Nodulation ofMedicago sativa in solution culture

Summary The time course of increase in nodule number was observed in solution cultures of controlled composition, over periods of 5 to 7 weeks following the inoculation of 8-day-old seedlings. The plants produced successive crops of nodules two to three weeks apart. Maintaining nitrate at concentrations of 0.2 or 0.5mM greatly reduced the number of nodules in the first crop but only slightly or not at all in the larger following crops, relative to the numbers in nil-nitrate control cultures. This was not due to any decrease in the sensitivity of nodulation to inhibition by nitrate. Nodulation in the nil-nitrate controlplants was restrained following their abundant first-crop nodulation. The experiments ceased to be simple tests of effects of nitrate on rate of nodule production as soon as the first crop of nodules appeared. Subsequent nodule production was then inhibited approximately equally, by prior nodulation in the abundantly nodulated control-plants, and by nitrate in the sparsely nodulated plants in the nitrate treatment.When the effect of nitrate was tested on plants comparable in size and in number of previously established nodules, it inhibited nodule production by 70–80%, whether the plants were seedlings or month-old plants, previously nodulated or not.     

Shoot/root assimilate allocation and nodulation ofVigna unguiculata seedlings as influenced by shoot light environment

Spectral balance of light received by southern pea [Vigna unguiculata (L.) Walp.] seedling shoots affected photoassimilate allocation among leaves, stems and roots. A higher ratio of far-red (FR) relative to red (R) light resulted in longer stems, higher shoot/root biomass ratio, less massive roots and fewer nodules. The same response pattern to FR/R ratio was obtained in a controlled environment with artificial light sources, or in sunlight where the FR/R ratio was modified by reflection from different colored soil surfaces or by FR reflected from competing plants. The importance of early shoot/root photoassimilate allocation and nodulation may differ according to soil nitrogen availability and moisture content.     

Effects of boron and calcium nutrition on the establishment of the Rhizobium leguminosarum–pea (Pisum sativum) symbiosis and nodule development under salt stress

The effects of modifying boron (B) and calcium (Ca²⁺) concentrations on the establishment and development of rhizobial symbiosis in Pisum sativum plants grown under salt stress were investigated. Salinity almost completely inhibited the nodulation of pea plants by Rhizobium leguminosarum bv. viciae 3841. This effect was prevented by addition of Ca²⁺ during plant growth. The capacity of root exudates derived from salt‐treated plants to induce Rhizobium nod genes was not significantly decreased. However, bacterial adsorption to roots was highly inhibited in plants grown with 75 mM NaCl. Moreover, R. leguminosarum 3841 did not grow in minimal media containing such salt concentration. High Ca²⁺ levels enhanced both rhizobial growth and adsorption to roots, and increased nodule number in the presence of high salt. Nevertheless, the nodules developed were not functional unless the B concentration was also increased. Because B has a strong effect on infection and cell invasion, these processes were investigated by fluorescence microscopy in pea nodules harbouring a R. leguminosarum strain that expresses green fluorescent protein. Salt‐stressed plants had empty nodules and only those treated with high B and high Ca²⁺ developed infection threads and exhibited enhanced cell and tissue invasion by Rhizobium. Overall, the results indicate that Ca²⁺ promotes nodulation and B nodule development leading to an increase of salt tolerance of nodulated legumes.     

TRANSLOCATION OF C14-LABELED PHOTOSYNTHATE IN NODULATED LEGUMES AS INFLUENCED BY NITRATE NITROGEN

Translocation of C¹⁴-labeled photosynthate was studied in nodulated pea and subterranean-clover plants which were grown either continuously without combined nitrogen or exposed to NaNO₃ in the nutrient solution for five days prior to C¹⁴O₂ assimilation. Supplying combined nitrogen decreased the proportion of photosynthate translocated to nodules with a corresponding increase in the proportion going to roots. Nodules on plants grown without combined nitrogen had a higher radioactivity than nodules on plants treated with sodium nitrate.     

