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.

Merging genotypes: graft union formation and scion-rootstock interactions.

Apple dwarfing rootstocks cause earlier shoot termination and reduced root and shoot mass. To identify physiological factors responsible for rootstock-induced growth restriction, we compared vascular-enriched gene expression between two dwarfing rootstocks (‘M27’ and ‘M9’) and the vigorous rootstock ‘M793’ using RNA sequencing and quantitative reverse transcriptase PCR. Differentially expressed genes common to both dwarfing rootstocks belonged to five main biological processes: (1) primary metabolism, (2) cell wall synthesis and modification, (3) secondary metabolism, (4) hormone signalling and response and (5) redox homeostasis. Genes promoting the biosynthesis of amino acids, lipids and cell walls were downregulated in dwarfing rootstocks, whereas genes promoting the breakdown of these compounds were upregulated. The only exception to this trend was the upregulation of starch synthesis genes in dwarfing rootstocks. Non-structural carbohydrate analysis demonstrated that starch concentrations in ‘M9’ roots, stems and grafted ‘Royal Gala’ (‘RG’) scions were double that of equivalent tissues from ‘RG’ homo-grafted trees (‘RG’/‘RG’). Fructose and glucose concentrations were much lower in all three tissues of the ‘RG’/‘M9’ trees. Together, these data indicate that dwarfing rootstocks are in a state of sugar depletion and reduced cellular activity despite having large starch reserves. Another significant finding was the over-accumulation of flavonoids and the downregulation of auxin influx transporters MdAUX1 and MdLAX2 in dwarfing rootstocks. We propose that both factors reduce polar auxin transport. The results of this study contribute novel information about the physiological state of dwarfing rootstocks. Dwarfing rootstocks used to grow smaller apple trees have a gene expression profile consistent with reduced cellular activity. Toshi Foster and colleagues from the New Zealand Institute for Plant & Food Research compared gene expression levels between two dwarfing rootstocks (the part of the plant onto which buds or shoots are grafted) and one full-size rootstock. They found that, compared to the full-size rootstock, both the dwarfing rootstocks showed reduced expression of genes involved in metabolism, hormone signaling and cell wall synthesis, and an increased expression of genes involved in breaking down amino acids, lipids and other compounds involved in cellular metabolism. Genes involved in starch synthesis were more active though, suggesting that dwarfing rootstocks keep apple trees small because they are in a continuous state of low energy usage despite having large starch reserves.

/pdf/apple-dwarfing-rootstocks-exhibit-an-imbalance-in-jagq2uqawb.pdf

Apple dwarfing rootstocks exhibit an imbalance in carbohydrate allocation and reduced cell growth and metabolism

Branching out: new insights into the genetic regulation of shoot architecture in trees.

Methylation effect on IPT5b gene expression determines cytokinin biosynthesis in apple rootstock.

Grafting has been commonly practiced for many centuries in the cultivation of horticultural crops. The use of dwarfing rootstocks has enabled a high-density plantation to produce maximum yield. Rootstock regulates scion phenotype, including precocity, fruit size, yield, quality characteristics, and tolerance to various environmental stresses. This review summarizes the existing information on the influence of rootstocks on scion growth and dwarfing mechanisms induced by multiple factors, including hormone signaling, photosynthesis, mineral transport, water relations, anatomical characteristics, and genetic markers. It has been shown that the complex interactions between scion and rootstock can regulate plant development and its structure. This information will provide interesting insights for future research related to rootstock-mediated dwarfing mechanisms and accelerate the breeding progress of dwarfing rootstocks.

An insight into dwarfing mechanism: contribution of scion-rootstock interactions toward fruit crop improvement

Rootstock-induced dwarfing of apple scions revolutionized global apple production during the twentieth century, leading to the development of modern intensive orchards. A high root bark percentage (the percentage of the whole root area constituted by root cortex) has previously been associated with rootstock induced dwarfing in apple. In this study, the root bark percentage was measured in a full-sib family of ungrafted apple rootstocks and found to be under the control of three loci. Two QTL for root bark percentage were found to co-localise to the same genomic regions on chromosome 5 and chromosome 11 previously identified as controlling dwarfing, Dw1 and Dw2, respectively. A third QTL was identified on chromosome 13 in a region that has not been previously associated with dwarfing. The development of closely linked 3 Sequence-tagged site STS markers improved the resolution of allelic classes thereby allowing the detection of dominance and epistatic interactions between loci, with high root bark percentage only occurring in specific allelic combinations. In addition, we report a significant negative correlation between root bark percentage and stem diameter (an indicator of tree vigour), measured on a clonally propagated grafted subset of the mapping population. The demonstrated link between root bark percentage and rootstock-induced dwarfing of the scion leads us to propose a three-locus model that is able to explain levels of dwarfing from the dwarf ‘M.27’ to the semi-invigorating rootstock ‘M.116’. Moreover, we suggest that the QTL on chromosome 13 (Rb3) might be analogous to a third dwarfing QTL, Dw3 that has not previously been identified.

/pdf/a-new-three-locus-model-for-rootstock-induced-dwarfing-in-5e49y4jlxf.pdf

A new three-locus model for rootstock-induced dwarfing in apple revealed by genetic mapping of root bark percentage.

Fruits are an important source of vitamins, minerals and nutrients in the human diet. They also contain several compounds of nutraceutical importance that have significant antioxidant and anti-inflammatory roles, which can protect the consumer from diseases, such as cancer, and cardiovascular disease as well as having roles in reducing the build-up of LDL-cholesterol in blood plasma and generally reduce the risks of disease and age-related decline in health. Cherries contain high concentrations of bioactive compounds and minerals, including calcium, phosphorous, potassium and magnesium, and it is, therefore, unsurprising that cherry consumption has a positive impact on health. This review highlights the development of sweet cherry fruit, the health benefits of cherry consumption, and the options for increasing consumer acceptance and consumption.

/pdf/fruit-development-in-sweet-cherry-3vjrpj2v.pdf

Fruit Development in Sweet Cherry

Abstract Main conclusion This manuscript identifies cherry orthologues of genes implicated in the development of pericarpic fruit and pinpoints potential options and restrictions in the use of these targets for commercial exploitation of parthenocarpic cherry fruit. Abstract Cherry fruit contain a large stone and seed, making processing of the fruit laborious and consumption by the consumer challenging, inconvenient to eat ‘on the move’ and potentially dangerous for children. Availability of fruit lacking the stone and seed would be potentially transformative for the cherry industry, since such fruit would be easier to process and would increase consumer demand because of the potential reduction in costs. This review will explore the background of seedless fruit, in the context of the ambition to produce the first seedless cherry, carry out an in-depth analysis of the current literature around parthenocarpy in fruit, and discuss the available technology and potential for producing seedless cherry fruit as an ‘ultimate snacking product’ for the twenty-first century. 

/pdf/options-for-the-generation-of-seedless-cherry-the-ultimate-rgomai5p.pdf

Options for the generation of seedless cherry, the ultimate snacking product

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A new three-locus model for rootstock-induced dwarfing in apple revealed by genetic mapping of root bark percentage.

Fruit Development in Sweet Cherry

Options for the generation of seedless cherry, the ultimate snacking product