relation to the rest of theplant. Nodulated plants of Alnus and Myrica continued to fixfree nitrogen concurrently with the uptake of combined nitrogenwhen the latter was supplied in the rooting medium in amountsunlikely to be exceeded in the field. Isotopic tests on detachednodules of one of the species (Alnus glutinosa) showed thatfixation continued, and though much reduced as compared withthat shown by attached nodules it considerably exceeded thatexperienced by the present and previous authors with detachedlegume nodules, on the basis of fixation per unit of total nodulenitrogen. It is probable that detached Alnus nodules presentconvenient material for the further study of various aspectsof the fixation process.     

Isotopic Studies of Nitrogen Fixation in Non-Legume Root Nodules

Isotopic studies are presented on six of the eight recognizednon-legume nodule-forming genera of Angiosperms. Evidence hasbeen obtained of the occurrence of fixation of atmospheric nitrogenin the root nodules of Casuarina, Ceanothus, and Shepherdia,now examined isotopically for the first time. In further studiesof the nitrogen-fixing nodules of Hippopha and Alnus it is shownthat the fixation continues for a longer period after detachmentfrom the plant than is the case with legume nodules, and thatthe enrichment in ¹³N which they (and also detached Casuarinanodules) finally achieve after exposure to excess of the freeisotope considerably exceeds that shown by detached legume nodules.Fixation in detached Myrica nodules was not clearly affectedby reduction in oxygen supplied until the proportion of oxygenwas less than 5 per cent.     

The Cobalt Requirement of Non-legume Root Nodule Plants

As already shown for Alnus glutinosa, cobalt is found to beessential for the proper growth of nodulated plants of Casuarinacunninghamiana and Myrica gale in a nitrogen-free rooting medium.If cobalt is not supplied, the plants develop symptoms of nitrogendeficiency; under the conditions of the experiments such symptomsbecame pronounced during the second season of growth of theseperennial plants. No cobalt requirement could be detected innon-nodulated plants of Alnus and Myrica supplied with nitrateor ammonium-nitrogen, and this suggests that in nodulated plantsthe need for cobalt is confined to the nodules. Vitamin B₁₂analogues are shown to be present in the nodules in relativelylarge amounts when cobalt is supplied, their formation beingattributed to the endophytes, which may therefore require cobaltfor their growth. The great reduction in fixation of atmosphericnitrogen in cobalt-deficient nodules may be due to a retardedgrowth of the endophyte, though this is not the only possibility.The cobalt relation of these non-legumes appears to be basicallysimilar to that of legumes.     

A Quantitative Study of Fixation and Transfer of Nitrogen in Alnus

A quantitative study of the fixation of nitrogen and of itstransfer from the nodules to the remainder of the plant hasbeen made in Alnus plants during their first season of growth.Fixation per plant reached a maximum in late August but fellrapidly with the onset of autumn, while fixation per unit dryweight of nodule tissues was greatest in young nodules and wasof the same order as in nodulated legumes. Throughout the growthseason there was a steady transfer from the nodules of some90 per cent, of the nitrogen fixed. The rate of fixation relativeto the growth of the endophyte is much higher than in free-livingnitrogen-fixing organisms and is clearly governed by the nitrogenrequirements of the entire symbiotic system. These findingsare compatible with the view that the fixation process is extracellularto the endophyte.     

The Fixation of Nitrogen associated with the Root Nodules of Myrica gale L., with Special Reference to its pH Relation and Ecological Significance

The paper deals with the fixation of nitrogen by nodulated plantsof Myrica gale under experimental conditions, the fixation being,in the first year of development, of a magnitude comparableto that of legumes under similar conditions. The effect of acidityon the fixation has been investigated. The Myrica organism ismarkedly adapted to acid conditions. The observations are correlatedwith field data in Britain and help to explain the distributionand habits of the species.     

