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PubMed1975

Sodium-1,2-C Acetate Incorporation in Roots of Frost-hardy and Less Hardy Alfalfa Varieties under Hardening Conditions.

When the temperature of incorporation of sodium acetate-1, 2-(14)C into lipids of alfalfa (Medicago media Pers. var. Rambler and Medicago sativa L. var. Caliverde) roots was lowered from 22 C to 1 C, elongation and desaturation of fatty acids and the labeling of phosphatidylcholine were strongly stimulated.Controlled hardening of alfalfa stimulated strongly the incorporation of sodium acetate-1, 2-(14)C into root lipids of the hardy variety Rambler, but only slightly in the case of the frost-sensitive variety Caliverde. When incorporation was done at 1 C at various times of hardening, labeling decreased significantly in linoleic acid with a corresponding increase in oleic acid. Hardening, therefore, repressed specifically the initial low temperature stimulation of oleic acid desaturation, without affecting the stimulation of elongation of palmitic acid and the desaturation of stearic acid at low temperature. The radioactivity in linoleic acid was slightly greater in hardy Rambler than in Caliverde throughout hardening.When feedings were done at 1 C at various times of hardening, labeling of phosphatidylcholine increased in Rambler while it decreased in Caliverde. Throughout the hardening period, when incorporation was done at 1 C, linoleic acid represented a higher percentage of the label in phosphatidylcholine than in phosphatidylethanolamine or triglycerides and its specific radioactivity was much greater in phosphatidylcholine than in phosphatidylethanolamine and triglycerides and in Rambler than in Caliverde. Phosphatidylcholine seems, therefore, to play a special part in linoleic acid synthesis and in its control during the acquisition of frost hardiness.

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PubMed1977

A possible new nucleoside diphosphatase from the cytosol of soybean and alfalfa nodules.

Nucleoside diphosphatase activity has been found in the cytosol of soybean (Glycine max) and alfalfa (Medicago satavia) nodules. The enzyme differs from other diphosphatases in that it does not exhibit a strong preference for one nucleoside diphosphate over another. Very little, if any, diphosphatase activity was detected in root extracts of alfalfa and soybean seedlings.

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PubMed1978

Development of Bacteroids in Alfalfa (Medicago sativa) Nodules.

The morphology, acetylene reduction capability, and nucleic acid content of bacteroids in different regions of alfalfa (Medicago sativa var. Buffalo) nodules were studied by electron microscopy, gas chromatography, and laser flow microfluorometry, respectively. Bacteroids in the nodule tips were small (1 to 2.5 micrometers in length), had low nucleic acid content, and contained distinct central nucleoids. These bacteroids were comparatively inactive in acetylene reduction in situ. Bacteroids in the middle regions of alfalfa nodules were greatly enlarged (5 to 7 micrometers in length), had relatively high nucleic acid content, and did not possess central nucleoids. The bacteroids were very active in acetylene reduction. Bacteroids in the basal nodule region also were enlarged and without distinct nucleoid regions, but had relatively low nucleic acid content and low in situ acetylene-reducing activity.

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PubMed1982

Ploidy Effects in Isogenic Populations of Alfalfa : III. Chloroplast Thylakoid-Bound Coupling Factor 1 in Protoplasts and Leaves.

The influence of polyploidization on chloroplast coupling factor 1 (CF1) was examined in leaves and leaf protoplasts from isogenic diploid-tetraploid (DDC 2X-4X) and tetraploid-octoploid (IC 4X-8X) sets of alfalfa (Medicago sativa L.). Alfalfa CF1 was purified to homogeneity and was found to contain five subunits with molecular weights of 57,900, 54,300, 38,700, 23,100, and 15,200. Rocket immunoelectrophoresis with anti-spinach CF1 gamma immunoglobulins was used to quantify CF1 from protoplasts and leaves. In the DDC 2X-4X set, fresh weight per cell and cellular content of CF1, chlorophyll (Chl), and DNA doubled with ploidy. Ratios of CF1 to Chl of leaf protoplasts and leaves were similar between diploids and tetraploids. In the IC 4X-8X set, octoploid protoplasts were 90% higher in Chl than were comparable tetraploids, whereas octoploids were 50 to 60% higher than tetraploids in fresh weight per cell and cellular content of CF1 and DNA. Concentrations of CF1 and Chl in leaves and ratios of CF1 to DNA in protoplasts were similar across ploidy levels of both isogenic sets. Therefore, cellular content of CF1 increases proportionately with the amount of DNA per cell or gene dosage.

