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Molecular Assisted Selection for Pollination-Constant and Non-Astringent Type Without Male Flowers in Spanish Germplasm for Persimmon Breeding

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Molecular Assisted Selection for Pollination-Constant and Non-Astringent Type Without Male Flowers in Spanish Germplasm for Persimmon Breeding agronomy Article Molecular Assisted Selection for Pollination-Constant and Non-Astringent Type without Male Flowers in Spanish Germplasm for Persimmon Breeding Manuel Blasco 1, Francisco Gil-Muñoz 2, María del Mar Naval 1 and María Luisa Badenes 2,* 1 CANSO, Mestre Serrano, 1, 46250 L’Alcúdia, Valencia, Spain; [email protected] (M.B.); [email protected] (M.d.M.N.) 2 Instituto Valenciano de Investigaciones Agrarias, CV 315 km 10,5., 46113 Moncada, Valencia, Spain; [email protected] * Correspondence: [email protected] Received: 14 July 2020; Accepted: 7 August 2020; Published: 11 August 2020 Abstract: Persimmon (Diospyros kaki Thunb) species is a hexaploid genotype that has a morphologically polygamous gyonodioecious sexual system. D. kaki bears unisexual flowers. The presence of male flowers resulted in the presence of seeds in the varieties. The fruits of persimmon are classified according to their astringency and the pollination events that produced seeds and modify the levels of astringency in the fruit. The presence of seeds in astringent varieties as pollination variant astringent (PVA), pollination variant non-astringent (PVNA) and pollination constant astringent (PCA) resulted in fruits not marketable. Molecular markers that allow selection of the varieties according to the type of flowers at the plantlet stage would allow selection of seedless varieties. In this study, a marker developed in D. lotus by bulk segregant analysis (BSA) and amplified fragment length polymorphism (AFLP) markers, named DlSx-AF4, has been validated in a germplasm collection of persimmon, results obtained agree with the phenotype data. A second important trait in persimmon is the presence of astringency in ripened fruits. Fruits non-astringent at the ripen stage named pollination constant non-astringent (PCNA) are the objective of many breeding programs as they do not need removal of the astringency by a postharvest treatment. Astringency in the hexaploid persimmon is a dominant trait. The presence of at least one astringent allele confers astringency to the fruit. In this paper we checked the marker developed linked to the AST gene. Our goal has been to validate both markers in germplasm from different origins and to test the usefulness in a breeding program. Keywords: persimmon; sex determination; fruit astringency; molecular markers 1. Introduction Persimmon (Diospyros kaki Thunb) species is a hexaploid genotype that has a morphologically polygamous gyonodioecious sexual system [1]. D. kaki bears unisexual flowers, as do other Diospyros species. There are genotypes that bear only female flowers and genotypes bearing male and female flowers [2]. Furthermore, varieties bearing only male flowers were described in China [3] and occasional male flower formation was reported in varieties that usually bear only female flowers [4]. In addition to the unisexual flowers, some varieties or genotypes bear hermaphrodite flowers; however, these flowers do not function fully as female flowers [5]. Most of the commercial varieties present only female flowers [6], however the presence of the male flowers type is important in two scenarios: first when the production of seeded fruits is convenient and second in breeding activities in which crosses are requested. The fruits of persimmon are classified according to their astringency and the pollination events that resulted in different types of fruits. The PCNA (pollination constant non-astringent) varieties Agronomy 2020, 10, 1172; doi:10.3390/agronomy10081172 www.mdpi.com/journal/agronomy Agronomy 2020, 10, 1172 2 of 11 Agronomy 2020, 10, x FOR PEER REVIEW 2 of 11 are alwaysscenarios: not astringent first when at the maturity production regardless of seeded fruits of pollination is convenient events and second and in the breeding presence activities or absence of in which crosses are requested. seeds in the fruit. The presence of male flowers in these varieties is most convenient since it allows The fruits of persimmon are classified according to their astringency and the pollination events pollinationthat and resulted produces in different seeded types fruits of fruits. that The increase PCNA (pollination the fruit size constant and no weight.n-astringent) In Japan, varieties the are presence of seeds in thealways fruit not does astringent not aff ectat maturity the consumers regardless demand of pollination [7]. However,events and the in Europepresence andor absence western of countries consumersseeds prefer in the seedless fruit. The fruits. presence of male flowers in these varieties is most convenient since it allows pollination