The Development and Significance of the Root Nodules of Casaurina

Young plants of Casuarina cunninghamiana and of C. equisetifoliagrowing in water culture developed macroscopic nodules in 26to 35 days from inoculation, the nodulation being most successfulat pH near to neutrality and falling off much more rapidly atlower pH than in other non-legume nodule-forming genera, confirmingthe distinctiveness of the Casuarina organism. The roots springingfrom the nodule lobes are shown to be characterized by upwardgrowth and in this to resemble remarkably closely the correspondingroots of Myrica. Nodulated plants of Casuarina are able to growvigorously in culture solution free of combined nitrogen, showingthat fixation of atmospheric nitrogen occurs, amounting in C.cunninghamiana to 50 mg. per plant during 6 months of activegrowth. The evidence indicates that the fixation occurs in thenodules, and that these have exactly the same functional significanceas those of legumes.     

Nitrogen Fixation in Wild Legumes

Isolations of the rhizobia from nodules of field plants of Ulexeuropaeus and Medicago lupulina have been examined for effectivenessin fixation of nitrogen by inoculating them into host plantsgrowing in a medium free of combined nitrogen. The results indicatethat vigorous fixation may be presumed to occur in associationwith field plants of the Ulex species, but that fixation inMedicago is probably decreased by the presence of ineffectiverhizobia in some nodules. The general importance of fixationin wild legumes is discussed, and it is pointed out that dueattention must be paid to non-legume genera with root nodules.These genera are locally abundant, and in former periods werestill more prominent in some regions of the world.     

Relationships between Acetylene Reduction Activity, Hydrogen Evolution and Nitrogen Fixation in Nodules of Acacia spp.: Experimental Background to Assaying Fixation by Acetylene Reduction under Field Conditions

Hansen, A. P., Pate, J. S. and Atkins, C. A. 1987. Relationshipsbetween acetylene reduction activity, hydrogen evolution andnitrogen fixation in nodules of Acacia spp.: Experimental backgroundto assaying fixation by acetylene reduction under field conditions.—J.exp. Bot. 38: 1–12 Glasshouse grown, symbiotically-dependent seedlings of Acaciaalata R.Br., .A. extensa Lindl., and A. pulchella R.Br. wereexamined for acetylene reduction in closed assay systems usingundisturbed potted plants, excavated whole plants, nodulatedroots or detached nodules. Nitrogenase activity declined sharplyover the first hour after exposure of detached nodules to acetylene(10% v/v in air), less steeply or not at all over a 3 h periodin assays involving attached nodules. Using detached nodules,rates of acetylene reduction, nitrogen (¹⁵N₂) fixation, andhydrogen evolution in air (¹⁵N₂) and acetylene-containing atmosphereswere measured in comparable 30 min assays. Total electron flowthrough nitrogenase in air was determined from rates of nitrogen(¹⁵N₂) fixation ( ? 3) plus hydrogen evolution, that in thepresence of acetylene from rates of acetylene reduction andhydrogen evolution in air: acetylene. Values for the ratio ofelectron flow in air: acetylene to that in air ranged from 0?43to 0?83 in A. pulcheila, from 0?44 to 0?66 in A. alala and from0?37 to 0?70 in A. extensa, indicating substantial inhibitionof electron flow through nitrogenase of detached nodules byacetylene. Relative efficiencies of nitrogenase functioningbased on hydrogen evolution and acetylene reduction were from0?15 to 0?79, those based on nitrogen (¹⁵N₂) fixation and hydrogenevolution from 0?53 to 0?87. Molar ratios of acetylene reducedto nitrogen (¹⁵N₂) fixed were 2?82 ? 0?24, 201 ? 0?15, and 1?91? 0?11 (?s.e.; n = 7) for A. pulcheila,A. extensa and A. alata respectively A standard 5–10 min acetylene reduction assay, conductedon freshly detached unwashed nodules in daytime (12.00–14.00h), was calibrated for field use by comparing total N accumulationof seedlings with estimated cumulative acetylene reduction overa 7-week period of glasshouse culture. Molar ratios for acetylenereduced: nitrogen fixed using this arbitrary method were 3?58for A. alata, 4?82 for A. extensa and 1?60 for A. pulchella.The significance of the data is discussed. Key words: Acacia spp, nitrogenase functioning     

