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Grafting has been widely used to improve horticultural traits. It has also served increasingly as a tool to investigate the long-distance transport of molecules that is an essential part for key biological processes. Many studies have revealed the molecular mechanisms of graft-induced phenotypic variation in anatomy, morphology and production. Here, we review the phenomena and their underlying mechanisms by which macromolecules, including RNA, protein, and even DNA, are transported between scions and rootstocks via vascular tissues. We further propose a conceptual framework that characterizes and quantifies the driving mechanisms of scion–rootstock interactions toward vascular reconnection and regeneration.

Grafting is typically utilized to merge adapted seedling rootstocks with highly productive clonal scions. This process implies the interaction of multiple genomes to produce a unique tree phenotype. However, the interconnection of both genotypes obscures individual contributions to phenotypic variation (rootstock-mediated heritability), hampering tree breeding. Therefore, our goal was to quantify the inheritance of seedling rootstock effects on scion traits using avocado ( Persea americana Mill.) cv. Hass as a model fruit tree. We characterized 240 diverse rootstocks from 8 avocado cv. Hass orchards with similar management in three regions of the province of Antioquia, northwest Andes of Colombia, using 13 microsatellite markers simple sequence repeats (SSRs). Parallel to this, we recorded 20 phenotypic traits (including morphological, biomass/reproductive, and fruit yield and quality traits) in the scions for 3 years (2015–2017). Relatedness among rootstocks was inferred through the genetic markers and inputted in a “genetic prediction” model to calculate narrow-sense heritabilities ( h 2 ) on scion traits. We used three different randomization tests to highlight traits with consistently significant heritability estimates. This strategy allowed us to capture five traits with significant heritability values that ranged from 0.33 to 0.45 and model fits ( r ) that oscillated between 0.58 and 0.73 across orchards. The results showed significance in the rootstock effects for four complex harvest and quality traits (i.e., total number of fruits, number of fruits with exportation quality, and number of fruits discarded because of low weight or thrips damage), whereas the only morphological trait that had a significant heritability value was overall trunk height (an emergent property of the rootstock–scion interaction). These findings suggest the inheritance of rootstock effects, beyond root phenotype, on a surprisingly wide spectrum of scion traits in “Hass” avocado. They also reinforce the utility of polymorphic SSRs for relatedness reconstruction and genetic prediction of complex traits. This research is, up to date, the most cohesive evidence of narrow-sense inheritance of rootstock effects in a tropical fruit tree crop. Ultimately, our work highlights the importance of considering the rootstock–scion interaction to broaden the genetic basis of fruit tree breeding programs while enhancing our understanding of the consequences of grafting.

The ancient plant production practice of grafting which instantly imparts new physiological properties to the desirable scion still remains shrouded in mystery. Yet, grafting remains a widely used technique in the production of several horticultural species. In a composite grafted plant, rootstocks control many aspects of scion growth and physiology including yield and quality attributes as well as biotic and abiotic stress tolerance. Broadly, physical, physiological, biochemical and molecular mechanisms have been reviewed to develop an integrated understanding of this enigmatic process that challenges existing genetic paradigms. This review summarizes the reported mechanisms underlying some of the economically important traits and identifies several key points to consider when conducting rootstock scion interaction experiments. Study of the somatogenetic interactions between rootstock and scion is a field that is ripe for discovery and vast improvements in the coming decade. Further, utilization of rootstocks based on a better understanding of the somatogenetic interactions is highly relevant in the current agricultural environment where there is a need for sustainable production practices. Rootstocks may offer a non-transgenic approach to rapidly respond to the changing environment and expand agricultural production of annual and perennial crops where grafting is feasible in order to meet the global food, fiber and fuel demands of the future.