Enhancing the early performance of the leguminous shrub Retama sphaerocarpa (L.) Boiss.: fertilisation versus Rhizobium inoculation

We have investigated the effect on growth of fertilisation versus biological nitrogen fixation by rhizobial nodules in Retama sphaerocarpa(L.) Boiss, a leafless leguminous shrub native to the Iberian Peninsula and North-West Africa that has generated interest for revegation of dry Mediterranean habitats. Our main objective was to optimise the formation of root nodules under nursery conditions and to evaluate their influence on the first year of seedling growth in comparison with standard fertilisation. Seedlings of R. sphaerocarpa from two Spanish localities were grown under two levels of fertilisation, and half of each were inoculated with rhizobia isolated from adult Retama, Cytisus and Adenocarpusplants in the field. Although some promiscuity was observed, nodulation was significantly successful with specific rhizobia. At the end of the experiment, highly fertilised plants were taller and heavier and exhibited larger photosynthetic rates than either nodulated or non-nodulated plants under low fertilisation. High fertilisation enhanced seedling growth but inhibited both the nodulation and the nitrogenase activity of the nodules. Thus, physiological differences between nodulated and non-nodulated plants were observed in the low but not in the high fertilisation treatment. Nitrogen uptake and use was enhanced by root nodules, which translated into enhanced photosynthesis and growth. Since inoculation is simple, environmentally friendly and cheap, and nodulated plants are more likely to overcome transplant stress than non-nodulated ones, our results suggest that inoculation together with low, background fertilisation (instead of high fertilisation) should be used when producing high quality seedlings of this autochthonous Mediterranean shrub.     

Phosphorus-induced micronutrient disorders in hybrid poplar

The effect of calcium on the nodulation of lucerne was studied using EGTA, a specific calcium-chelator. First, the effects of the chelator were tested on hydroponically grown plants at pH 7.0. Optimal numbers of nodules were obtained in nutrient solution containing 0.2 mM CaCl₂. When 0.4 mM EGTA was given additionally, nodulation was completely inhibited. Nodulation was restored specifically with CaCl₂, but not with MgCl₂. For studies in an acid soil (pH-H₂O 5.2), lucerne seedlings were grown in rhizotrons. 67% of the seedlings became nodulated when the soil around the seed was neutralized locally with 1.0 μmol of K₂CO₃ in drops of 12 μL volume. When native calcium was removed with 2 μmol of EGTA, nodulation was reduced to 12%. However, addition of EGTA to soil resulted in a drop of pH from 6.1 to 5.2. A phosphate buffer could also not keep soil-pH sufficiently stable. Such pH-decreases could be avoided by placing agar blocks containing 6 μmol of EGTA for three hours on freshly developed roots. This treatment reduced nodulation from 87% to 32%, with soil-pH lowering only from 6.2 to 6.0. Nodulation could be restored by adding 2 μmol of CaCl₂. The depletion of soil-calcium could depress nodule formation only during the first day after inoculation.     

The nodulation and nitrogen fixation of isolated roots of Phaseolus vulgaris L.

Summary The exposure of isolated bean roots to white light immediately before inoculation with nodule bacteria stimulated nodulation during a subsequent 7-day dark period. Strong white light applied to inoculated roots for 3 days or longer suppressed nodulation except when the light was applied immediately after inoculation and the duration of the light period did not exceed 3 days. Alternating periods of 12 hours light and 12 hours darkness were almost as inhibitory to nodulation as a continuous exposure to light. A combination of blue and far-red light inhibited nodule formation more than a combination of red and far-red light. Light treatments that inhibited nodulation also suppressed the production of lateral roots, but to a lesser extent. The effect of light on nodulation and lateral root formation was not influenced by the presence or absence of an attached hypocotyl segment in the explant.     

Involvement of cytokinin in the stimulation of nodulation by low concentrations of ammonium in Pisum sativum

Nodulation in pea (Pisum sativum L.) grown in hydroponic and sand culture systems is stimulated by low concentrations (<1.0 mM) of ammonium, but the physiological mechanisms underlying this stimulation are unknown. The current study involves a series of experiments, which investigate if the ammonium‐induced stimulation of nodulation involves changes in endogenous hormone (auxin and cytokinin) levels. P. sativum L. cv. Express was grown in growth pouches for 1 week with mineral N (0.5 and 2.0 mM NH₄⁺ or NO₃^–) or for 3 weeks exposed to exogenous indole‐3‐acetic acid (IAA) or 6‐benzylaminopurine (BAP) at a range of concentrations (10^‐9?10^‐5 M). Ammonium enhanced nodulation on the basis of both early whole plant (nodules plant^?1) and specific nodulation (nodules g^?1 root DW), especially in 0.5 mM treatment in which nodulation was approximately 4‐fold of the mineral‐N‐free control 1 week after inoculation. Correspondingly, the roots treated with ammonium contained much higher levels of t‐zeatin (Z) and lower t‐zeatin riboside (ZR) than that the control or nitrate‐treated plants. There was no significant difference in IAA levels between the control and ammonium treatments. Exogenous application of BAP for 3 weeks at concentrations of 10^‐7?10^‐5 M strongly inhibited nodulation. However, 10^?9 M BAP, but not IAA, significantly enhanced nodulation. These data support the theory that a relatively high ratio of cytokinin:auxin in roots is favourable for nodule initiation, but that an excessively high level of cytokinin inhibits nodulation. Based on these results we propose that stimulation of nodulation by low concentrations of ammonium may be mediated through increasing Z level in roots, which alters the balance of cytokinin and auxin, which in turn induces cortical cell divisions leading to nodule initiation.     