Fine Structure of Peanut Root Nodules Induced by Nod+Fix- Strains of Bradyrhizobium with Special Reference to Lipid Bodies

Ineffective nodules of peanut induced by two nod⁺fix^–strains of Bradyrhizobium sp. were compared with the ones inducedby nod⁺fix⁺ strain NC92. One of the fix^– strains, 639is a transconjugate Tn5 mutant of NC92, while the other, 7091,is an isolate from ICRISAT soil. Both induce small nodules lackingleghemoglobin and nitrogen-fixing activity. Ultrastructuralobservations revealed that the nodules of 639, have enlargedperibacteroid space and lack persistence of nodule function.The 7091-induced nodules showed impediment in bacteroid releaseand differentiation. In both the ineffective nodules large amountsof lipid bodies were found to accumulate several times in excess,compared to the effective NC92 nodules. The large lipid accumulationin absence of nitrogen fixation supports the hypothesis thatin peanut nodules lipid bodies are utilized as a supplementarysource of carbon and energy for nitrogen fixation. Peanut, lipid bodies, nitrogen fixation, nod⁺fix^– Bradyrhizobium, ultrastructure     

Cyclic AMP in Rhizobia and Symbiotic Nodules

Cyclic adenosine 3',5'-monophosphate (cAMP) content of variouscultured rhizobia strains and tissues of legumes and non-leguminousplants was measured by enzyme immunoassays. Most rhizobia, culturedfor 44 to 165 h, contained cAMP ranging from 0.6 to 5 pmol mg^-1proteinexcept forAzorhizobium caulinodansORS571. The culture mediaalso contained varying amounts of cAMP depending on the strainof rhizobia.Azorhizobiumcells and their media contained no detectablecAMP. Nodules from most legumes and non-legumes had cAMP contentsranging from 2–70 pmol g^-1f.wt. However, nodules fromSesbaniarostrata,Crotalaria spectabilisandParasponia andersoniishowedundetectable cAMP levels, and those fromGlycine maxandVignaangularisoccasionally showed levels below the detection limit.The leaves of non-legumes mostly had cAMP levels below detectionlimit (approx. 1.0 pmol g^-1 f.wt), while the leaves ofa few legumes occasionally had detectable cAMP. The possiblerole of cAMP as a symbiotic signal is discussed. cAMP; legumes; modules; rhizobia; symbiosis     

Nodule Function in Symbiotic Bambara Groundnut (Vigna subterraneaL.) and Kersting's Bean (Macrotyloma geocarpumL.) is Tolerant of Nitrate in the Root Medium

Nitrogen-fixing activity in two nodulated African legumes, Bambaragroundnut (Vigna subterraneaL.) and Kersting's bean (MacrotylomageocarpumL.), was assessed in the presence of nitrate (NO₃^-)ions in the rooting medium. Nitrogenase activity was unimpairedby the supply of 5 mol m^-3NO₃to both species. Also, large concentrationsof ureides dominated the transpiration stream of NO₃-fed plants.Compared to other symbiotic legumes cultured with similar NO₃concentrations,nodule functioning in the tested landraces of Bambara groundnutand Kersting's bean is tolerant of NO₃ions in the rhizosphere.The potential benefits of such naturally occurring NO₃-tolerantsymbioses are substantial, as they would permit inorganic Nfertilizer application in intercropping systems without inhibitingN₂fixation in the associated legumes.Copyright 1998 Annals ofBotany Company NO₃tolerance, Bambara groundnut, Kersting's bean, nitrogenase activity, xylem ureides.     

