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Research article · 2023 · Full text

International Journal of Horticulture and Food Science

Rootstock breeding in fruit crops: Advances in selection, compatibility and stress tolerance mechanisms

Satheesh B

Volume 5(2) · Pages 123–127 · DOI: 10.33545/26631067.2023.v5.i2b.465 ↗

Research areasPlant Disease Management TechniquesPlant Physiology and Cultivation StudiesPhytoplasmas and Hemiptera pathogens

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Abstract

Author-written

Rootstock selection fundamentally influences fruit crop performance through effects on tree vigor, stress tolerance, nutrient acquisition, and scion productivity. This review examines recent advances in rootstock breeding approaches, graft compatibility mechanisms, and stress tolerance characteristics across major fruit crop species. Traditional selection from wild populations and chance seedlings has given way to systematic breeding programs employing hybridization, mutation breeding, and marker-assisted selection to develop improved rootstocks meeting specific production objectives. Analysis of breeding progress reveals that dwarfing characteristics have received greatest attention, with successful development of size-controlling rootstocks enabling high-density planting systems in apple, pear, and citrus. Drought tolerance breeding has achieved 40-60% improvement in water use efficiency through selection of genotypes with enhanced root architecture and osmotic adjustment capacity. Salinity tolerance varies substantially among rootstock genotypes, with screening protocols identifying materials maintaining productivity at electrical conductivity levels exceeding 6 dS/m. Disease resistance breeding has produced rootstocks resistant to major soil-borne pathogens including Phytophthora, nematodes, and bacterial diseases in several fruit species. Graft compatibility mechanisms involve complex interactions between rootstock and scion at anatomical, physiological, and molecular levels that influence vascular connectivity and long-term graft success. Incompatibility symptoms ranging from poor initial union formation to delayed failure after years of apparent success reflect disruptions in these interaction processes. Molecular markers linked to rootstock traits enable early selection in breeding populations, substantially accelerating genetic gain compared to traditional field evaluation methods. Climate change adaptation will require rootstocks combining multiple stress tolerances, driving breeding strategies toward trait pyramiding approaches. This synthesis provides framework for understanding rootstock influences on fruit production and guides future breeding priorities.

Summary

Source-checked

Plain-language reading of the full paper, prepared with AI and checked against the text. Each point links to the authors' own words.

  1. Why it matters

    • Selecting the right rootstock is critical because it directly affects a fruit tree's vigor, how it acquires nutrients, its productivity, and its ability to withstand stress. View evidence ↗

      Why it matters

      Selecting the right rootstock is critical because it directly affects a fruit tree's vigor, how it acquires nutrients, its productivity, and its ability to withstand stress.

      In the paper's own words

      “Rootstock selection fundamentally influences fruit crop performance through effects on tree vigor, stress tolerance, nutrient acquisition, and scion productivity.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • In modern farming, rootstocks are essential for determining the physical structure of the orchard, how quickly trees bear fruit, and the overall economic success of the crop. View evidence ↗

      Why it matters

      In modern farming, rootstocks are essential for determining the physical structure of the orchard, how quickly trees bear fruit, and the overall economic success of the crop.

      In the paper's own words

      “Modern fruit production relies extensively on grafted trees where rootstock selection determines orchard architecture, precocity, stress tolerance, and ultimately economic viability.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
  2. What they did

    • The authors organized the gathered information by specific breeding goals and crop types to allow for comparisons between different species. View evidence ↗

      What they did

      The authors organized the gathered information by specific breeding goals and crop types to allow for comparisons between different species.

      In the paper's own words

      “Information was organized by breeding objective and fruit crop to facilitate comparative analysis across species.”

      Page 2

      Found in: Methods

      Read it in the full textOpen PDF at page 2 ↗
  3. What they found

    • Dwarfing traits have been the primary focus of breeding, leading to size-controlling rootstocks that allow for high-density planting in citrus, pear, and apple crops. View evidence ↗

      What they found

      Dwarfing traits have been the primary focus of breeding, leading to size-controlling rootstocks that allow for high-density planting in citrus, pear, and apple crops.