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PubMed1983

Ploidy Effects in Isogenic Populations of Alfalfa (Medicago sativa L.) : IV. Similarity in Physical and Kinetic Properties of Ribulose-1,5-bisphosphate Carboxylase/Oxygenase.

Ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCase) was isolated from isogenic diploid-tetraploid and tetraploid-octoploid sets of alfalfa (Medicago sativa L.) leaves. Molecular weights of RuBPCase subunits were similar across ploidy levels of both isogenic sets with subunits of 52,000 and 14,000. Apparent K(m)(CO(2)) values and substrate specificity factors (V(c)K(o)/V(o)K(c)) of RuBPCase were similar across ploidy levels of both isogenic sets. These results indicate that ploidy had no effect on the kinetic properties of RuBPCase in alfalfa.

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PubMed1984

Alfalfa Root Nodule Carbon Dioxide Fixation : II. Partial Purification and Characterization of Root Nodule Phosphoenolpyruvate Carboxylase.

A nonphotosynthetic phosphoenolpyruvate carboxylase (EC 4.1.1.31) was partially purified from the cytosol of root nodules of alfalfa. The enzyme was purified 86-fold by ammonium sulfate fractionation, DEAE-cellulose, hydroxylapatite chromatography, and reactive agarose with a final yield of 32%. The enzyme exhibited a pH optimum of 7.5 with apparent K(m) values for phosphoenolpyruvate and magnesium of 210 and 100 micromolar, respectively. Two isozymes were resolved by nondenaturing polyacrylamide disc gel electrophoresis. Subsequent electrophoresis of these isozymes in a second dimension by sodium dodecyl sulfate slab gel electrophoresis yielded identical protein patterns for the isozymes with one major protein band at molecular weight 97,000. Malate and AMP were slightly inhibitory (about 20%) to the partially purified enzyme. Phosphoenolpyruvate carboxylase comprised approximately 1 to 2% of the total soluble protein in actively N(2)-fixing alfalfa nodules.

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PubMed1985

Photosynthate supply and utilization in alfalfa : a developmental shift from a source to a sink limitation of photosynthesis.

Long-term carbon dioxide enrichment, (14)CO(2) feeding, and partial defoliation were employed as probes to investigate source/sink limitations of photosynthesis during the development of symbiotically grown alfalfa. In the mature crop, long-term CO(2) enrichment does not affect the rates of net photosynthesis, relative growth, (14)C export to nonphotosynthetic organs, or the rates of (14)C label incorporation into leaf sucrose, starch, or malate. The rate of glycolate labeling is, however, substantially reduced under these conditions. When the mature crop was partially defoliated, a considerable increase in net photosynthesis occurred in the remaining leaves. In the seedling crop, long-term CO(2) enrichment increased dry matter accumulation, primarily as a result of increases in leaf starch content. Although the higher rates of starch synthesis are not maintained, the growth enhancement of the enriched plants persisted throughout the experimental period. These results imply a source limitation of seedling photosynthesis and a sink limitation of photosynthesis in more mature plants. Consequently, both the supply and the utilization of photosynthate may limit seasonal photosynthesis in alfalfa.

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PubMed1985

Interorgan photosynthate partitioning in alfalfa.

The net partitioning of current photosynthate among vegetative organs of nodulated alfalfa (Medicago sativa L.) was investigated by determining radiolabel distribution from upper and lower source leaves, at different times of day, and during successively longer chase periods. Photosynthate was exported sooner and more completely by a fully expanded lower than by a fully expanded upper source leaf. The radiolabel pulse from the lower source leaf peaked in the main stem within 1.5 hours, in the crown and nodules after 3 hours, in the unexpanded leaves and apex of the main stem after 6 hours, and in the shoots growing from leaf axils on the main stem after 24 hours. The results suggest that the crown, apex, axillary shoots, and nodules both imported and mobilized photosynthate originating at a lower source leaf, while roots and shoots growing from the crown showed net accumulation. The pulse from the upper source leaf was initially rapidly exported by the main stem and imported by the root between 1.5 and 3 hours, but there was no net change of label content of these organs during the ensuing 21 hours. Rapidly growing organs had the highest concentrations, and the largest organs had the highest content of radiolabel. These results provide new information about the accumulation and circulation of photosynthate within the alfalfa plant.

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PubMed1985

Regeneration in alfalfa tissue culture: stimulation of somatic embryo production by amino acids and N-15 NMR determination of nitrogen utilization.

