Root exudates drive interspecific facilitation by enhancing nodulation and N2 fixation.
ABSTRACT: Plant diversity in experimental systems often enhances ecosystem productivity, but the mechanisms causing this overyielding are only partly understood. Intercropping faba beans (Vicia faba L.) and maize (Zea mays L.) result in overyielding and also, enhanced nodulation by faba beans. By using permeable and impermeable root barriers in a 2-y field experiment, we show that root-root interactions between faba bean and maize significantly increase both nodulation and symbiotic N2 fixation in intercropped faba bean. Furthermore, root exudates from maize promote faba bean nodulation, whereas root exudates from wheat and barley do not. Thus, a decline of soil nitrate concentrations caused by intercropped cereals is not the sole mechanism for maize promoting faba bean nodulation. Intercropped maize also caused a twofold increase in exudation of flavonoids (signaling compounds for rhizobia) in the systems. Roots of faba bean treated with maize root exudates exhibited an immediate 11-fold increase in the expression of chalcone-flavanone isomerase (involved in flavonoid synthesis) gene together with a significantly increased expression of genes mediating nodulation and auxin response. After 35 d, faba beans treated with maize root exudate continued to show up-regulation of key nodulation genes, such as early nodulin 93 (ENOD93), and promoted nitrogen fixation. Our results reveal a mechanism for how intercropped maize promotes nitrogen fixation of faba bean, where maize root exudates promote flavonoid synthesis in faba bean, increase nodulation, and stimulate nitrogen fixation after enhanced gene expression. These results indicate facilitative root-root interactions and provide a mechanism for a positive relationship between species diversity and ecosystem productivity.
Project description:Intercropping, which grows at least two crop species on the same pieces of land at the same time, can increase grain yields greatly. Legume-grass intercrops are known to overyield because of legume nitrogen fixation. However, many agricultural soils are deficient in phosphorus. Here we show that a new mechanism of overyielding, in which phosphorus mobilized by one crop species increases the growth of a second crop species grown in alternate rows, led to large yield increases on phosphorus-deficient soils. In 4 years of field experiments, maize (Zea mays L.) overyielded by 43% and faba bean (Vicia faba L.) overyielded by 26% when intercropped on a low-phosphorus but high-nitrogen soil. We found that overyielding of maize was attributable to below-ground interactions between faba bean and maize in another field experiment. Intercropping with faba bean improved maize grain yield significantly and above-ground biomass marginally significantly, compared with maize grown with wheat, at lower rates of P fertilizer application (<75 kg of P(2)O(5) per hectare), and not significantly at high rate of P application (>112.5 kg of P(2)O(5) per hectare). By using permeable and impermeable root barriers, we found that maize overyielding resulted from its uptake of phosphorus mobilized by the acidification of the rhizosphere via faba bean root release of organic acids and protons. Faba bean overyielded because its growth season and rooting depth differed from maize. The large increase in yields from intercropping on low-phosphorus soils is likely to be especially important on heavily weathered soils.
Project description:Wheat (Triticum aestivum L.)/faba bean (Vicia faba L.) intercropping shows significant overyielding and high nitrogen (N)-use efficiency, but the dynamics of plant interactions have rarely been estimated. The objective of the present study was to investigate the temporal dynamics of competitive N acquisition between intercropped wheat and faba bean with the logistic model. Wheat and faba bean were grown together or alone with limited N supply in pots. Data of shoot and root biomass and N content measured from 14 samplings were fitted to logistic models to determine instantaneous rates of growth and N uptake. The superiority of instantaneous biomass production and N uptake shifted from faba bean to wheat with their growth. Moreover, the shift of superiority on N uptake occurred 7-12 days earlier than that of biomass production. Interspecific competition stimulated intercropped wheat to have a much earlier and stronger superiority on instantaneous N uptake compared with isolated wheat. The modeling methodology characterized the temporal dynamics of biomass production and N uptake of intercropped wheat and faba bean in different planting systems, which helps to understand the underlying process of plant interaction for intercropping plants.
Project description:Rhizosphere processes stimulate overyielding and facilitative phosphorus (P) uptake in cereal/legume intercropping systems. However, little is known about when and how rhizosphere alteration of legumes plays a role in improving P uptake by cereals. Wheat was grown isolated, monocropped or intercropped with faba bean in pots with low-P soil. The biomass, P content, carboxylates and phosphatases activity were measured in 15 destructive samplings. Intraspecific competition of the biomass and P uptake of monocropped wheat was not significant before 40 and 36 days after sowing (DAS), whereas there was interspecific competition of biomass of intercropped wheat before 66 DAS. However, afterwards, the increments of the biomass and P uptake of the intercropped wheat were 1.3-1.9 and 1.9-2.3 times of increment of monocropped wheat. Meanwhile, the concentrations of malate and citrate and the acid phosphatase activity in the rhizospheres of intercropped wheat were significantly increased, which suggested that wheat/faba bean intercropping is efficient in P utilization due to complementary P uptake in the early growth stage and the positive interactions of the rhizosphere processes when the soil P was depleted.
