Common bean (Phaseolus vulgaris) has become a cosmopolitan crop, but was originally domesticated in the Americas and has been grown in Latin America for several thousand years. Consequently an enormous diversity of bean nodulating bacteria have developed and in the centers of origin the predominant species in bean nodules is R. etli. In some areas of Latin America, inoculation, which normally promotes nodulation and nitrogen fixation is hampered by the prevalence of native strains. Many other species in addition to R. etli have been found in bean nodules in regions where bean has been introduced. Some of these species such as R. leguminosarum bv. phaseoli, R. gallicum bv. phaseoli and R. giardinii bv. phaseoli might have arisen by acquiring the phaseoli plasmid from R. etli. Others, like R. tropici, are well adapted to acid soils and high temperatures and are good inoculants for bean under these conditions. The large number of rhizobia species capable of nodulating bean supports that bean is a promiscuous host and a diversity of bean-rhizobia interactions exists. Large ranges of dinitrogen fixing capabilities have been documented among bean cultivars and commercial beans have the lowest values among legume crops. Knowledge on bean symbiosis is still incipient but could help to improve bean biological nitrogen fixation.

Diversity of Rhizobium-Phaseolus vulgaris symbiosis: overview and perspectives

Biological N2 fixation represents the major source of N input in agricultural soils including those in arid regions. The major N2-fixing systems are the symbiotic systems, which can play a significant role in improving the fertility and productivity of low-N soils. The Rhizobium-legume symbioses have received most attention and have been examined extensively. The behavior of some N2-fixing systems under severe environmental conditions such as salt stress, drought stress, acidity, alkalinity, nutrient deficiency, fertilizers, heavy metals, and pesticides is reviewed. These major stress factors suppress the growth and symbiotic characteristics of most rhizobia; however, several strains, distributed among various species of rhizobia, are tolerant to stress effects. Some strains of rhizobia form effective (N2-fixing) symbioses with their host legumes under salt, heat, and acid stresses, and can sometimes do so under the effect of heavy metals. Reclamation and improvement of the fertility of arid lands by application of organic (manure and sewage sludge) and inorganic (synthetic) fertilizers are expensive and can be a source of pollution. The Rhizobium-legume (herb or tree) symbiosis is suggested to be the ideal solution to the improvement of soil fertility and the rehabilitation of arid lands and is an important direction for future research.

Rhizobium-Legume Symbiosis and Nitrogen Fixation under Severe Conditions and in an Arid Climate

Introduction, Allen V. Barker and David J. Pilbeam Essential Elements-Macronutrients Nitrogen, Allen V. Barker and Gretchen M. Bryson Phosphorus, Charles A. Sanchez Potassium, Konrad Mengel Calcium, David J. Pilbeam and Philip S. Morley Magnesium, Donald J. Merhaut Sulfur, Silvia Haneklaus, Elke Bloem, Ewald Schnug, Luit J. de Kok and Ineke Stulen Essential Elements-Micronutrients Boron, Umesh C. Gupta Chlorine, Joseph R. Heckman Copper, David E. Kopsel and Dean A. Kopsell Iron, Volker Roemheld and Miroslav Nikolic Manganese, Julia M. Humphries, James C.R. Stangoulis, and Robin D. Graham Molybdenum, Russell L. Hamlin Nickel, Patrick H. Brown Zinc, J. Benton Storey Beneficial Elements Aluminum, Susan C. Miyasaka, N.V. Hue, and Michael A. Dunn Cobalt, Geeta Talukder and Archana Sharmaz Selenium, Dean A. Kopsell and David E. Kopsell Silicon, George H. Snyder, Vladimir V. Matichenkov and Lawrence E. Datnoff Sodium, John Gorham Vanadium, David J Pilbeam and K Drihem Conclusion, Allen V. Barker and David J Pilbeam