The production of somatic embryos in alfalfa (Medicago sativa L., cv Regen S) is increased 5- to 10-fold by alanine and proline. However, utilization of nitrogen for synthesis of protein from alanine, proline, glutamate, and glycine is not qualitatively different, even though the latter two amino acids do not increase somatic embryo formation. These determinations were made by (15)N labeling with detection by nuclear magnetic resonance. Overall metabolism of the nitrogen of proline, alanine, glutamate, and glycine is also similar in two regenerating and nonregenerating genotypes with similar germplasm, except that the levels of free amino acids are consistently higher in the nonregenerating line. In addition, when regeneration is suppressed in either of the two regenerating lines, the level of intracellular free amino acids increases. This increased level of metabolites is the only direct evidence provided by analysis of nitrogen metabolism of differences between the regenerating and nonregenerating states in alfalfa.

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PubMed1987

Nonphotosynthetic CO(2) Fixation by Alfalfa (Medicago sativa L.) Roots and Nodules.

The dependence of alfalfa (Medicago sativa L.) root and nodule nonphotosynthetic CO(2) fixation on the supply of currently produced photosynthate and nodule nitrogenase activity was examined at various times after phloem-girdling and exposure of nodules to Ar:O(2). Phloemgirdling was effected 20 hours and exposure to Ar:O(2) was effected 2 to 3 hours before initiation of experiments. Nodule and root CO(2) fixation rates of phloem-girdled plants were reduced to 38 and 50%, respectively, of those of control plants. Exposure to Ar:O(2) decreased nodule CO(2) fixation rates to 45%, respiration rates to 55%, and nitrogenase activities to 51% of those of the controls. The products of nodule CO(2) fixation were exported through the xylem to the shoot mainly as amino acids within 30 to 60 minutes after exposure to (14)CO(2). In contrast to nodules, roots exported very little radioactivity, and most of the (14)C was exported as organic acids. The nonphotosynthetic CO(2) fixation rate of roots and nodules averaged 26% of the gross respiration rate, i.e. the sum of net respiration and nonphotosynthetic CO(2) assimilation. Nodules fixed CO(2) at a rate 5.6 times that of roots, but since nodules comprised a small portion of root system mass, roots accounted for 76% of the nodulated root system CO(2) fixation. The results of this study showed that exposure of nodules to Ar:O(2) reduced nodule-specific respiration and nitrogenase activity by similar amounts, and that phloem-girdling significantly reduced nodule CO(2) fixation, nitrogenase activity, nodule-specific respiration, and transport of (14)C photoassimilate to nodules. These results indicate that nodule CO(2) fixation in alfalfa is associated with N assimilation.

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PubMed1988

Abscisic Acid-regulated gene expression in relation to freezing tolerance in alfalfa.

A comparison of abscisic acid (ABA)-induced and cold-acclimation-induced freezing tolerance in two alfalfa cultivars (Medicago falcata cv Anik and Medicago sativa v Trek) indicates that ABA alone can increase freezing tolerance to some extent, but for the development of maximum tolerance, cold acclimation is essential. Analysis of in vivo-labeled proteins of ABA-treated seedlings reveals that ABA causes several changes in the pattern of protein synthesis. While some of these changes appear to be similar to those induced by cold acclimation, others seem to be specific to ABA treatment. From a cDNA library constructed against poly(A(+)) RNA of a freezing-tolerant alfalfa cultivar, Anik, a cDNA clone, pSM1409, has been isolated. Expression of the gene corresponding to this clone, as determined by northern hybridization, is regulated most likely at the transcriptional level by cold acclimation and exogenously supplied ABA. However, the increase in the transcript level is much greater in the freezing-tolerant cultivar Anik than in the relatively freezing-sensitive cultivar, Trek. The role of ABA in the acquisition of freezing tolerance is discussed.

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PubMed1988

Alfalfa Root Exudates and Compounds which Promote or Inhibit Induction of Rhizobium meliloti Nodulation Genes.

Using a plate induction assay, we demonstrate that alfalfa exudes inducer of Rhizobium meliloti nodulation genes. The inducer is exuded from the infectible zone of the root, accumulates to at least 1 micromolar, and is not affected by 10 millimolar nitrate. No zones of inhibition are observed. A nodulation minus mutant line of alfalfa, MN-1008, exudes normal levels of inducer. R. meliloti grown in rich medium requires ten-fold higher concentrations of luteolin to achieve half-maximal induction as compared to cells grown in a minimal medium. Flavonoids other than luteolin are found to have activity in R. meliloti nodulation gene induction assays. The compounds apigenin and eriodictyol have activities two-fifths and one-seventh that of luteolin, respectively. Several of the flavonoids tested (morin = naringenin > kaempferol = chrysin > quercetin = fisetin = hesperitin) demonstrate antagonistic activity toward induction by luteolin. The most effective antagonist is the coumarin, umbelliferone.