The ethylene-inhibitor aminoethoxyvinylglycine restores normal nodulation by Rhizobium leguminosarum biovar. viciae on Vicia sativa subsp. nigra by suppressing the ‘Thick and short roots’ phenotype

Nodulation of Vicia sativa subsp. nigra L. by Rhizobium bacteria is coupled to the development of thick and short roots (Tsr). This root phenotype as well as root-hair induction (Hai) and root-hair deformation (Had) are caused by a factor(s) produced by the bacteria in response to plant flavonoids. When very low inoculum concentrations (0.5–5 bacteria·ml^-1) were used, V. sativa plants did not develop the Tsr phenotype and became nodulated earlier than plants with Tsr roots. Furthermore, the nodules of these plants were located on the primary root in contrast to nodules on Tsr roots, which were all located at sites of lateral-root emergence. The average numbers of nodules per plant were not significantly different for these two types of nodulation. Root-growth inhibition and Hai, but not Had, could be mimicked by ethephon, and inhibited by aminoethoxyvinylglycine (AVG). Addition of AVG to co-cultures of Vicia sativa and the standard inoculum concentration of 5·10⁵ bacteria·ml^-1 suppressed the development of the Tsr phenotype and restored nodulation to the pattern that was observed with very low concentrations of bacteria (0.5–5 bacteria·ml^-1). The delay in nodulation on Tsr roots appeared to be caused by the fact that nodule meristems did not develop on the primary root, but only on the emerging laterals. The relationship between Tsr, Hai, Had, and nodulation is discussed.Abbreviations AVG aminoethoxyvinylglycine - cfu colonyforming units - Had root-hair deformation - Hai root-hair induction - NB naringenin-bacteria filtrate - Tsr Thick and short roots     

Plant genetic suppression of the non-nodulation phenotype of Rhizobium meliloti host-range nodH mutants: gene-for-gene interaction in the alfalfa-Rhizobium symbiosis?

Summary Rhizobium nodulation genes can produce active extracellular signals for legume nodulation. The R. meliloti host-range nodH gene has been postulated to mediate the transfer of a sulfate to a modified lipo-oligosaccharide, which in its sulfated form is a specific nodulation factor for alfalfa (Medicago sativa L.). We found that alfalfa was capable of effective nodulation with signal-defective and non-nodulating nodH mutants (Nnr) defining a novel gene-for-gene interaction that conditions nodulation. Bacteria-free nodules that formed spontaneously at about a 3–5% rate in unselected seed populations of alfalfa cv Vernal in the total absence of Rhizobium (Nar) exhibited all the histological, regulatory and ontogenetic characteristics of alfalfa nodules. Inoculation of such populations with nodH mutants, but not with nodA or nodC mutants, produced a four- to five-fold increase in the percentage of nodulated plants. Some 10–25% of these nodulated plants formed normal pink nitrogen-fixing nodules instead of white empty nodules. About 70% of the S₁ progeny of such Nnr⁺ plants retained the parental phenotype; these plants were also able to form nodules in the absence of Rhizobium. If selected Nar⁺ plants were self-pollinated almost the entire progeny exhibited the parental Nar⁺ phenotype. Segregation analysis of S₁ and S₂ progeny from selected Nar⁺ plants suggests that the Nar character is monogenic dominant and that the nodulation phenotype is controlled by a gene dose effect. The inoculation of different S₁ Nar⁺ progeny with nodH mutant bacteria gave only empty non-fixing nodules. Our results indicate that certain alfalfa genotypes can be selected for suppression of the non-nodulation phenotype of nodH mutants. The fact that the Nnr plant phenotype behaved as a dominant genetic trait and that it directly correlated with the ability of the selected plants to form nodules in the absence of Rhizobium suggests that the interaction of plant and bacterial alleles occurs early during signal transduction through the alteration of a signal reception component of the plant so that it responds to putative signal precursors.Abbreviations Nara Nodulation in the absence of Rhizobium - Nara nodulation with non-nodulating Rhizobium     