Hydrogen Effect on Nitrogenase (C2H2 Reduction) Response to Oxygen in Bradyrhizobium japonicum-Phaseoleae Symbioses

The influence of hydrogenase in Bradyrizobum-Phaseoleae symbioseswas studied ex-planta and in-planra in soybean (Glycine max)and cowpea (Vigna unguiculata). The hydrogenase was activatedby the addition of hydrogen in the incubation gas phase whichmodified the response of nitrogenase activity of Hup⁺ (hydrogenuptake positive) symbiosis to the external oxygen partial pressure.For bacteroids the hydrogenase expression increased nitrogenaseactivity at supraoptimal pO₂, acting possibly as a respiratoryprotection of nitrogenase. However, at suboptimal pO₂, nitrogenaseactivity of Hup⁺ bacteroids decreased with hydrogen, a phenomenonattributed to the lower efficiency of ATP synthesis from hydrogenthan from carbon substrates oxidation. For undisturbed nodules,the hydrogenase expression in soybean increased the optimalpO₂ for ARA (COP), from 35.3 to 40.3 kPa O₂, and the ARA atsupraoptimal pO₂; at suboptimal PO₂ there was a negative effectof hydrogenase on ARA, although this inhibition was less thanon bacteroids and was not detected if plants were grown at 15°C rather than 20 °C root temperature. No H₂ effectwas detected on cowpea nodules. The results on soybean nodulesare consistent with the concept that symbiotic nitrogen fixationis oxygen-limited and that hydrogenase activity has no beneficialeffect on nitrogen fixation in O₂ limitation. Key words: Glycine max, hydrogenase, nitrogenase, nitrogen fixation, nodules, Vigna unguiculata     

Identification and Localization of Frankia Strains in Alnus Nodules by In Situ Hybridization of nif H mRNA with Strain-Specific Oligonucleotide Probes

In situ hybridization of Frankia mRNA with specific probes wasused to localize the strains Arl3 and AcoN24d in Alnus nodulesobtained after inoculation with one or both strains. The probesconsisted of 18-mer oligonucleotides, complementary to strain-specificsequences located within the nif H gene. Sections of nodulesinoculated with only one strain revealed a specific hybridizationbetween the probe and the corresponding Frankia strain mRNA.In sections of dually-inoculated nodules the presence of thestrain AcoN24d in the nodule was clearly shown whereas thoseof the strain Arl3 could not be detected. This suggests thatthe strain Arl3 is less infective than the strain AcoN24d andis not present within the nodule. Key words: Nitrogen fixation, actinorhizae, autoradiography, histochemistry     

Translocation of Nitrogenous Compounds in Symbiotic and Nitrate-Fed Amide-Exporting Legumes

Peoples, M. B., Sudin, M. N. and Herridge, D. F. 1987. Translocationof nitrogenous compounds insymbiotic and nitrate-fed amide-exportinglegumes.–J. exp. Bot. 38: 567–579. The transport of nitrogen from the roots and nodules of chickpea(Cicer anetinum L.), lentil (Lens culinaris Medic), faba bean(Vicia faba L.) and pea (Pisum sativum L.) was examined in glasshouse-grownplants supplied either with nitrate-free nutrients or with nutrientssupplemented with 1,2,4 or 8 mol m^-3153N-nitrate. A sixth treatmentcomprised uninoculated plants supplied with 8–0 mol m^-31513N-nitrate. For each species, more than 75% of the nitrogenwas exported from the nodules as the amides, asparagine andglutamine. In fully symbiotic plants, the amides also dominatednitrogen transport to the shoot When N2 fixation activity wasdecreased by the addition of nitrate to the rooting medium,the N-composition of xylem exudate and stem solutes changedconsiderably. The relative concentrations of asparagine tendedto increase in the xylem whilst those of glutamine were reduced;the levels of nitrate increased in both xylem exudate and thesoluble nitrogen pool of the stem with a rise in nitrate supply.The changes in relative nitrate contents reflected generallythe contributions of root and shoot to overall nitrate reductaseactivity at the different levels of nitrate used. The relationshipsbetween the relative contents of xylary or stem nitrate andamino nitrogen and the plants' reliance on N2 fixation (determinedby the ¹⁵N isotope dilution procedure) were examined. Data suggestthat compositional relationships based on nitrate may be reasonableindicators of symbiotic dependence for all species under studyexcept faba bean when greater than 25% of plant nitrogen wasderived from N2 fixation. Key words: Nitrogen, translocation, legumes     