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PubMed1989

Molecular cloning and relationship to freezing tolerance of cold-acclimation-specific genes of alfalfa.

Cold-acclimation-specific (CAS) gene expression has been examined by screening a cDNA library prepared from poly(A)(+) RNA of cold-acclimated seedlings of a freezing-tolerant variety of alfalfa (Medicago falcata cv Anik). Three CAS cDNA clones, pSM784, pSM2201, and pSM2358, representing different sequence species, have been used to investigate the relative abundance and time-course of accumulation of corresponding transcripts. Results obtained show that the expression of these CAS genes is regulated in a coordinated manner most likely at the level of transcription. The expression of genes, as measured by mRNA abundance corresponding to the three CAS cDNA clones, is not stimulated or induced by heat shock, water stress, abscisic acid, or wounding. A positive correlation is observed between the expression of these cloned sequences and the degree of freezing-tolerance in four alfalfa cultivars.

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PubMed1989

Differential Histone Acetylation in Alfalfa (Medicago sativa) Due to Growth in NaCl : Responses in Salt Stressed and Salt Tolerant Callus Cultures.

The steady state distribution of histone variant proteins and their modifications by acetylation were characterized in wild type and salinity stress adapted alfalfa (Medicago sativa). Isotopic labeling detected dynamic acetylation at four sites in the histone H3 variants and five sites in histones H4 and H2B. Histone variant H3.2 was the most highly acetylated histone with 25% higher steady state acetylation and a two- to threefold higher acetylation labeling than histone H3.1. Histone phosphorylation was limited to histone variants H1.A, H1.B, and H1.C and to histone H2A.3, which was also acetylated. Histone variant composition was unaffected by cellular exposure to NaCl. Histone acetylation was qualitatively similar in salt-tolerant and salt-sensitive cells under normal growth conditions. However, short term salt stress in salt sensitive cells or continued growth at 1% NaCl in salt tolerant cells led to major increases in the multiacetylated forms of histone H4 and the two variants of histone H3. These changes were more pronounced in the diploid than in the tetraploid alfalfa strains. The increase in multiacetylation of core histones serves as an in vivo reporter suggesting an altered intranuclear ionic environment in the presence of salt. It may also represent an adaptive response in chromatin structure to permit chromatin function in a more saline intranuclear environment.

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PubMed1989

Purification and Characterization of NADH-Glutamate Synthase from Alfalfa Root Nodules.

Glutamate synthase (GOGAT), a key enzyme in the pathway for the assimilation of symbiotically fixed dinitrogen (N(2)) into amino acids in alfalfa (Medicago sativa L.) root nodules, was purified and used to produce high titer polyclonal antibodies. Purification resulted in a 208-fold increase in specific activity to 13 micromole per minute per milligram of protein and an activity yield of 37%. Further purification to near homogeneity was achieved by fast protein liquid chromatography, but with substantial loss of activity. Enzymic activity was highly labile, losing 3% per hour even when substrates, stabilizers, and reducing agents were included in buffers. However, activity could be partially stabilized for up to 1 month by storing GOGAT at -80 degrees C in 50% glycerol. The subunit molecular weight of GOGAT was estimated at 200 +/- 7 kilodaltons with a native molecular weight of 235 +/- 16 kilodaltons, which suggested that GOGAT is a monomer of unusually high molecular weight. The pl was estimated to be 6.6. The K(m) values for glutamine, alpha-ketoglutarate, and NADH were 466, 33, and 4.2 micromolar, respectively. Antibodies were produced to NADH-GOGAT. Specificity of the antibodies was shown by immunotitration of GOGAT activity. Alfalfa nodule NADH-GOGAT antibodies cross-reacted with polypeptides of a similar molecular weight in a number of legume species. Western blots probed with anti-GOGAT showed that the high GOGAT activity of nodules as compared to roots was associated with increased levels of GOGAT polypeptides. Nodule NADH-GOGAT appeared to be highly expressed in effective nodules and little if any in other organs.

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PubMed1989

Aspartate aminotransferase in alfalfa root nodules : I. Purification and partial characterization.