Nodulation gene regulation and quorum sensing control density-dependent suppression and restriction of nodulation in the Bradyrhizobium japonicum-soybean symbiosis

The nodulation of Glycine max cv. Lambert and the nodulation-restricting plant introduction (PI) genotype PI 417566 by wild-type Bradyrhizobium japonicum USDA110 is regulated in a population-density-dependent manner. Nodulation on both plant genotypes was suppressed (inhibited) when plants received a high-density inoculum (10(9) cells/ml) of strain USDA110 grown in complex medium, and more nodules were produced on plants receiving a low-cell-density inoculum (10(5) cells/ml). Since cell-free supernatants from strain USDA110 grown to high cell density in complex medium decreased the expression of an nodY-lacZ fusion, this phenomenon was attributed to bradyoxetin-induced repression of nod gene expression. Inoculation of either the permissive soybean genotype (cv. Lambert) or PI 417566 with 10(9) cells/ml of the nodD2, nolA, nodW, and nwsB mutants of USDA110 enhanced nodulation (up to 24%) relative to that seen with inoculations done with 10(5) cells/ml of the mutants or the wild-type strain, indicating that these genes are involved in population-density-dependent nodulation of soybeans. In contrast, the number of nodules produced by an nodD1 mutant on either soybean genotype was less than those seen with the wild-type strain inoculated at a low inoculum density. The nodD2 mutant outcompeted B. japonicum strain USDA123 for nodulation of G. max cv. Lambert at a high or low inoculum density, and the results of root-tip-marking and time-to-nodulate studies indicated that the nolA and nodD2 mutants nodulated this soybean genotype faster than wild-type USDA110. Taken together, the results from these studies indicate that the nodD2 mutant of B. japonicum may be useful to enhance soybean nodulation at high inoculum densities and that NodD2 is a key repressor influencing host-controlled restriction of nodulation, density-dependent suppression of nodulation, perception of bradyoxetin, and competitiveness in the soybean-B. japonicum symbiosis.     

Photosynthate partitioning and nodule formation in soybean plants that received red or far-red light at the end of the photosynthetic period

The influence of row orientation on spectral distribution of light received by growing soybean [ Glycine max (L.) Merr. (cv. Coker 338)] plants was measured under field conditions, and light spectrum effects on photosynthate partitioning were studied under controlled environments. Light received by leaves under field conditions differed among those grown in north-south vs east-west oriented rows. In morning and late afternoon, the far-red/ red ratio received by leaves at the surface of the canopy differed about 3-fold from the east to west sides of north-south rows, but only 1.3-fold from the south to north sides of east-west rows.
In controlled environments, brief exposures to red or far-red light at the end of the photosynthetic period influenced partitioning of photosynthate among leaves, stems and roots. The top/root ratios differed significantly between the red and far-red treated plants. Red treated plants partitioned less photosynthate to stems and more to roots than did those treated with far-red. Also, plants with larger root systems developed more nodules. Phytochrome effects on photosynthate partitioning between tops and roots may influence yield of soybean plants grown in soils with low water-holding capacity.     

Manipulation of Ethylene Synthesis in Roots Through Bacterial ACC Deaminase for Improving Nodulation in Legumes

Symbiotic association between rhizobia and legumes results in the development of unique structures on roots, called nodules. Nodulation is a very complex process involving a variety of genes that control NOD factors (bacterial signaling molecules), which are essential for the establishment, maintenance and regulation of this process and development of root nodules. Ethylene is an established potent plant hormone that is also known for its negative role in nodulation. Ethylene is produced endogenously in all plant tissues, particularly in response to both biotic and abiotic stresses. Exogenous application of ethylene and ethylene-releasing compounds are known to inhibit the formation and functioning of nodules. While inhibitors of ethylene synthesis or its physiological action enhance nodulation in legumes, some rhizobial strains also nodulate the host plant intensively, most likely by lowering endogenous ethylene levels in roots through their 1-aminocyclopropane-1-carboxylate (ACC) deaminase activity. Co-inoculation with ACC deaminase containing plant growth promoting rhizobacteria plus rhizobia has been shown to further promote nodulation compared to rhizobia alone. Transgenic rhizobia or legume plants with expression of bacterial ACC deaminase could be another viable option to alleviate the negative effects of ethylene on nodulation. Several studies have well documented the role of ethylene and bacterial ACC deaminase in development of nodules on legume roots and will be the primary focus of this critical review.     

