Characterization of fruit quality traits for organic acids content and profile in a large peach germplasm collection
Introduction
Peach [Prunus persica L. (Batsch.)] is a perennial crop setting climacteric fleshy fruits widely cultivated in the temperate regions, mainly. Annual peach production in Europe has showed a decreasing trend in the last decade (from about 4 Mt in 2008 to 3.5 Mt in 2018), following a reduction of fresh consumption, likely due to the overall low quality of commercialized fruits (Etienne et al., 2002; Cirilli et al., 2016). Many efforts have been done in the last years in developing novel varieties with improved external fruit quality, related to size, texture and skin colour (Dirlewanger et al., 1999; Fideghelli et al., 1998; Moreno, 2005; Cantín et al., 2009, 2010). Nevertheless, a poor taste or lack of the characteristic “peach” aroma still remains the major consumers’ complaint (Bassi and Selli, 1990; Cirilli et al., 2016). Among basic tastes, sweetness and sourness are the prominent eating quality attributes in peach (Crisosto et al., 2006). Their perception mostly depends on the relative abundance of sugars (the most abundant water-soluble compound) and acidity levels (dependent to the organic acid content, particularly malic and citric) (Vizzotto et al., 2002; Colaric et al., 2005; Batista-Silva et al., 2018). Acidity is a major driver of fruit quality perception in peach directly affecting consumers’ liking degree. Peach varieties are usually classified as normal or low-acid, based on fruit pH respectively below 3.8 or higher than 4 (Yoshida, 1970; Dirlewanger et al., 1998). The low-acid (LA) type has been often classified as the so called ‘honey peach’ group, widely spread in the Far-East countries (China, Korea and Japan) (Reimer, 1906). In contrast, most of the European and USA varieties tend to have normal-acid (NA) fruits. The low-acid trait has been further elucidated, revealing a simple, dominant genetic inheritance (locus D, for Douceur) (Monet, 1979; Boudehri et al., 2009), making this phenotype a relatively easy breeding target (Byrne, 2002). Apart from the D locus, titratable acidity and pH still show a remarkable variability, as suggested by an early study of Yoshida (1970). The content and profile of organic acids (OAs) seem to be quantitatively and qualitatively variable among accessions. OAs are responsible of protons bulk in mesocarp cells that determines the titratable acidity of fruits (Famiani et al., 2020). Acidity correlates with taste and aroma perception: sweetness is influenced by citric and shikimic acids and sugars/organic acids ratio, while aroma correlates with total acidity and malic acid content (Colaric et al., 2005). Peach accumulates mainly malate (dicarboxylic acid) and citrate (tricarboxylic acid) in the flesh, while shikimate, fumarate, oxalate, succinate, quinate and ascorbate are reported at low contents (Moing and Svanella, 1998; Etienne et al., 2013; Nowicka et al., 2019; Zheng et al., 2021). Total OAs content range from about 400 to over 3000 mg L−1 (Cao et al., 2016), being malic and citric acids the most abundant averaging from 300 to 600 and 100–300 mg kg−1 of fresh weight, respectively. The OAs content and profile are markedly regulated by fruit development and ripening, as well as post-harvest storage conditions and treatments, although generally well-correlated across seasons (Cano-Salazar et al., 2013; Desnoues et al., 2014). In NA accessions, malate content remains relatively constant along fruit development and ripening, while in LA accessions it tends to decrease. Citrate tends to be accumulated from pre-climateric stage onwards in NA, and only in reduced amount in LA (Moing and Svanella, 1998; Lombardo et al., 2011). Differences among varieties for both titratable acidity and OAs content seem to be mainly dependent to their aptitude to store OAs in the vacuole of mesocarp cells rather than metabolic synthesis or degradation (Moing and Svanella, 1998), although other factors are also reported to play a role in citrate accumulation (Zheng et al., 2021). In LA fruits, some divergence in expression of vacuole proton pumps may explain the cells’ reduced capability to sequester OAs, thereby making them accessible to catabolic enzymes (Etienne et al., 2002). Higher levels of PEPC activity and reduced accumulation of sugars in the LA fruits during ripening may suggest that the acids are constantly recycled (Moing et al., 2000).
This study reports an analysis of fruit acidity and OAs content in a large and genetically diverse peach germplasm collection with the aim of providing valuable information on this important class of metabolites to support eating quality-oriented breeding programmes.
Section snippets
Plant material and experimental design
The peach collection used in this study contains genetically diversified materials of different origins, including traditional accessions, breeding selection and modern varieties (Laurens et al., 2018). The collection was maintained at the ‘M. Neri’ farm of Centro Ricerche Produzioni Vegetali (CRPV) located in Imola (Bologna county, Northern Italy; 44.162628 °N, 12.219135 °E). Trees were grown on GF- 677 rootstock. A total of 138 and 183 peach accessions were assessed for several fruit quality
Fruit titratable acidity
A number of 117 and 180 accessions and selections were recorded for titratable acidity (TA) in the seasons 2017 and 2018, respectively, for a total of 201 unique accessions (Supplemental Table A. 1). In the 96 accessions common to both seasons, TA was highly correlated, indicating low seasonal influence on this trait (Supplemental Table A. 2). The distribution and frequency of across-year average TA values observed for the 201 unique genotypes are reported in Fig. 1. According with the density
Discussion
An extended characterization of fruit quality traits in a large collection of peach accessions and selections was described in this study. The primary purpose was to provide a cutting-edge exploration of the phenotypic diversity associated with parameters contributing to fruit eating quality, in particular titratable acidity and organic acids content. Titratable acidity (TA) is an important quality indicator and a major driver of consumer’s preference; it directly affects sweetness and aroma
Conclusions
The narrow diversity of the recent peach varieties released on the market has hampered the effort of increasing market quality. This study has revealed a high diversity for acidity-related traits in the peach, providing a pool of phenotypic data for both the selection of existing varieties and their development. Beyond malate and citrate, this work highlighted the presence and contribution of other organic acids showing a genotype-dependent profile. Their contribution to sensory quality and
Author contributions
IB: performed phenotypic analysis, analysed the data collected and drafted the manuscript; CR: performed phenotypic analysis; BD: provided plant materials, supported work planning and revised manuscript; GC: helped in phenotypic analysis; CM and SA conceived the work and revised manuscript.
Funding
This work has been partially funded in the framework of the MAS.PES (Italian project aimed at apricot and peach breeding) for design of the study, plant material and phenotyping and partially supported by the PRIMA-FREECLIMB international project.
Declaration of Competing Interest
The authors report no declarations of interest.
Acknowledgements
The authors wish to thank S. Foschi and M. Lama (CRPV, Cesena, Italy) for technical assistance in field operations.
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