Biological and Cultural Characteristics of the Effective Frankia Strain HFPCcl3 (Actinomycetales) from Casuarina cunninghamiana (Casuarinaceae)

From homogenates prepared from surface-sterilized nodules ofseedlings of Casuarina cunninghamiana grown aeroponically, astrain of Frankia designated HFPCc13 was isolated and has beengrown in pure filamentous culture in a defined synthetic nutrientmedium. Vesicle and sporangium formation can be induced by removalof combined nitrogen from the medium.Frankia strain HFPCc13nodulates young seedlings of C. cunninghamiana and C. equisetifoliawithin three weeks of inoculation with an optimum root mediumpH of 6–7 for nodulation and optimum temperature of 30–35^°C. The presence of combined nitrogen in the root mediuminhibits nodulation with NH₄⁺ more inhibitory than NO₃^–.Frankia HFPCc13 does not nodulate Allocasuarina species withinthe same family nor several other possible actinorhizal plantstested. Thus this strain is quite precise in its host specificity.The rate of acetylene reduction was greater in C. cunninghamianathan the closely related species C. equisetifolia. In neitherof these host species were vesicles observed to occur withinthe infected root nodules which had been demonstrated to beactively fixing dinitrogen. Root nodules were shown to be activein acetylene reduction over a range of O₂ concentration in thegaseous environment with an optimum at about 20 per cent O₂,the ambient P_O2 of the air. The mechanism(s) for oxygen protectionof nitrogenase within the filamentous form of Frankia withinthese nodules remains to be explained. Casuarina, Frankia, nodulation, nitrogen fixation     

Photosynthesis and Formation of Fats in a Diatom

A study has been made, using C¹⁴ as a tracer, of the variationswhich occur according to the physiological condition of thecells in the distribution of carbon fixed, both in the lightand in the dark, by the diatom Navicula pelliculosa (Bréb.)Hilse during periods of 30 to 300 seconds. Fixation into thefollowing cell fractions was determined: (A) material solublein 80 per cent. ethanol but insoluble in benzene, (B) materialsoluble both in 80 per cent. ethanol and in benzene, and (C)material insoluble in 80 per cent. ethanol. Carbon fixed inphotosynthesis was incorporated, rapidly and in amounts representingup to 70 per cent. of the total fixation, into fractions B andC, as well as into fraction A. Considerable variation was foundin the proportions of carbon entering the three fractions inthe light; in actively growing cells the proportion enteringfraction C preponderated over that in B, corresponding to thesynthesis of protein, whereas in nitrogen-deficient cells fixationin B was the greater, corresponding to the synthesis of fat.These patterns changed only slowly, over periods of days, followingthe transfer of cells to altered conditions of nitrate supply.However, when ammonium nitrogen was supplied to nitrogen-deficientcells a marked change in distribution of carbon fixed occurredwithin 5 minutes, fixation in fraction B falling to a low valueand that in A rising correspondingly. In cells subjected toprolonged nitrogen-deficiency, fixation in fraction B fell toa relatively low value but the proportion which this fractionformed of the total dry matter in the cells rose as a resultof an increased rate of loss from the cells of constituentsother than lipides. The distribution of carbon fixed was alsodependent on light intensity. Fixation in fractions B and Crose relatively to that in A as light intensity was increasedup to 100 foot-candles but fell again at the highest intensityused, 2,000 foot-candles. These results are discussed with particularreference to the relationship between fat accumulation and photosynthesisin algae.     

Does legume nitrogen fixation underpin host quality for the hemiparasitic plant Rhinanthus minor?