A comparison of the nodulation of diploid and tetraploid varieties of red clover inoculated with different Rhizobial strains

Summary The effectiveness of three different strains ofRhizobium trifolii on a diploid and a tetraploid variety of red clover (Trifolium pratense) grown in aseptic mineral-agar culture was compared. It was found that with two of the rhizobial strains tested, the tetraploid plants were slower to nodulate than the diploid plants, but that a hundred per cent nodulation was obtained in both cases. With the third strain, however, a sixty-six per cent nodulation of the diploid plants was reduced to an eight per cent nodulation of the tetraploids.In general, over a given period, fewer nodules were formed on the tetraploid plants, but these tended to be larger than on the diploids. The fresh weight of the tetraploid plants always exceeded that of the diploids, but less so in cases of poorly nodulated plants.     

Medicago sativa - Sinorhizobium meliloti Symbiosis Promotes the Bioaccumulation of Zinc in Nodulated Roots

In this study we investigated effects of Zn supply on germination, growth, inorganic solutes (Zn, Ca, Fe, and Mg) partitioning and nodulation of Medicago sativa This plant was cultivated with and without Zn (2 mM). Treatments were plants without (control) and with Zn tolerant strain (S532), Zn intolerant strain (S112) and 2 mM urea nitrogen fertilisation. Results showed that M. sativa germinates at rates of 50% at 2 mM Zn. For plants given nitrogen fertilisation, Zn increased plant biomass production. When grown with symbionts, Zn supply had no effect on nodulation. Moreover, plants with S112 showed a decrease of shoot and roots biomasses. However, in symbiosis with S532, an increase of roots biomass was observed. Plants in symbiosis with S. meliloti accumulated more Zn in their roots than nitrogen fertilised plants. Zn supply results in an increase of Ca concentration in roots of fertilised nitrogen plants. However, under Zn supply, Fe concentration decreased in roots and increased in nodules of plants with S112. Zn supply showed contrasting effects on Mg concentrations for plants with nitrogen fertilisation (increase) and plants with S112 (decrease). The capacity of M. sativa to accumulate Zn in their nodulated roots encouraged its use in phytostabilisation processes.     

Far-Red Light Reflection from Green Leaves and Effects on Phytochrome-Mediated Assimilate Partitioning under Field Conditions

The influence of plant spacing and row orientation on spectral distribution of light received by growing soybean (Gylcine max [L.] Merr.) plants was measured under field conditions. Light absorption, reflection and transmission of individual leaves showed that most of the blue and red was absorbed while most of the far-red was either reflected or transmitted. Plants growing in the field received different ratios of far-red relative to red, depending on nearness and/or orientation of other vegetation. Plants grown in close-spaced rows, or high population densities, received higher far-red/red ratios than did those grown in wide rows, or sparse populations. Heliotropic movements of the leaves also contributed to the far-red reflection patterns associated with row orientation. Under field conditions, differences in far-red/red ratios associated with nearness of competing vegetation became more pronounced with low solar angle near the end of the day. Plants exposed to far-red for 5 minutes at the end of each day in controlled environments, and those grown in close-spaced rows in the field, developed longer internodes and fewer branches. Red, far-red photoreversibility in the controlled environment study indicated involvement of phytochrome. Dry matter partitioning among plant components in the field was related to far-red/red light ratio received during growth and development.     