Handbook of plant nutrition

As a major contributor to the reduced nitrogen pool in the biosphere, symbiotic nitrogen fixation by legumes plays a critical role in a sustainable production system. However this legume contribution varies with the physico-chemical and biological conditions of the nodulated-root rhizosphere. In order to assess the abiotic and biotic constrains that might limit this symbiosis at the agroecosystem level, a nodular diagnosis is proposed with common bean as a model grain-legume, and a major source of plant proteins for world human nutrition. The engineering of the legume symbiosis is addressed by participatory assessment of bean recombinant inbred lines contrasting for their efficiency in use of phosphorous for symbiotic nitrogen fixation. With this methodology, in field-sites chosen with farmers of an area of cereal-cropping in the Mediterranean basin, a large spatial and temporal variation in the legume nodulation was found. Soil P availability was a major limiting factor of the rhizobial symbiosis. In order to relate the field measurements with progress in functional genomics of the symbiosis, in situ RT-PCR on nodule sections has been implemented showing that the phytase gene is expressed in the cortex with significantly higher number of transcripts in P-efficient RILs. It is concluded that various tools and indicators are available for developing the ecological engineering of the rhizobial symbiosis, in particular for its beneficial contribution to the bio-geochemical cycle of N, and also P and C.

Nodular diagnosis for ecological engineering of the symbiotic nitrogen fixation with legumes

Rapid development and wide applications of nanotechnology brought about a significant increment on the number of engineered nanomaterials (ENs) inevitably entering our living system. Plants comprise of a very important living component of the terrestrial ecosystem. Studies on the influence of engineered nanomaterials (carbon and metal/metal oxides based) on plant growth indicated that in the excess content, engineered nanomaterials influences seed germination. It assessed the shoot-to-root ratio and the growth of the seedlings. From the toxicological studies to date, certain types of engineered nanomaterials can be toxic once they are not bound to a substrate or if they are freely circulating in living systems. It is assumed that the different types of engineered nanomaterials affect the different routes, behavior, and the capability of the plants. Furthermore, different, or even opposing conclusions, have been drawn from most studies on the interactions between engineered nanomaterials with plants. Therefore, this paper comprehensively reviews the studies on the different types of engineered nanomaterials and their interactions with different plant species, including the phytotoxicity, uptakes, and translocation of engineered nanomaterials by the plant at the whole plant and cellular level.

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Effects of Engineered Nanomaterials on Plants Growth: An Overview

The efficacy of the alumina system for differentiating between bean (Phaseolus vulgaris L.) genotypes for growth at different levels of phosphorus availability was determinated. In addition to response to P levels, comparisons were made between plants receiving N either from fertilizer or nitrogen fixation. When the cv. Carioca was provided with either 100 ppm of N or inoculated withRhizobium leguminosarum biovarphaseoli, differences in shoot dry weight and nodule number were related to P level. There was a greater proportion of green, ineffectivevs. red, active nodules at the low P concentration than at the higher P concentration. In a second experiment, two cvs., Puebla 152 and Carioca and the breeding line UW 24-21, either were inoculated with rhizobia or provided with 150ppm of N. Each genotype-nitrogen combination was grown at 8 levels of P. There was a positive effect of P level on shoot dry weight, nodule number and nodule mass. Root mass was affected less than nodule or shoot mass by the P level of the growth medium. Nodule mass, but not P concentration in the nodules, was affected by P level, whereas in the other plant tissues, P concentrations were lower at lower P levels in the media.

Nitrogen fixation and plant growth of common bean ( Phaseolus vulgaris L.) at different levels of phosphorus availability

Phosphorus is one of several factors which affect N2 fixation and along with N, is a principal yield-limiting nutrient in many regions of the world. Since the legume plant is an essential partner in symbiotic N2 fixation, knowledge of host genotype variability for this process at different levels of P availability will be useful when breeding bean cultivars for enhanced N2 fixation. The objective of this study was to obtain common bean (Phaseolus vulgaris L.) lines able with enhanced ability to support biological N2 fixation under different levels of available phosphorus. Experiments were conducted in a growth room using a sand-alumina system to provide different levels of available P and in the field on a low-N soil. In the growth room studies, P availability strongly affected plant growth and traits related to N2 fixation. No significant interaction was detected for P levels × bean lines, indicating that bean lines performed similarly at both high and low P levels. Total shoot N was used as a direct and indirect measure of N2-fixation potential under growth room (N-free media) and field (low-N soil) conditions, respectively. Based on this criterion, two of the 41 and 54 inbred backcross lines of a segregating population evaluated in the growth room and the field, respectively, contained greater shoot N content than the recurrent parent and N shoot contents similar to the donor parent. Variability of N2 fixation under low available P was observed, and high N2 fixing and high yielding progeny lines were detected.