      In the paper's own words

      “Analysis of breeding progress reveals that dwarfing characteristics have received greatest attention, with successful development of size-controlling rootstocks enabling high-density planting systems in apple, pear, and citrus.”

      Page 1

      Found in: Discussion

      Read it in the full textOpen PDF at page 1 ↗
    • Breeding for drought tolerance has improved water use efficiency (WUE) by 40-60% by selecting for osmotic adjustment and better root architecture. View evidence ↗

      What they found

      Breeding for drought tolerance has improved water use efficiency (WUE) by 40-60% by selecting for osmotic adjustment and better root architecture.

      In the paper's own words

      “Drought tolerance breeding has achieved 40-60% improvement in water use efficiency through selection of genotypes with enhanced root architecture and osmotic adjustment capacity.”

      Page 1

      “Drought | Deep rooting, osmotic adjustment | Controlled deficit | 40-60% WUE improvement”

      Page 3

      Found in: Table 2

      Read it in the full textOpen PDF at page 1 ↗
    • Some rootstock genotypes have been identified that can maintain productivity even when soil electrical conductivity exceeds 6 dS/m. View evidence ↗

      What they found

      Some rootstock genotypes have been identified that can maintain productivity even when soil electrical conductivity exceeds 6 dS/m.

      In the paper's own words

      “Salinity tolerance varies substantially among rootstock genotypes, with screening protocols identifying materials maintaining productivity at electrical conductivity levels exceeding 6 dS/m.”

      Page 1

      “Salinity | Ion exclusion, compartmentation | Saline irrigation | Tolerant to 6 dS/m”

      Page 3

      Found in: Table 2

      Read it in the full textOpen PDF at page 1 ↗
    • Breeding for cold tolerance has successfully produced rootstocks that are hardy down to -30°C. View evidence ↗

      What they found

      Breeding for cold tolerance has successfully produced rootstocks that are hardy down to -30°C.

      In the paper's own words

      “Cold | Supercooling, bark hardiness | Freeze tests | Hardy to -30°C achieved”

      Page 3

      Found in: Table 2

      Read it in the full textOpen PDF at page 3 ↗
    • Breeding efforts have created rootstocks resistant to several major soil-borne threats, such as nematodes, bacterial diseases, and Phytophthora. View evidence ↗

      What they found

      Breeding efforts have created rootstocks resistant to several major soil-borne threats, such as nematodes, bacterial diseases, and Phytophthora.

      In the paper's own words

      “Disease resistance breeding has produced rootstocks resistant to major soil-borne pathogens including Phytophthora, nematodes, and bacterial diseases in several fruit species.”

      Page 1

      Found in: Introduction

      Read it in the full textOpen PDF at page 1 ↗
    • The use of molecular markers allows breeders to select traits at an early stage, which speeds up genetic improvement compared to waiting for field results. View evidence ↗

      What they found

      The use of molecular markers allows breeders to select traits at an early stage, which speeds up genetic improvement compared to waiting for field results.

      In the paper's own words

      “Molecular markers linked to rootstock traits enable early selection in breeding populations, substantially accelerating genetic gain compared to traditional field evaluation methods.”

      Page 1

      Found in: Discussion

      Read it in the full textOpen PDF at page 1 ↗
    • Graft incompatibility can range from an immediate failure to form a union to a delayed vascular failure that occurs years after the graft seemed successful. View evidence ↗

      What they found

      Graft incompatibility can range from an immediate failure to form a union to a delayed vascular failure that occurs years after the graft seemed successful.

      In the paper's own words

      “Incompatibility manifests through various mechanisms ranging from poor callus formation at the graft union to long-term vascular dysfunction appearing years after apparently successful grafting.”

      Page 4

      Found in: Discussion

      Read it in the full textOpen PDF at page 4 ↗
  4. What it means

    • The authors recommend that future efforts focus on combining multiple stress tolerances for climate adaptation, improving graft success through better compatibility research, and using molecular tools. View evidence ↗

      What it means

      The authors recommend that future efforts focus on combining multiple stress tolerances for climate adaptation, improving graft success through better compatibility research, and using molecular tools.

