Legumes are renowned for their distinctive biological characteristic of forming symbiotic associations with soil bacteria, mostly belonging to the Rhizobiaceae familiy, leading to the establishment of symbiotic root nodules. Within these nodules, rhizobia play a pivotal role in converting atmospheric nitrogen into a plant-assimilable form. However, it has been discerned that root nodules of legumes are not exclusively inhabited by rhizobia; non-rhizobial endophytic bacteria also reside within them, yet their functions remain incompletely elucidated. This comprehensive review synthesizes available data, revealing that Bacillus and Pseudomonas are the most prevalent genera of nodule endophytic bacteria, succeeded by Paenibacillus, Enterobacter, Pantoea, Agrobacterium, and Microbacterium. To date, the bibliographic data available show that Glycine max followed by Vigna radiata, Phaseolus vulgaris and Lens culinaris are the main hosts for nodule endophytic bacteria. Clustering analysis consistently supports the prevalence of Bacillus and Pseudomonas as the most abundant nodule endophytic bacteria, alongside Paenibacillus, Agrobacterium, and Enterobacter. Although non-rhizobial populations within nodules do not induce nodule formation, their presence is associated with various plant growth-promoting properties (PGPs). These properties are known to mediate important mechanisms such as phytostimulation, biofertilization, biocontrol, and stress tolerance, emphasizing the multifaceted roles of nodule endophytes. Importantly, interactions between non-rhizobia and rhizobia within nodules may exert influence on their leguminous host plants. This is particularly shown by co-inoculation of legumes with both types of bacteria, in which synergistic effects on plant growth, yield, and nodulation are often measured. Moreover these effects are pronounced under both stress and non-stress conditions, surpassing the impact of single inoculations with rhizobia alone.

The potential of rhizobacteria in assisting plants used in the phytostabilization or re-vegetation of soils contaminated by heavy metals is gaining interest all around the world. In this context, six rhizobacterial strains isolated from highly heavy metal-contaminated soils situated in abandoned mining sites around the Oujda region (Morocco) were tested with Sulla spinosissima (L.), a native leguminous plant expanding in this area. The strains used were multi-resistant to heavy metals and possessed multiple plant growth-promoting traits. Potential beneficial effects of the strains were also evaluated in planta by measuring various growth and physiological parameters of inoculated Sulla plants grown in sterilized sand. Inoculation with the Rhodococcus qingshengii strain LMR340 boosted plant biomass (39% to 83% increase compared to uninoculated plants), chlorophyll and carotenoid content (up to 29%), and antioxidant enzyme activities (15% to 80% increase). Based on these interesting findings, selected strains were inoculated into plants growing in a heavy metal, multi-polluted, and poor soil. Under these conditions, non-inoculated plants and those inoculated with the strain LMR250 were unable to grow, while the other five bacterial inoculants restored plant growth. The best performing strain, Pseudarthrobacter oxydans LMR291, could be considered as a good biofertilizer and/or biostimulant candidate to be used for promoting the growth of selected plants in re-vegetation and/or phytostabilization programs of degraded and contaminated soils.

The phenotypic characteristics of 48 isolates obtained from root nodules of four Acacia species (Acacia cyanophylla, A. gummifera, A. horrida and A. raddiana) growing in soils collected from the arid...