Aspartate aminotransferase (l-aspartate:2-oxoglutarate aminotransferase, EC 2.6.1.1 [AAT]), a key enzyme in the assimilation of C and N compounds, was purified from the cytosol of alfalfa (Medicago sativa L.) root nodules. Isoforms that increased during nodule development, AAT-2a, AAT-2b, and AAT-2c, were purified greater than 447-fold to apparent homogeneity, and high titer polyclonal antibodies were produced. The native molecular weight of the AAT-2 isoforms was approximately 80 kilodatons with a subunit molecular weight of 40 kilodatons, indicating that the holoenzymes are dimers. The AAT-2 isoforms comprised approximately 0.4% of the total soluble nodule protein. The AAT specific activity was measured in leaf, stem, root, and nodule organs, and zymograms of each were compared. Enzyme activity was 4- to 37-fold greater in effective (nitrogen fixing) nodules than in leaves, stems, and roots. Effective nodule AAT-specific activity was 3- to 8-fold greater than that of plant-controlled ineffective nodules. No differences in K(m) were observed between AAT-1 and AAT-2. Antibodies raised against AAT-2 were more selective against AAT-2 than AAT-1. Evidence obtained from zymograms suggests that the expression of alfalfa nodule AAT is controlled at two different gene loci, AAT-1 and AAT-2, resulting in different dimeric isoforms.

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PubMed1989

Interactions among Flavonoid nod Gene Inducers Released from Alfalfa Seeds and Roots.

Alfalfa (Medicago sativa L.) seeds and roots can create complex rhizosphere effects by releasing flavonoids that induce nodulation (nod) genes in Rhizobium meliloti. Previous reports identified luteolin and 4,4'-dihydroxy-2'-methoxychalcone as strong inducers that are released from seeds and roots, respectively, and 4',7-dihydroxyflavone and 4',7-dihydroxyflavanone as weaker inducers which are exuded by roots. As a first step toward identifying flavonoid interactions that may occur in the rhizosphere, combinations of these molecules were tested for transcriptional effects on a nodABC-lacZ fusion in R. meliloti. At low concentrations (e.g. 8.4 nanomolar), interactions of the three nod gene inducers from root exudate were additive. When the strong inducers 4,4'-dihydroxy-2'-methoxychalcone and luteolin were present separately at higher concentrations (e.g. 21 nanomolar), their effect could be decreased significantly by the weaker inducers 4',7-dihydroxyflavone and 4',7-dihydroxyflavanone. In contrast, when low concentrations of luteolin from seed rinses and 4,4'-dihydroxy-2'-methoxychalcone from root exudate were present together, they produced synergistic increases in nod gene transcription. Tests with mixtures of the three nod gene inducers from root exudate indicated that alfalfa seedlings might easily decrease the strong inductive effect of the chalcone by releasing modest amounts of the weaker inducers. In addition, mixtures of luteolin and the nod gene inducers in root exudate suggested that interactions between nod gene inducers from seeds and roots may create a zone highly favorable to root nodule formation near the top of the primary root.

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PubMed1989

Nitrogen Assimilating Enzyme Activities and Enzyme Protein during Development and Senescence of Effective and Plant Gene-Controlled Ineffective Alfalfa Nodules.

Effective (N(2)-fixing) alfalfa (Medicago sativa L.) and plant-controlled ineffective (non-N(2)-fixing) alfalfa recessive for the in(1) gene were compared to determine the effects of the in(1) gene on nodule development, acetylene reduction activity (ARA), and nodule enzymes associated with N assimilation and disease resistance. Effective nodule ARA reached a maximum before activities of glutamine synthetase (GS), glutamate synthase (GOGAT), aspartate aminotransferase (AAT), asparagine synthetase (AS), and phosphoenolpyruvate carboxylase (PEPC) peaked. Ineffective nodule ARA was only 5% of effective nodule ARA. Developmental profiles of GS, GOGAT, AAT, and PEPC activities were similar for effective and ineffective nodules, but activities in ineffective nodules were lower and declined earlier. Little AS activity was detected in developing ineffective nodules. Changes in GS, GOGAT, AAT, and PEPC activities in developing and senescent effective and ineffective nodules generally paralleled amounts of immunologically detectable enzyme polypeptides. Effective nodule GS, GOGAT, AAT, AS, and PEPC activities declined after defoliation. Activities of glutamate dehydrogenase, malate dehydrogenase, phenylalanine ammonia lyase, and caffeic acid-o-methyltransferase were unrelated to nodule effectiveness. Maximum expression of nodule N-assimilating enzymes appeared to require the continued presence of a product associated with effective bacteroids that was lacking in in(1) effective nodules.

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