      In the paper's own words

      “Future priorities should emphasize climate adaptation through combined stress tolerance, understanding of compatibility mechanisms to improve graft success rates, and integration of molecular tools with conventional breeding to accelerate genetic gain.”

      Page 4

      Found in: Conclusion

      Read it in the full textOpen PDF at page 4 ↗
    • The review may underrepresent underutilized fruit crops because it focuses on species with a long history of rootstock breeding. View evidence ↗

      What it means

      The review may underrepresent underutilized fruit crops because it focuses on species with a long history of rootstock breeding.

      In the paper's own words

      “This review emphasizes species with substantial rootstock breeding history, potentially underrepresenting opportunities in underutilized fruit crops.”

      Page 4

      Found in: Limitations

      Read it in the full textOpen PDF at page 4 ↗
    • A limitation of the available data is that many proprietary breeding programs do not publish much of their development information. View evidence ↗

      What it means

      A limitation of the available data is that many proprietary breeding programs do not publish much of their development information.

      In the paper's own words

      “Proprietary breeding programs contribute substantially to rootstock development but publish limited information.”

      Page 4

      Found in: Limitations

      Read it in the full textOpen PDF at page 4 ↗

Visual summary

1 visuals

Illustrations explain the paper; they do not show study data.

AI-generated illustration
AI-generated conceptual illustration (grafted apple tree). It explains the idea and does not show study data.↗ Enlarge
Rootstock Influence and Breeding Goals
AI-generated conceptual illustration (grafted apple tree). It explains the idea and does not show study data.

AI-generated conceptual illustration (grafted apple tree). It explains the idea and does not show study data.

Open full-size image ↗

Rootstock Influence and Breeding Goals

AI-generated conceptual illustration (grafted apple tree). It explains the idea and does not show study data.

View source evidence ↗

Figures & tables

From the original paper
Summary of rootstock breeding objectives and progress in major fruit cropsTable 1Page 2Comparative stress tolerance responses of tolerant versus susceptible rootstock genotypes under progressive stress conditionsFigure 1Page 2Stress tolerance mechanisms in selected rootstock genotypesTable 2Page 3Graft compatibility matrix showing interaction scores between rootstock and scion combinationsFigure 2Page 3Relative breeding progress toward major rootstock improvement objectives across fruit crop programsFigure 3Page 3Conceptual model integrating genetic resources, selection criteria, and evaluation methods in rootstock breedingFigure 4Page 4
FULL TEXT Original article · reflowed for reading

Introduction

The practice of grafting fruit trees onto rootstocks of different genetic origin dates back thousands of years, enabling propagation of desirable scion varieties while exploiting rootstock characteristics for adaptation to challenging soil and environmental conditions [1] . Modern fruit production relies extensively on grafted trees where rootstock selection determines orchard architecture, precocity, stress tolerance, and ultimately economic viability. The diversity of rootstock genotypes available for major fruit species reflects accumulated selection from wild populations, controlled hybridization, and more recently, molecular breeding approaches [2] . Rootstock influences on scion performance operate through multiple mechanisms including water and nutrient acquisition efficiency, hormone signaling across the graft union, and physical restriction of assimilate transport [3] . Understanding these mechanisms enables targeted selection for specific production objectives and prediction of rootstock-scion interactions in novel combinations. The complexity of these interactions explains the extended evaluation periods traditionally required for rootstock development [4] . Climate change adaptation represents an emerging priority for rootstock breeding, demanding combined tolerance to drought, heat stress, and in some regions, salinity or flooding [5] . Traditional approaches relying on phenotypic selection in target environments require decades to develop adapted rootstocks, stimulating interest in marker-assisted methods that could accelerate genetic improvement. Integration of genomic tools with conventional breeding offers promise for addressing these complex breeding objectives [6] . This review synthesizes current understanding of rootstock breeding methods, graft compatibility mechanisms, and stress tolerance characteristics to provide framework for future research and breeding priorities [7] . The analysis encompasses major fruit crop species while identifying principles applicable across the broader range of grafted crops.