Rhizobium leguminosarum strains forming nitrogen-rich nodules inPisum sativum

The nodules which developed on the roots ofPisum sativum after inoculation withRhizobium leguminosarum bv.viciae strains showing a Hup⁺ symbiotic phenotype and/or a high relative efficiency of electron transfer to dinitrogen (RE) had a high nitrogen content of their tissue. In comparison with the nodules initiated by a strain possessing the Hup⁻ symbiotic phenotype or by an indigenous soilRhizobium population, the nitrogen-rich root nodules contained up to four times more nitrogen (9.2% of dry mass). In the nitrogen-rich nodules, total amino acid, and especially Ala, Lys and Phe contents were significantly increased. The nitrogen-rich nodule symbiotic phenotype (Nrn) was well expressed, irrespective of pea cultivars and host plant age. If a strain showing the Nrn phenotype was applied in double-strain inocula, a significant correlation was found between increasing rates of the strain and increasing percentage nitrogen content of nodule tissue in the nodulated plants.     

Signaling and Gene Expression for Water-Tolerant Legume Nodulation

In the symbiotic interaction with rhizobia, legumes develop nodules in which nitrogen fixation takes place. Upon submersion, most temperate legumes are incapable of nodulation, but tropical legumes that grow in waterlogged soils have acquired water stress tolerance for growth and nodulation. One well-studied model plant, the tropical, semi-aquatic Sesbania rostrata, develops stem-located adventitious root primordia that grow out into adventitious roots upon submergence and develop into stem nodules after inoculation with the microsymbiont, Azorhizobium caulinodans. Sesbania rostrata also has a nodulated underground root system. On well-aerated roots, nodules form via root hair curling infection in the zone, just above the root tip, where root hairs develop; on hydroponic roots, an alternative process is used, recruiting a cortical intercellular invasion program at the lateral root bases that skips the epidermal responses. This intercellular cortical invasion entails infection pocket formation, a process that involves cell death features and reactive oxygen species. The plant hormones ethylene and gibberellin are the major signals that act downstream from the bacterial nodulation factors in the nodulation and invasion program. Both hormones block root hair curling infection, but cooperate to stimulate lateral root base invasion and play a role in infection thread formation, meristem establishment, and differentiation of meristem descendants.     

Mycorrhizal improvement in non-leguminous nitrogen fixing associations with particular reference toHippophaë rhamnoides L.

Summary The roots ofHippophaë rhamnoides which regularly bear actinomycete induced nodules when growing on Scottish sand dunes have also been found to support an endomycorrhizal association withGlomus fasciculatus. Ultrastructural and cytochemical studies carried out on the indigenous infections of establishedHippophaë mycorrhizal roots would support the postulate that transport is indeed occurring between the fungal symbiont and the host plant and vice versa in respect of phosphate and carbohydrate. Experiments using various inoculation regimes, demonstrated the significant improvement in the mycorrhizal/nodulated plants compared to the nodulated-only and the mycorrhizal-only plants with respect to plant growth, uptake of phosphate and nitrogenase activity, when grown in a medium poor in combined nitrogen and soluble phosphate. Preliminary work onAlnus andMyrica species growing in Central Scotland indicates that the mycorrhizae associated with these nodulated root systems exhibit a different interaction pattern which may be dependent on habitat type and associated angiosperm species.     

Soil and environmental factors related to nodulation in Cowania and Purshia

Summary Nitrogen fixing trees and shrubs may be useful in revegetation efforts. The possibility that soil and environmental factors may influence a soil's capability to produce nodulated seedlings was explored.Purshia tridentata andCowania mexicana var. Stansburiana seedlings were grown in greenhouse trials using ten soils from native sites for each of the two genera. Treatments included a control and a six mmole nitrogen amendment as NH₄NO₃ for both surface and subsurface samples. Nodulation was often sparse for seedlings grown in surface collected samples. Although nodulation was usually better in subsoil samples, even some subsoils produced few or no nodules. Nitrogen additions inhibit nodulation and although soil nitrogen may be inhibitory in some unamended surface soils it is probably not a general cause of sparse nodulation. Nodule masses showed the same trends as nodule number but varied less with treatment and depth of soil source. Seedlings compensated for sparse nodulation with an increase in mass per nodule. Incidence of nodulation was related to some soil and environmental factors. Multiple regression analysis explained a substantial portion of nodulation variability. Soils from lower elevations with less precipitation did not produce well nodulated seedlings even in well watered greenhouse trials. Micronutrient cations, potassium, and phosphorus are positively correlated with nodulation incidence. The two genera were generally similar in nodulation responses to soil and environmental factors.Support for portions of this study was provided by the National Science Foundation (PCM-8204885). Any opinions, findings conclusions or recommendations expressed are those of the authors and do not necessarily reflect the views of the sponsor. Oregon Agricultural Experiment Station technical paper number 7293.