Grafting is a centuries-old technique used in plants to obtain economic benefits. Grafting increases nutrient uptake and utilization efficiency in a number of plant species, including fruits, vegetables, and ornamentals. Selected rootstocks of the same species or close relatives are utilized in grafting. Rootstocks absorb more water and ions than self-rooted plants and transport these water and ions to the aboveground scion. Ion uptake is regulated by a complex communication mechanism between the scion and rootstock. Sugars, hormones, and miRNAs function as long-distance signaling molecules and regulate ion uptake and ion homeostasis by affecting the activity of ion transporters. This review summarizes available information on the effect of rootstock on nutrient uptake and utilization and the mechanisms involved. Information on specific nutrient-efficient rootstocks for different crops of commercial importance is also provided. Several other important approaches, such as interstocking (during double grafting), inarching, use of plant-growth-promoting rhizobacteria, use of arbuscular mycorrhizal fungi, use of plant growth substances (e.g., auxin and melatonin), and use of genetically engineered rootstocks and scions (transgrafting), are highlighted; these approaches can be combined with grafting to enhance nutrient uptake and utilization in commercially important plant species. Whether the rootstock and scion affect each other's soil microbiota and their effect on the nutrient absorption of rootstocks remain largely unknown. Similarly, the physiological and molecular bases of grafting, crease formation, and incompatibility are not fully identified and require investigation. Grafting in horticultural crops can help reveal the basic biology of grafting, the reasons for incompatibility, sensing, and signaling of nutrients, ion uptake and transport, and the mechanism of heavy metal accumulation and restriction in rootstocks. Ion transporter and miRNA-regulated nutrient studies have focused on model and non-grafted plants, and information on grafted plants is limited. Such information will improve the development of nutrient-efficient rootstocks.

Generation of stable gene-edited plant lines using clustered regularly interspaced short palindromic repeats (CRISPR)–CRISPR-associated protein 9 (Cas9) requires a lengthy process of outcrossing to eliminate CRISPR–Cas9-associated sequences and produce transgene-free lines. We have addressed this issue by designing fusions of Cas9 and guide RNA transcripts to tRNA-like sequence motifs that move RNAs from transgenic rootstocks to grafted wild-type shoots (scions) and achieve heritable gene editing, as demonstrated in wild-type Arabidopsis thaliana and Brassica rapa . The graft-mobile gene editing system enables the production of transgene-free offspring in one generation without the need for transgene elimination, culture recovery and selection, or use of viral editing vectors. We anticipate that using graft-mobile editing systems for transgene-free plant production may be applied to a wide range of breeding programs and crop plants. Gene-edited plants free of CRISPR-associated sequences are generated by grafting.

The appropriate selection of rootstock-scion combinations to improve yield and fully realize grafting benefits requires an in-depth understanding of rootstock-scion synergy. Toward this end, we grafted two determinate-type scions [grape tomato (‘BHN 1022') and beefsteak tomato (‘Skyway')] onto four rootstocks with different characteristics to examine plant growth, yield performance, biomass production, and fruit mineral nutrient composition. The study was conducted during two growing seasons (spring and fall plantings in Florida) under organic production in high tunnels with the non-grafted scions as controls. Rootstocks had previously been designated as either “generative” (‘Estamino') or “vegetative” (‘DR0141TX') by some commercial suppliers or had not been characterized [‘RST-04-106-T' and ‘SHIELD RZ F1 (61-802)']. Also, ‘Estamino', ‘DR0141TX', and ‘RST-04-106-T' had been described as more vigorous than ‘SHIELD RZ F1 (61-802)'. In both planting seasons (with low levels of soilborne disease pressure), the “vegetative” and “generative” rootstocks increased marketable and total fruit yields for both scions except for the beefsteak tomato grafted with the “vegetative” rootstock in fall planting. Positive effects of ‘RST-04-106-T' on fruit yield varied with scions and planting seasons, and were most manifested when grafted with the beefsteak tomato scion in fall planting. ‘SHIELD RZ F1 (61-802)' led to similar yields as the non-grafted controls except for grafting with the grape tomato scion in fall planting. For vegetative and fruit biomass, both the “vegetative” and “generative” rootstocks had positive impacts except for the beefsteak tomato in fall planting. For fruit mineral composition, the “vegetative” and “generative” rootstocks, both highly vigorous, consistently elevated fruit P, K, Ca, Zn, and Fe contents on a dry weight basis, whereas the other rootstocks did not. Overall, although the more vigorous rootstocks enhanced tomato plant productivity and fruit minerals, the evidence presented here does not support the suggestion that the so-called “vegetative” and “generative” rootstocks have different impacts on tomato scion yield, biomass production, or fruit mineral contents. More studies with different production systems and environmental conditions as well as contrasting scion genotypes are needed to further categorize the impacts of rootstocks with different vigor and other characteristics on plant biomass production and their implications on fruit yield development.