Selection of common bean (Phaseolus vulgaris L.) for N2 fixation at different levels of available phosphorus under field and environmentally-controlled conditions

Fifteen lettuce cultivars representing three different morphological types were grown in a sand alumina system under conditions of low (deficient) and high (sufficient) P supply. An efficient plant was defined as one that produced a large shoot fresh weight under low P conditions. Cultivars within their respective groups varied significantly for some traits that appeared to be important in determining adaptation to P. This led to the conclusion that accumulation of P in shoot tissue or the total plant was the main difference between efficient and inefficient cultivars. Accumlation of P seemed to be due to a greater absorption capability of roots or greater root mass (weight), depending on the different lettuce groups. Differences in internal use of P did not contribute to differences in shoot fresh weight.

Variability among lettuce cultivars grown at two levels of available phosphorus

The role of lateral root nodules in N2 fixation and the relationships between total shoot N and several traits which influence or control N2 fixation in common bean (Phaseolus vulgaris L.)i.e., acetylene reduction value, specific nodule activity, leghemoglobin concentration, total leghemoglobin and nodule mass, were investigated in field studies. Significant variation among bean lines was observed for all the traits measured. Lines varied for the proportion of total N accumulated up to the R3 growth state, thus measurements of total shoot N near maturity (e.g., R7) provided a better estimate of total N2 fixation than measurements taken at an early growth stage. Nodule mass was correlated with acetylene reduction and total leghemoglobin, and total leghemoglobin was correlated with acetylene reduction value. Total shoot N at R7 was correlated with seasonal means of nodule mass and number, acetylene reduction value and total leghemoglobin. For all traits except total leghemoglobin, values for lateral roots were more highly correlated with total shoot N than were values for either crown roots or the whole root system. Seed yield was most highly correlated with nodule mass of the lateral roots. These results will be useful in devising breeding strategies for improved N2 fixation of the host plant.

Indirect measures of N2 fixation in common bean (Phaseolus vulgaris L.) under field conditions: The role of lateral root nodules

Improvement of dinitrogen fixation in beans (Phaseolus vulgaris L.) will depend on the selection of superior plant genotypes and the presence of efficient rhizobial strains. This study was conducted to evaluate diverse bean lines for N2 fixation potential using the15N-depleted dilution technique under field conditions in Wisconsin, USA. Plants of 21 bean lines and three non-nodulating isolines of soybean received appliin Wisconsin, USA. Plants of 21 bean lines and three non-nodulating isolines of soybean received applications of15N-depleted ammonium sulphate. Shoots harvested at the V6, R3 and R7 stages and dry seeds were analyzed for total N using the Kjeldahl procedure, and the ratio of15N to14N was determind on a MAT 250 mass spectrometer. Nodule occupancy of the applied strain ofR. leguminosarum biovarphaseoli, CIAT 899, was determined in five of the bean lines. Total shoot N content showed a pattern of accumulation similar to shoot dry weight and fixed N2 in the shoot. Based on shoot total N, N2 fixed in the shoot and shoot dry weight Riz 30 and Preto Cariri were identified as being as good fixers as Puebla 152 and Cargamanto appear to begin N2 fixation early. Furthermore, some bean lines that fixed considerable N2 did not translocate a large amount of N to the grains. Preto Cariri accumulated 21.2 kg N ha−1 in the seeds compared to Puebla 152 which accumulated 43.8 kg N ha−1 of the fixed N2 into the grains. At the early sampling, Puebla 152 and 22–27 had a considerable higher percentage of their crown nodules formed by the inoculant strain CIAT 899, than did Rio Tibagi which has been considered a poor N2 fixer.

F. A. Bliss

Papers

Nitrogen fixation and plant growth of common bean ( Phaseolus vulgaris L.) at different levels of phosphorus availability

Selection of common bean (Phaseolus vulgaris L.) for N2 fixation at different levels of available phosphorus under field and environmentally-controlled conditions

Variability among lettuce cultivars grown at two levels of available phosphorus

Indirect measures of N2 fixation in common bean (Phaseolus vulgaris L.) under field conditions: The role of lateral root nodules

15N-determined dinitrogen fixation potential of genetically diverse bean lines (Phaseolus vulgaris L.)