The high quality of leguminous hosts for the parasitic plantRhinanthus minor (in terms of growth and fecundity), comparedwith forbs (non-leguminous dicots) has long been assumed tobe a function of the legume's ability to fix atmospheric nitrogen(N) from the air and the potential for direct transfer of compatibleamino compounds to the parasite. Using associations betweenRhinanthus minor and Vicia faba (Fabaceae) that receive N eitherexclusively via symbiotic associations with rhizobia supplyingorganic N fixed from N₂ or exclusively through the supply ofinorganic nitrate to the substrate, the underlying reasons forthe quality of legumes as hosts for this parasite are unravelled.It is shown that sole dependence of the host, V. faba, on Nfixation results in lower growth of the attached parasite thanwhen the host is grown in a substrate supplied exclusively withinorganic N. In contrast, the host plants themselves achieveda similar biomass irrespective of their N source. The physiologicalbasis for this is investigated in terms of N and abscisic acid(ABA) partitioning, haustorial penetration, and xylem sap aminoacid profiles. It is concluded that legume N fixation does notunderpin the quality of legumes as hosts for Rhinanthus butrather the well-developed haustorium formed by the parasite,coupled with the lack of defensive response of the host tissuesto the invading haustorium and the presence of sufficient nitrogenouscompounds in the xylem sap accessible to the parasite haustoria,would appear to be the primary factors influencing host qualityof the legumes. Key words: ABA, haustorium, legume, nitrogen fixation, nodules, parasitic plant Received 14 November 2007; Revised 7 January 2008 Accepted 8 January 2008     

Young Reproductive Structures Promote Nitrogen Fixation in Soya Bean

In many legumes the transition from the vegetative to the reproductivephase of development is associated with a marked increase inthe rate of symbiotic nitrogen fixation. In soya bean [Glycinemax (L.) Merr.), the removal of reproductive parts at differentstages of their development showed that the increase in nitrogenfixation rate was primarily due to the presence of flower buds.The increase in the fixation rate of intact reproductive plantswas accompanied by a rapid increase in the weight of noduleson lateral roots and it is suggested that these nodules areresponsible for much of the nitrogen fixation which occurs duringreproductive growth. Maintaining plants in the vegetative stateprovided evidence which suggests that it is the flower budsand not the flowering stimulus which are responsible for theincrease in fixation rate. The marked effects on vegetativegrowth of removing reproductive parts suggests that the mechanisminvolved in the promotion of nitrogen fixation may be hormonal. Glycine max (L.) Merr., soya bean, nitrogen fixation     

Nitrogen Economy of Post-fire Stands of Shrub Legumes in Jarrah (Eucalyptus marginata Donn ex Sm.) Forest of S.W. Australia

Hansen, A. P., Pate, J. S., Hansen, A. and Bell, D. T. 1987.Nitrogen economy of post-fire stands of shrub legumes in jarrah(Eucalyptus marginata Donn ex Sm.) forest of S.W. Australia.—J.exp. Bot. 38: 26–41. Growth, demography and N economy of 1–6-year-old standsof Acacia pulchella, A. alata, A. extensa and Bossiaea aquifoliumwere examined using population sampling to assess annual incrementsof N as living biomass, and returns of N as litter, seed anddead plants. Dependence on nitrogen fixation was assessed fromseasonal profiles of acetylene reduction, employing data fromprevious calibrations to convert C₂H₂ reduced to N₂ fixed. After2 years of slow growth and minimal reproduction all speciesgrew rapidly to achieve maximum or near maximum size and seedproduction. Intense self-thinning of stands occurred in thesecond and third years, especially in the highly dense standsof the smallest species, A. alata. Annual turnover in standsranged from 0?3 to 127 kg N ha^–1 year^–1, dependingon species current age and density of a stand. Returns of Nas litter and shed seed comprised small proportions of the annualbudgets, but returns due to plant death equalled or exceededincrements of living biomass in years when stands were thinningrapidly. Proportional dependencies of the species on fixed N₂were relatively high (13–61%) in first year seedlings,and then declined markedly to 1?1–3?4percnt; in the second,0?3–1?6% in the third, and, with one exception, to wellbelow 1% in the fourth and sixth year stands. Mean annual ratesof N₂ fixation over the 6-year post-fire period were 1?6 kgN ha^–1 year^–1 for A. alata, 0–49 for A. pulchella,0?19 for B. aquifolium and 0-10 for A. extensa Key words: Shrub legumes, post-fire N economy     

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