Project description:Populations of Rhizobium leguminosarum biovar viciae were sampled from two bulk soils, rhizosphere, and nodules of host legumes, fava bean (Vicia faba) and pea (Pisum sativum) grown in the same soils. Additional populations nodulating peas, fava beans, and vetches (Vicia sativa) grown in other soils and fava bean-nodulating strains from various geographic sites were also analyzed. The rhizobia were characterized by repetitive extragenomic palindromic-PCR fingerprinting and/or PCR-restriction fragment length polymorphism (RFLP) of 16S-23S ribosomal DNA intergenic spacers as markers of the genomic background and PCR-RFLP of a nodulation gene region, nodD, as a marker of the symbiotic component of the genome. Pairwise comparisons showed differences among the genetic structures of the bulk soil, rhizosphere, and nodule populations and in the degree of host specificity within the Vicieae cross-inoculation group. With fava bean, the symbiotic genotype appeared to be the preponderant determinant of the success in nodule occupancy of rhizobial genotypes independently of the associated genomic background, the plant genotype, and the soil sampled. The interaction between one particular rhizobial symbiotic genotype and fava bean seems to be highly specific for nodulation and linked to the efficiency of nitrogen fixation. By contrast with bulk soil and fava bean-nodulating populations, the analysis of pea-nodulating populations showed preferential associations between genomic backgrounds and symbiotic genotypes. Both components of the rhizobial genome may influence competitiveness for nodulation of pea, and rhizosphere colonization may be a decisive step in competition for nodule occupancy.
Project description:Root morphological/physiological modifications are important for phosphorus (P) acquisition of plants under P deficiency, but strategies differ among plant species. Detailed studies on the response of maize roots to P deficiency are limited. Nitrogen (N) form influences root morphology/physiology, and thus may influence root responses to P deficiency. This work investigated adaptive mechanisms of maize roots to low P by comparison with white lupin and faba bean supplied with two N forms. Plants were grown for 7-16 days in hydroponics with sufficient (250 µmol L(-1)) and deficient P supply (1 µmol L(-1)) under supply of NH4NO3 or Ca(NO3)2 Plant growth and P uptake were measured, and release of protons and organic acid anions, and acid phosphatase activity in the root were monitored. The results showed that P deficiency significantly decreased shoot growth while increased root growth and total root length of maize and faba bean, but not white lupin. It enhanced the release of protons and organic acid anions, and acid phosphatase activity, from the roots of both legumes but not maize. Compared with Ca(NO3)2, NH4NO3 dramatically increased proton release by roots but did not alter root morphology or physiology of the three species in response to low P. It is concluded that the N form did not fundamentally change root morphological/physiological responses of the three species to P deficiency. Morphological variation in maize and morpho-physiological modifications in white lupin and faba bean were the main adaptive strategies to P deficiency.
Project description:BACKGROUND: Within-field multiple crop species intercropping is well documented and used for disease control, but the underlying mechanisms are still unclear. As roots are the primary organ for perceiving signals in the soil from neighboring plants, root behavior may play an important role in soil-borne disease control. PRINCIPAL FINDINGS: In two years of field experiments, maize/soybean intercropping suppressed the occurrence of soybean red crown rot, a severe soil-borne disease caused by Cylindrocladium parasiticum (C. parasiticum). The suppressive effects decreased with increasing distance between intercropped plants under both low P and high P supply, suggesting that root interactions play a significant role independent of nutrient status. Further detailed quantitative studies revealed that the diversity and intensity of root interactions altered the expression of important soybean PR genes, as well as, the activity of corresponding enzymes in both P treatments. Furthermore, 5 phenolic acids were detected in root exudates of maize/soybean intercropped plants. Among these phenolic acids, cinnamic acid was released in significantly greater concentrations when intercropped maize with soybean compared to either crop grown in monoculture, and this spike in cinnamic acid was found dramatically constrain C. parasiticum growth in vitro. CONCLUSIONS: To the best of our knowledge, this study is the first report to demonstrate that intercropping with maize can promote resistance in soybean to red crown rot in a root-dependent manner. This supports the point that intercropping may be an efficient ecological strategy to control soil-borne plant disease and should be incorporated in sustainable agricultural management practices.