Material and Methods Material

This review synthesizes published research on fruit crop rootstock breeding from peer-reviewed journals, conference proceedings, and institutional technical reports [8] . Literature searches covered major databases using keywords related to specific rootstock traits, breeding methods, and fruit species. Studies were evaluated for relevance, experimental rigor, and practical applicability.

Methods

Information was organized by breeding objective and fruit crop to facilitate comparative analysis across species. Quantitative data on breeding progress and trait expression were compiled where available. Recommendations for future research priorities were formulated based on identified knowledge gaps and emerging production challenges [9] .

Genetic Resources

Wild species related to cultivated fruits represent primary genetic resources for rootstock breeding, harboring genes for stress tolerance and disease resistance absent in domesticated germplasm [10] . Systematic collection and evaluation of these materials has expanded the breeding gene pool available for hybridization programs. Interspecific hybridization creates novel genetic combinations that can express transgressive segregation for target traits, producing offspring exceeding both parents in stress tolerance or dwarfing ability.

Results

Results

Discussion

Graft compatibility represents a fundamental consideration in rootstock breeding that has received inadequate research attention relative to its practical importance [11] . Incompatibility manifests through various mechanisms ranging from poor callus formation at the graft union to long-term vascular dysfunction appearing years after apparently successful grafting. Understanding the molecular basis of compatibility would enable early screening of breeding populations, avoiding investment in genotypes destined to fail in commercial combinations. Molecular marker applications in rootstock breeding have expanded rapidly with decreasing genotyping costs and improving marker-trait associations [12] . Markers linked to dwarfing genes, disease resistance loci, and stress tolerance QTLs enable selection at seedling stages before field expression of mature phenotypes. This acceleration of genetic gain particularly benefits perennial crops where generation intervals traditionally span decades. Climate change adaptation requires rootstocks combining multiple stress tolerances that rarely occur together in existing germplasm [13] . Trait pyramiding through markerassisted backcrossing offers potential for assembling favorable alleles from diverse sources into elite backgrounds. However, the complex genetic architecture of most stress tolerances complicates marker-based selection, often requiring genomic selection approaches that consider genome-wide effects.

Limitations

This review emphasizes species with substantial rootstock breeding history, potentially underrepresenting opportunities in underutilized fruit crops. The variable quality of published research limits quantitative comparison across studies. Proprietary breeding programs contribute substantially to rootstock development but publish limited information. Long evaluation periods inherent in perennial crop assessment mean that newer breeding advances may not yet be fully documented in accessible literature.

Conclusion

Rootstock breeding has achieved substantial progress in developing dwarfing rootstocks for high-density systems, disease-resistant materials reducing pesticide dependence, and stress-tolerant genotypes expanding production into challenging environments. These advances have transformed fruit production practices and enabled significant productivity gains in adopting regions. Future priorities should emphasize climate adaptation through combined stress tolerance, understanding of compatibility mechanisms to improve graft success rates, and integration of molecular tools with conventional breeding to accelerate genetic gain. Collaboration among breeding programs to share germplasm and evaluation data would enhance efficiency across the research community. The framework presented in this review provides guidance for prioritizing research investments and breeding strategies in rootstock improvement programs seeking to address current and emerging production challenges in fruit crop cultivation.

Figures and tables from the paper

Table 1. Summary of rootstock breeding objectives and progress in major fruit crops
Fruit CropPrimary ObjectiveProgress LevelKey Achievements
AppleDwarfingHighM9, M26, G series rootstocks
CitrusDisease resistanceModerateTrifoliate hybrids, Swingle
Stone fruitsSize controlModerateKrymsk, Gisela series
GrapePhylloxera resistanceHighAmerican species hybrids
AvocadoRoot rot toleranceLow-ModerateDusa, Latas selections
MangoPolyembryonyLowSeed propagated selections

Table text extracted from the original PDF; superscript letters are shown inline. The original table image is below.

