Breeding Approaches for Enhancing Heavy Metal Tolerance in Legume: A Comprehensive Review
摘要
Breeding approaches for improving heavy metal stress tolerance in legume crops have become increasingly important in response to the rising contamination of agricultural soils with toxic metals such as cadmium, lead, arsenic, nickel, and aluminum. Conventional breeding has laid the foundation by exploiting natural genetic variation for traits related to metal exclusion, sequestration, and tolerance; however, its effectiveness is often limited by the complex, quantitative inheritance and strong environmental influence of heavy metal stress responses. To overcome these constraints, modern breeding strategies now integrate marker-assisted selection, genomic selection, QTL mapping, and candidate gene-based approaches to enhance selection precision and accelerate genetic gains. In parallel, transgenic and genome-editing approaches enable functional validation and targeted manipulation of key genes involved in metal transport, chelation, antioxidant defense, and stress signaling pathways. Moreover, microRNAs (miRNAs) have emerged as critical post-transcriptional regulators that modulate the expression of multiple stress-responsive genes under heavy metal stress. Plastid transformation is a powerful transgenic approach for homologous recombination, site-specific integration of foreign genes into the plastid genome and stable expression of traits related to metal tolerance. Phytoremediation strategies using transgenic legumes further present eco-friendly and cost-effective solutions for the extraction, sequestration, or stabilization of heavy metals from contaminated soils. The combined application of these breeding approaches allows the development of legume cultivars with improved productivity, reduced metal accumulation in edible parts, and enhanced adaptability to contaminated soils. This review focusing on integrated breeding strategies to provide a robust and forward-looking framework for sustaining legume production, food quality, and supporting environmental remediation under heavy metal stress conditions.