and produces seeded fruits that increase the fruit size and weight. In Japan, the presence Three additional variety types can be distinguished: PVNA-type (pollination variant of seeds in the fruit does not affect the consumers demand [7]. However, in Europe and western non-astringent),countries which consumers are prefer non-astringent seedless fruits. varieties when seeds are present; PVA-type (pollination variant astringent),Three additional which are variety astringent types varietiescan be distin inguished: most parts PVNA-type of the fruit(pollination and non-astringent variant non- around the seedsastringent), if they are which present, are non-astringent and PCA-type varieties (pollination when seeds are constant present; PVA-type astringent), (pollination which variant are astringent astringent), which are astringent varieties in most parts of the fruit and non-astringent around the varieties regardless the presence of seeds. The loss of astringency in these types of persimmons is seeds if they are present, and PCA-type (pollination constant astringent), which are astringent associatedvarieties to the abilityregardless of thethe presence seeds to of produce seeds. The acetaldehyde. loss of astringency This in productionthese types of resulted persimmons in browningis of the flesh aroundassociated the to the seeds ability (Figure of the s1eeds), which to produce interfere acetaldehyde. with theThis postharvest production resulted treatment in browning for removing the astringencyof the flesh in thearound fruit, the seeds all together (Figure 1), the which PVA inte andrfere PCAwith the type postharvest fruits aretreatment unmarketable for removing if they are the astringency in the fruit, all together the PVA and PCA type fruits are unmarketable if they are pollinatedpollinated and the fruitsand the present fruits present seeds [seeds8,9]. In[8,9]. PVA In andPVA PCAand PCA varieties varieties it is it crucialis crucial to avoidto avoid pollination, hence thepollination, presence hence of male the presence flowers of in male the flowers variety in andthe variety in the and vicinity in the vicinity of the of crop the crop should should be avoided. The abilitybe of avoided. the production The ability ofof the male production flowers of is male a genetic flowers trait is a genetic that should trait that be should determined be determined in the varieties for avoidingin the seeds varieties in astringent for avoiding varieties seeds in astringent or improving varieties the or presence improving ofthe them presence in non-astringent of them in non- varieties. astringent varieties. Figure 1. Phenotypes of the traits selected. (A,B) flowers from the variety ‘Cal Fuyu’, female and male respectively and (C,D) results of pollination on astringent fruits from ‘Rojo Brillante’ a pollination variant astringent (PVA) variety: (C) parthenocarpic non-pollinated fruits and (D) pollinated fruits in which presence of seeds resulted in no marketable fruits. Elucidation of the genetic and molecular basis of sex expression in D. kaki leading to the development of molecular markers would allow selection of the varieties according to the type of flowers, being a great contribution for persimmon production and Diospyros breeding. The hexaploidy of D. kaki made elucidation of this question more difficult than in diploid genotypes. Since the genus includes more than 700 species with different levels of polyploidy, the diploid Diospyros lotus was Agronomy 2020, 10, 1172 3 of 11 used for investigation of the sex expression into the genus [10]. These authors described the model of inheritance and developed molecular markers associated to sex expression. Later small RNA acting as a sex determinant was identified [11]. Development of markers used the bulk segregant analysis (BSA) and amplified fragment length polymorphism (AFLP). An AFLP marker identified as DlSx-AF4 was sequence-characterized and converted into the sequence characterized amplified region (SCAR) [10]. In this study the marker has been tested in a germplasm selection of varieties phenotyped for sex expression and a backcross population obtained at Instituto Valenciano de Investigaciones Agrarias (IVIA). The results provide evidence of the usefulness of molecular marker assistance in identifying the genetic potential of production of male flowers in persimmon, an important trait in breeding. PCNA varieties are highly desired because their mature fruits are not astringent, as they stop accumulating tannins at early steps of fruit development [12]. In Japanese varieties, the PCNA trait is recessive to the non-PCNA trait [13] and is controlled by a single locus, AST [14]. Due to persimmon being a hexaploid, the PCNA type should contain six recessive ast alleles [15]. In breeding programs aimed at obtaining PCNA cultivars, the hexaploidy of persimmon along with the recessive inheritance of the non-astringency trait led to breeders to develop crosses
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