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Summary of rootstock breeding objectives and progress in major fruit crops↗ Enlarge
Table 1
Summary of rootstock breeding objectives and progress in major fruit crops

Summary of rootstock breeding objectives and progress in major fruit crops

Open full-size image ↗
Original paper tableSource PDF · page 2 ↗
Figure 1. Comparative stress tolerance responses of tolerant versus susceptible rootstock genotypes under progressive stress conditions
Comparative stress tolerance responses of tolerant versus susceptible rootstock genotypes under progressive stress conditions↗ Enlarge
Figure 1
Comparative stress tolerance responses of tolerant versus susceptible rootstock genotypes under progressive stress conditions

Comparative stress tolerance responses of tolerant versus susceptible rootstock genotypes under progressive stress conditions

Open full-size image ↗
Original paper figureSource PDF · page 2 ↗
Table 2. Stress tolerance mechanisms in selected rootstock genotypes
Stress TypeTolerance MechanismScreening MethodBreeding Progress
DroughtDeep rooting, osmotic adjustmentControlled deficit40-60% WUE improvement
SalinityIon exclusion, compartmentationSaline irrigationTolerant to 6 dS/m
FloodingAdventitious roots, aerenchymaWaterlogging trialsModerate progress
ColdSupercooling, bark hardinessFreeze testsHardy to -30°C achieved
HeatMembrane stabilityHigh temperature exposureLimited progress

Table text extracted from the original PDF; superscript letters are shown inline. The original table image is below.

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Stress tolerance mechanisms in selected rootstock genotypes↗ Enlarge
Table 2
Stress tolerance mechanisms in selected rootstock genotypes

Stress tolerance mechanisms in selected rootstock genotypes

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Original paper tableSource PDF · page 3 ↗
Figure 2. Graft compatibility matrix showing interaction scores between rootstock and scion combinations
Graft compatibility matrix showing interaction scores between rootstock and scion combinations↗ Enlarge
Figure 2
Graft compatibility matrix showing interaction scores between rootstock and scion combinations

Graft compatibility matrix showing interaction scores between rootstock and scion combinations

Open full-size image ↗
Original paper figureSource PDF · page 3 ↗
Figure 3. Relative breeding progress toward major rootstock improvement objectives across fruit crop programs
Relative breeding progress toward major rootstock improvement objectives across fruit crop programs↗ Enlarge
Figure 3
Relative breeding progress toward major rootstock improvement objectives across fruit crop programs

Relative breeding progress toward major rootstock improvement objectives across fruit crop programs

Open full-size image ↗
Original paper figureSource PDF · page 3 ↗
Figure 4. Conceptual model integrating genetic resources, selection criteria, and evaluation methods in rootstock breeding
Conceptual model integrating genetic resources, selection criteria, and evaluation methods in rootstock breeding↗ Enlarge
Figure 4
Conceptual model integrating genetic resources, selection criteria, and evaluation methods in rootstock breeding

Conceptual model integrating genetic resources, selection criteria, and evaluation methods in rootstock breeding

Open full-size image ↗
Original paper figureSource PDF · page 4 ↗

References

14 in the original paper
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  9. Gregory PJ, Atkinson CJ, Bengough AG. Contributions of roots and rootstocks to sustainable fruit production. Journal of Experimental Botany. 2013;64(5):12091222.
  10. Castle WS. A career perspective on citrus rootstocks. HortScience. 2010;45(1):11-15.
  11. Goldschmidt EE. Plant grafting: New mechanisms, evolutionary implications. Frontiers in Plant Science. 2014;5:727.
  12. Pilcher RL, Celton JM, Gardiner SE. Genetic markers linked to the dwarfing trait in apple rootstocks. Journal of the American Society for Horticultural Science. 2008;133(1):100-106.
  13. Serra I, Strever A, Myburgh PA. Review: The interaction between rootstocks and cultivars. Australian Journal of Grape and Wine Research. 2014;20(1):1-14.
  14. Reig G, Lordan J, Fazio G. Horticultural performance and elemental nutrient concentrations on Fuji grafted on apple rootstocks. Scientia Horticulturae. 2018;241:7178.