Etudes physiologiques et génétiques de caractères morpho-physico-chimiques des fruits d’agrumes au cours de la maturation jusqu’à l’abscission

Fruit quality control is an agronomic, genetics and production research objective. The concept of quality is crucial for fruit produced for the fresh fruit market. In citrus, which are non-climacteric fruit, fruit quality traits in the tree is set to trigger the harvest time that must precede fruit abscission. However, this fruit drop is sometimes very close to the stage of maturity and therefore causes losses because of shorter time to ensure the harvest. This is usually observed in orange grown in Spain and Tunisia. Furthermore, the quality of a citrus fruit is often defined by the skin color, fruit size, lack of seeds, aroma and juice content, sugar and acidity. If seedlessness can be obtained by induced mutations or triploidy, the control of the change in other characters is based on many factors: variety, interaction with the rootstock, environmental impact and cultivation techniques. Among them, the varietal genotype, which means the genetic structure inherited from the parents, is a key factor to achieve the expected level of improvement in breeding. In citrus, obtaining a genetic structure adapted to production targets is not easy because the characteristics associated with reproduction such as polyembryony, juvenility or gamete self-incompatibility. In the present work, we aimed to develop knowledge on the variation of fruit quality traits by including abscission, heritability and traits inheritance to facilitate breeding programs and breeding.We first studied the variation of the abscission in several varieties of orange on 3 sites, Tunisia, Spain and Corsica. This study showed that for the same varieties, the process of abscission measured by investigating the decrease the fruit detachment force (FDF) required to separate the fruit from the calyx, was very dependent on the environment, but not on the fruit quality traits. Interestingly, Corsica does not seem favorable to the expression of this trait. Among the environmental factors described on the three locations, the increase of the number of days that favor growth (average temperature ≥ 13 °C) in late winter seems to be the cause of the sudden and massive fruit drop. Nevertheless, in Corsica, the FDF can also decrease during maturation in other citrus than oranges.The analysis of inheritance of fruit quality traits and abscission was achieved by investigating the segregation of QTLs in a population of 116 hybrids resulting from a backcross (clementine × mandarin), clementine being itself a hybrid (mandarin × orange). Analyzes were replicated two consecutive years at several maturity dates which were bounded by maturity dates of both parents. Most traits presented a significant variation and a transgressive inheritance arising from the high heterozygosity of parental genomes. Three genetic maps have been developed (parental and consensus) using SSR and SNP markers covering about 75% of the reference genome. After estimating the random effect on the variance of the traits (BLUP), QTLs of each trait were detected (1 QTL for acidity, citrate, juice content, TSS, FDF as well as 5 QTLs for the a* color index). Most of them were detected at a single date of maturation.Interannual and inter-population stability of these QTLs will be checked before any possible use of the linked markers in breeding programs.Le contrôle de la qualité des fruits est un objectif de recherche agronomique, génétique mais également de production. La notion de qualité est cruciale pour les fruits destinés au commerce du fruit frais. Chez les agrumes, fruitier non climactérique, elle est définie sur l'arbre et utilisée pour déclencher la récolte qui doit précéder l'abscission des fruits. Néanmoins cette chute de fruit est parfois très proche du stade de maturité et occasionne donc des pertes du fait de délais trop courts pour assurer la récolte. Ceci est généralement observé chez les orangers cultivés en Espagne et en Tunisie. Par ailleurs, la qualité d'un agrume est souvent définie par la couleur de peau, la taille du fruit, l'absence de pépin, l'arôme et les teneurs en jus, en sucres et en acidité. Si l'absence de pépin peut être obtenue par la mutation induite ou par la triploïdie, le contrôle de la variation des autres caractères repose sur de nombreux facteurs: la variété, l'interaction avec le porte-greffe, les effets de l'environnement et les techniques culturales. Parmi eux le génotype variétal, c'est-à-dire sa structure génétique héritée de ses parents est un facteur prépondérant pour atteindre le niveau d'amélioration attendu. L'obtention d'une structure génétique adaptée aux objectifs de production n'est pas aisée car les caractéristiques liées à la reproduction telles que la polyembryonie, la phase juvénile ou l'auto-incompatibilité gamétique, sont contraignantes chez les agrumes. Dans le cadre de ce travail, nous avons eu pour objectif de développer des connaissances sur la variation des caractères de la qualité du fruit, en y incluant l'abscission, mais également sur leur héritabilité et leur hérédité afin de faciliter les programmes de sélection et de création variétale. Nous avons tout d'abord étudié la variation de l'abscission chez plusieurs variétés d'orangers sur 3 sites, en Tunisie en Espagne et en Corse. Cette étude démontre que pour des variétés identiques, le processus d'abscission mesurée via la diminution de la force de détachement du fruit du pédoncule (FDF), est très dépendante de l'environnement et non des caractères de qualité du fruit. De façon originale la Corse ne semble pas être favorable à l'expression de ce caractère. Parmi les facteurs environnementaux décrits sur les trois sites, celui de l'augmentation du nombre de jours favorable à la croissance (température moyenne ≥ 13°C) en fin d'hiver semble être à l'origine de la chute massive et soudaine des fruits. Néanmoins, en Corse, la FDF peut aussi diminuer au cours de la maturation sur d'autres agrumes que les oranges. L'analyse de l'hérédité des caractères de la qualité des fruits et de leur abscission a été réalisée par une approche de la ségrégation de QTLs dans une population de 116 hybrides issus d'un croisement de type backcross (clémentinier × mandarinier), le clémentinier étant déjà un hybride (mandarinier x oranger). Les analyses ont été reproduites sur deux campagnes de production à plusieurs dates de maturité et bornées par celles des deux parents. La plupart des caractères ont une variation importante et une hérédité transgressive découlant de l'hétérozygotie élevée des génomes parentaux. Trois cartes génétiques ont été développées (les parentales et la consensus) à l'aide de marqueurs SNP et SSR couvrant environ 75% du génome de référence. Après estimation de l'effet aléatoire sur la variance des caractères (BLUP), des QTLs de chacun des caractères ont été détectés (1 QTL pour l'acidité, le citrate, la teneur en jus, la TSS, la FDF à 5 QTLs pour l'indice a* de coloration). La plupart d'entre eux ne sont détectés qu'à une seule date de maturation. La stabilité interannuelle et inter-population de ces QTLs devra être vérifiée avant une possible utilisation des marqueurs liés dans les programmes d'amélioration

