Abstract

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 (h2) 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.

Highlights

  • How different genomes interact to shape a unique phenotype has been one of the most pervasive questions in quantitative genetics and molecular evolution (Lynch, 2007; Bijma, 2013; Fisher and Mcadam, 2019)

  • The only morphological trait that we found having a significant genetic-estimated heritability value was trunk height, likely an emergent property of the rootstock–scion interaction in orchards of the same age only subjected to comparable annual light correctional pruning

  • There were significant rootstock effects for various harvest and quality traits such as total number of fruits, number of fruits with exportation quality, and number of fruits discarded because of low weight and damage by thrips. These findings suggest the inheritance of rootstock effects on a wide spectrum of “Hass” avocado traits relevant for yield, which will be critical to meet the demands of the growing worldwide market

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Summary

Introduction

How different genomes interact to shape a unique phenotype has been one of the most pervasive questions in quantitative genetics and molecular evolution (Lynch, 2007; Bijma, 2013; Fisher and Mcadam, 2019). A commonly disregarded yet ancient process that produces genetic chimeras is grafting, which refers to the agricultural practice that joins the root system (rootstock) of one plant, usually a woody crop, to the shoot (scion) of another (Warschefsky et al, 2016; Gautier et al, 2019; Bartusch and Melnyk, 2020). Modern grafting is crucial for the clonal propagation of fruit trees (e.g., avocado, citrus, grapevine and peach) and the establishment of seed orchards for the wood industry—i.e., pines, teak (Tuskan et al, 2018). Grafting is common in a phylogenetically diverse assortment of fruit and forest tree species, so it offers an irreplaceable experimental playground to study the rootstock– scion interaction (Albacete et al, 2015)

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