Although epigenetic modifications have been intensely investigated over the last decade due to their role in crop adaptation to rapid climate change, it is unclear which epigenetic changes are heritable and therefore transmitted to their progeny. The identification of epigenetic marks that are transmitted to the next generations is of primary importance for their use in breeding and for the development of new cultivars with a broad-spectrum of tolerance/resistance to abiotic and biotic stresses. In this review, we discuss general aspects of plant responses to environmental stresses and provide an overview of recent findings on the role of transgenerational epigenetic modifications in crops. In addition, we take the opportunity to describe the aims of EPI-CATCH, an international COST action consortium composed by researchers from 28 countries. The aim of this COST action launched in 2020 is: (1) to define standardized pipelines and methods used in the study of epigenetic mechanisms in plants, (2) update, share, and exchange findings in epigenetic responses to environmental stresses in plants, (3) develop new concepts and frontiers in plant epigenetics and epigenomics, (4) enhance dissemination, communication, and transfer of knowledge in plant epigenetics and epigenomics.

Salinity is a major limiting factor in crop productivity worldwide. Medicago sativa L. is an important fodder crop, broadly cultivated in different environments, and it is moderately tolerant of salinity. Medicago arborea L. is considered a stress-tolerant species and could be an important genetic resource for the improvement of M. sativa’s salt tolerance. The aim of the study was to evaluate the seedling response of M. sativa, M. arborea, and their hybrid (Alborea) to salt shock and salt stress treatments. Salt treatments were applied as follows: salt stress treatment at low dose (50 mM NaCl), gradual acclimatization at 50–100 and 50–100–150 mM NaCl, and two salt shock treatments at 100 and 150 mM NaCl. Growth rates were evaluated in addition to transcriptional profiles of representative genes that control salt uptake and transport (NHX1 and RCI2A), have an osmotic function (P5CS1), and participate in signaling pathways and control cell growth and leaf function (SIMKK, ZFN, and AP2/EREB). Results showed that the studied population of M. sativa and M. arborea performed equally well under salt stress, whereas that of M. sativa performed better under salt shock. The productivity of the studied population of Alborea exceeded that of its parents under normal conditions. Nevertheless, Alborea was extremely sensitive to all initial salt treatments except the low dose (50 mM NaCl). In addition, significantly higher expression levels of all the studied genes were observed in the population of M. arborea under both salt shock and salt stress. On the other hand, in the population of M. sativa, NHX1, P5CS1, and AP2/EREB were highly upregulated under salt shock but to a lesser extent under salt stress. Thus, the populations of M. sativa and M. arborea appear to regulate different components of salt tolerance mechanisms. Knowledge of the different parental mechanisms of salt tolerance could be important when incorporating both mechanisms in Alborea populations.

Global climate change is one of the major constraints limiting plant growth, production, and sustainability worldwide. Moreover, breeding efforts of the past years have focused on improving certain favorable crop traits, leading to genetic bottlenecks. The use of crop wild relatives (CWRs) to expand genetic diversity and improve crop adaptability seems to be a promising and sustainable approach for crop improvement in the context of the ongoing climate challenges. In this review, we present the progress that has been achieved towards CWRs exploitation for enhanced resilience against major abiotic stressors (e.g., water deficiency, increased salinity, and extreme temperatures) in crops of high nutritional and economic value, such as tomato, legumes, and several woody perennial crops. The advances in -omics technologies have facilitated the elucidation of the molecular mechanisms that may underlie abiotic stress tolerance. Comparative analyses of whole genome sequencing (WGS) and transcriptomic profiling (RNA-seq) data between crops and their wild relative counterparts have unraveled important information with respect to the molecular basis of tolerance to abiotic stressors. These studies have uncovered genomic regions, specific stress-responsive genes, gene networks, and biochemical pathways associated with resilience to adverse conditions, such as heat, cold, drought, and salinity, and provide useful tools for the development of molecular markers to be used in breeding programs. CWRs constitute a highly valuable resource of genetic diversity, and by exploiting the full potential of this extended allele pool, new traits conferring abiotic-stress tolerance may be introgressed into cultivated varieties leading to superior and resilient genotypes. Future breeding programs may greatly benefit from CWRs utilization for overcoming crop production challenges arising from extreme environmental conditions.

Increased soil salinity, and therefore accumulation of ions, is one of the major abiotic stresses of cultivated plants that negatively affect their growth and yield. Among Medicago species, only Medicago truncatula, which is a model plant, has been extensively studied, while research regarding salinity responses of two important forage legumes of Medicago sativa (M. sativa) and Medicago arborea (M. arborea) has been limited. In the present work, differences between M. arborea, M. sativa and their hybrid Alborea were studied regarding growth parameters and metabolomic responses. The entries were subjected to three different treatments: (1) no NaCl application (control plants), (2) continuous application of 100 mM NaCl (acute stress) and (3) gradual application of NaCl at concentrations of 50-75-150 mM by increasing NaCl concentration every 10 days. According to the results, M. arborea maintained steady growth in all three treatments and appeared to be more resistant to salinity. Furthermore, results clearly demonstrated that M. arborea presented a different metabolic profile from that of M. sativa and their hybrid. In general, it was found that under acute and gradual stress, M. sativa overexpressed saponins in the shoots while M. arborea overexpressed saponins in the roots, which is the part of the plant where most of the saponins are produced and overexpressed. Alborea did not perform well, as more metabolites were downregulated than upregulated when subjected to salinity stress. Finally, saponins and hydroxycinnamic acids were key players of increased salinity tolerance.

The accumulation of ions due to increased salinity in the soil is one of the major abiotic stressors of cultivated plants that negatively affect their productivity. The model plant, Medicago truncatula, is the only Medicago species that has been extensively studied, whereas research into increased salinity adaptation of two important forage legumes, M. sativa and M. arborea, has been limited. In the present study, the expression of six genes, namely SOS1, SOS3, NHX2, AKT, AVP and HKT1 was monitored to investigate the manner in which sodium ions are blocked and transferred to the various plant parts. In addition, in silico miRNA analysis was performed to identify miRNAs that possibly control the expression of the genes studied. The following treatments were applied: (1) salt stress, with initial treatment of 50 mM NaCl and gradual acclimatization every 10 days, (2) salt shock, with continuous application of 100 mM NaCl concentration and (3) no application of NaCl. Results showed that M. arborea appeared to overexpress and activate all available mechanisms of resistance in conditions of increased salinity, while M. sativa acted in a more targeted way, overexpressing the HKT1 and AKT genes that contribute to the accumulation of sodium ions, particularly in the root. Regarding miRNA in silico analysis, five miRNAs with significant complementarity to putative target genes, AKT1, AVP and SOS3 were identified and served as a first step in investigating miRNA regulatory networks. Further miRNA expression studies will validate these results. Our findings contribute to the understanding of the molecular mechanisms underlying salt-responsiveness in Medicago and could be used in the future for generating salt-tolerant genotypes in crop improvement programs.