Project description:BACKGROUND AND AIMS:During their domestication, maize, bean and squash evolved in polycultures grown by small-scale farmers in the Americas. Polycultures often overyield on low-fertility soils, which are a primary production constraint in low-input agriculture. We hypothesized that root architectural differences among these crops causes niche complementarity and thereby greater nutrient acquisition than corresponding monocultures. METHODS:A functional-structural plant model, SimRoot, was used to simulate the first 40 d of growth of these crops in monoculture and polyculture and to determine the effects of root competition on nutrient uptake and biomass production of each plant on low-nitrogen, -phosphorus and -potassium soils. KEY RESULTS:Squash, the earliest domesticated crop, was most sensitive to low soil fertility, while bean, the most recently domesticated crop, was least sensitive to low soil fertility. Nitrate uptake and biomass production were up to 7 % greater in the polycultures than in the monocultures, but only when root architecture was taken into account. Enhanced nitrogen capture in polycultures was independent of nitrogen fixation by bean. Root competition had negligible effects on phosphorus or potassium uptake or biomass production. CONCLUSIONS:We conclude that spatial niche differentiation caused by differences in root architecture allows polycultures to overyield when plants are competing for mobile soil resources. However, direct competition for immobile resources might be negligible in agricultural systems. Interspecies root spacing may also be too large to allow maize to benefit from root exudates of bean or squash. Above-ground competition for light, however, may have strong feedbacks on root foraging for immobile nutrients, which may increase cereal growth more than it will decrease the growth of the other crops. We note that the order of domestication of crops correlates with increasing nutrient efficiency, rather than production potential.
Project description:Even though the faba bean (Vicia faba L.) is among the most ubiquitously cultivated crops, very little is known about its origins. Here, we report discoveries of charred faba beans from three adjacent Neolithic sites in the lower Galilee region, in the southern Levant, that offer new insights into the early history of this species. Biometric measurements, radiocarbon dating and stable carbon isotope analyses of the archaeological remains, supported by experiments on modern material, date the earliest farming of this crop to ~10,200?cal?BP. The large quantity of faba beans found in these adjacent sites indicates intensive production of faba beans in the region that can only have been achieved by planting non-dormant seeds. Selection of mutant-non-dormant stock suggests that the domestication of the crop occurred as early as the 11(th) millennium cal BP. Plant domestication| Vicia faba L.| Pre-Pottery Neolithic B| radiocarbon dating| ?(13)C analysis.
Project description:Cereal/legume intercropping is likely to achieve the optimal exploitation of soil and atmospheric nitrogen (N) sources to maintain high production and quality levels with low N inputs, as an attempt to eliminate underlying environmental effects. Nevertheless, the extent of the effect of cereal/legume intercropping on nodulation and N fixation of intercropped legumes in root interaction requires extensive verification. In the present study, root interaction of alfalfa/triticale intercropping was simulated in pots with the use of root separation types (pot with no barrier (A-T), pot with nylon mesh barrier (NA-T), pot with plastic barrier (PA-T), and alfalfa alone (SA)) in pots. Moreover, the experiment was measured at a range of N levels (N21, N210) and growing stages (branching, budding and initial flowering stages) in growth chamber. As alfalfa was growing, the total nodule number (TNN), effective nodule number (ENN) and nitrogenase activity (NA) of alfalfa with A-T and other cropping systems more noticeably differed from each other at higher N levels, whereas their diversification was reduced at lower N levels. As alfalfa was growing continuously, fresh nodule weight per plant (PNW) and single fresh nodule weight (SNW) with A-T and other cropping systems were amplified more significantly. The nodulation and N fixation ability under N21 were more significant than those under N210. Dry weight of plant per pot (TDW) and nitrogen accumulation of plant per pot (TNA) with A-T were obviously higher than those with other systems in the initial flowering stage, except for TNA under N21. The parameters regarding the nodulation and N fixation ability were significantly positively correlated on the whole. However, SNW and TNA were not significantly correlated, neither were SNW and TDW. According to the mentioned results, the closer root interaction, the better the nodulation form and N fixation ability of alfalfa will be, and the higher the biomass and N accumulation of all plants in pots will be. Interspecific facilitation in alfalfa/triticale intercropping system resulted in a greater yield and N accumulation; it also ultimately enhanced nodulation and N fixation ability, which can be applied in sustainable systems to avoid N loss to the environment and enhance N use efficiency.
Project description:The understanding of crop domestication is dependent on tracking the original geographical distribution of wild relatives. The faba bean (Vicia faba L.) is economically important in many countries around the world; nevertheless, its origin has been debated because its ancestor could not be securely identified. Recent investigations in the site of el-Wad (Mount Carmel, Israel), provide the first and, so far, only remains of the lost ancestor of faba bean. X-ray CT scan analysis of the faba beans provides the first set of measurements of the biometry of this species before its domestication. The presence of wild specimens in Mount Carmel, 14,000 years ago, supports that the wild variety grew nearby in the Lower Galilee where the first domestication was documented for Neolithic farmers 10,200 years ago.