Key messageRootstock provides more abscisic acid (ABA) content to scions to increase the chilling tolerance of seedlings. H2O2 is involved in ABA regulation of grafting-induced chilling tolerance of cucumber.Here we examined the role of ABA in the response of grafted cucumber to chilling stress. The data showed chilling induced an increase in leaf and root ABA content and there was a positive correlation between ABA content and the chilling tolerance of the varieties. The increase of ABA content and NCED mRNA abundance in the leaf of both Cs/Cs (self-root) and Cs/Cm (grafted with pumpkin as rootstock) showed a delay under aerial stress compared with those under whole plant and root-zone stress. Intriguingly, an increase in ABA in xylem was found under whole-plant and root-zone chilling stress but was not detected under aerial stress, implying the increases in ABA content in leaves were mainly from root ABA transportation. Compared to Cs/Cs, a higher ABA content and NCED mRNA abundance were observed in Cs/Cm, which showed that Cm could output more ABA than Cs. The removal of endogenous ABA decreased the difference in chilling tolerance induced by Cm, as evidenced by the observed similar oxidative stress levels and photosynthetic capacity between Cs/Cs and Cs/Cm after chilling stress. Moreover, we found that the H2O2 signal in grafted cucumber could respond to chilling stress earlier than the H2O2 signal in self-rooted cucumber. The inhibition of endogenous H2O2 decreased the chilling tolerance of grafted cucumber induced by ABA by reducing photosynthesis and the mRNA abundance of CBF1 and COR. Thus, our results indicate that H2O2, as the downstream signal, participated in the rootstock-induced chilling tolerance of grafted seedlings induced by ABA.

Vegetable grafting is extensively used today in agricultural production to control soil-borne pathogens, abiotic and biotic stresses and to improve phenotypic characteristics of the scion. Commercial vegetable grafting is currently practiced in tomato, watermelon, melon, eggplant, cucumber, and pepper. It is also regarded as a rapid alternative to the relatively slow approach of breeding for increased environmental-stress tolerance of fruit vegetables. However, even though grafting has been used for centuries, until today, there are still many issues that have not been elucidated. This review will emphasize on the important mechanisms taking place during grafting, especially the genomic interactions between grafting partners and the impact of rootstocks in scion’s performance. Special emphasis will be drawn on the relation between vegetable grafting, epigenetics, and the changes in morphology and quality of the products. Recent advances in plant science such as next-generation sequencing provide new information regarding the molecular interactions between rootstock and scion. It is now evidenced that genetic exchange is happening across grafting junctions between rootstock and scion, potentially affecting grafting-mediated effects already recorded in grafted plants. Furthermore, significant changes in DNA methylation are recorded in grafted scions, suggesting that these epigenetic mechanisms could be implicated in grafting effects. In this aspect, we also discuss the process and the molecular aspects of rootstock scion communication. Finally, we provide with an extensive overview of gene expression changes recorded in grafted plants and how these are related to the phenotypic changes observed. Τhis review finally seeks to elucidate the dynamics of rootstock-scion interactions and thus stimulate more research on grafting in the future. In a future where sustainable agricultural production is the way forward, grafting could play an important role to develop products of higher yield and quality in a safe and “green” way.

Grafting has been utilised for at least the past 7000 years. Historically, grafting has been developed by growers without particular interest beyond the agronomical and ornamental effects, and thus knowledge about grafting has remained largely empirical. Much of the commercial production of fruit, and increasingly vegetables, relies upon grafting with rootstocks to provide resistance to soil-borne pathogens and abiotic stresses as well as to influence scion growth and performance. Although there is considerable agronomic knowledge about the use and selection of rootstocks for many species, we know little of the molecular mechanisms underlying rootstock adaptation to different soil environments and rootstock-conferred modifications of scion phenotypes. Furthermore, the processes involved in the formation of the graft union and graft compatibility are poorly understood despite over a hundred years of scientific study. In this paper, we provide an overview of what is known about grafting and the mechanisms underlying rootstock-scion interactions. We highlight recent studies that have advanced our understanding of graft union formation and outline subjects that require further development.

Drought, one of the most severe and complex abiotic stresses, is increasingly occurring due to global climate change and adversely affects plant growth and yield. Grafting is a proven and effective tool to enhance plant drought resistance ability by regulating their physiological and molecular processes. In this review, we have summarized the current understanding, mechanisms, and perspectives of the drought stress resistance of grafted plants. Plants resist drought through adaptive changes in their root, stem, and leaf morphology and structure, stomatal closure modulation to reduce transpiration, activating osmoregulation, enhancing antioxidant systems, and regulating phytohormones and gene expression changes. Additionally, the mRNAs, miRNAs and peptides crossing the grafted healing sites also confer drought resistance. However, the interaction between phytohormones, establishment of the scion-rootstock communication through genetic materials to enhance drought resistance is becoming a hot research topic. Therefore, our review provides not only physiological evidences for selecting drought-resistant rootstocks or scions, but also a clear understanding of the potential molecular effects to enhance drought resistance using grafted plants.