Abscission of Orange Fruit (Citrus sinensis (L.) Osb.) in the Mediterranean Basin Depends More on Environmental Conditions Than on Fruit Ripeness

Orange fruit abscission usually occurs at the final stage of fruit maturation but in some areas of citrus production, in advance of the usual harvest period, and sometimes suddenly and intensely. The reasons for this precocious citrus fruit abscission remains unclear. Therefore, the aim of this study was to try to clarify what the determinants of this phenomenon are. A multi-site experimentation was carried out on six orange cultivars, in Corsica, Spain and Tunisia where the phenomenon of early massive fruit drop varies. Climatic parameters, fruit maturity parameters and fruit detachment force (FDF) were recorded along the fruit maturation period. Respectively to the fruit drop, the FDF decreased in Tunisia and in Spain until the fruit falls, whereas in Corsica, it remained relatively constant throughout fruit maturation. Although data on fruit maturity parameters (rind color, acidity and total soluble solids) differed at the three sites, their evolution was similar during the period of maturation. FDF was not related to changes in any fruit maturity parameters, and more likely depended on changes in temperatures on days when the mean temperature was above 13 °C. Massive fruit abscission could be linked to the earlier more rapid restart of vegetative growth in Tunisia and Spain than in Corsica

Physiological and genetic studies of morpho-physico-chemical characteristics of citrus fruit during the ripening until abscission

Le contrôle de la qualité des fruits est un objectif de recherche agronomique, génétique mais également de production. La notion de qualité est cruciale pour les fruits destinés au commerce du fruit frais. Chez les agrumes, fruitier non climactérique, elle est définie sur l'arbre et utilisée pour déclencher la récolte qui doit précéder l'abscission des fruits. Néanmoins cette chute de fruit est parfois très proche du stade de maturité et occasionne donc des pertes du fait de délais trop courts pour assurer la récolte. Ceci est généralement observé chez les orangers cultivés en Espagne et en Tunisie. Par ailleurs, la qualité d'un agrume est souvent définie par la couleur de peau, la taille du fruit, l'absence de pépin, l'arôme et les teneurs en jus, en sucres et en acidité. Si l'absence de pépin peut être obtenue par la mutation induite ou par la triploïdie, le contrôle de la variation des autres caractères repose sur de nombreux facteurs: la variété, l'interaction avec le porte-greffe, les effets de l'environnement et les techniques culturales. Parmi eux le génotype variétal, c'est-à-dire sa structure génétique héritée de ses parents est un facteur prépondérant pour atteindre le niveau d'amélioration attendu. L'obtention d'une structure génétique adaptée aux objectifs de production n'est pas aisée car les caractéristiques liées à la reproduction telles que la polyembryonie, la phase juvénile ou l'auto-incompatibilité gamétique, sont contraignantes chez les agrumes. Dans le cadre de ce travail, nous avons eu pour objectif de développer des connaissances sur la variation des caractères de la qualité du fruit, en y incluant l'abscission, mais également sur leur héritabilité et leur hérédité afin de faciliter les programmes de sélection et de création variétale. Nous avons tout d'abord étudié la variation de l'abscission chez plusieurs variétés d'orangers sur 3 sites, en Tunisie en Espagne et en Corse. Cette étude démontre que pour des variétés identiques, le processus d'abscission mesurée via la diminution de la force de détachement du fruit du pédoncule (FDF), est très dépendante de l'environnement et non des caractères de qualité du fruit. De façon originale la Corse ne semble pas être favorable à l'expression de ce caractère. Parmi les facteurs environnementaux décrits sur les trois sites, celui de l'augmentation du nombre de jours favorable à la croissance (température moyenne ≥ 13°C) en fin d'hiver semble être à l'origine de la chute massive et soudaine des fruits. Néanmoins, en Corse, la FDF peut aussi diminuer au cours de la maturation sur d'autres agrumes que les oranges. L'analyse de l'hérédité des caractères de la qualité des fruits et de leur abscission a été réalisée par une approche de la ségrégation de QTLs dans une population de 116 hybrides issus d'un croisement de type backcross (clémentinier × mandarinier), le clémentinier étant déjà un hybride (mandarinier x oranger). Les analyses ont été reproduites sur deux campagnes de production à plusieurs dates de maturité et bornées par celles des deux parents. La plupart des caractères ont une variation importante et une hérédité transgressive découlant de l'hétérozygotie élevée des génomes parentaux. Trois cartes génétiques ont été développées (les parentales et la consensus) à l'aide de marqueurs SNP et SSR couvrant environ 75% du génome de référence. Après estimation de l'effet aléatoire sur la variance des caractères (BLUP), des QTLs de chacun des caractères ont été détectés (1 QTL pour l'acidité, le citrate, la teneur en jus, la TSS, la FDF à 5 QTLs pour l'indice a* de coloration). La plupart d'entre eux ne sont détectés qu'à une seule date de maturation. La stabilité interannuelle et inter-population de ces QTLs devra être vérifiée avant une possible utilisation des marqueurs liés dans les programmes d'amélioration.Fruit quality control is an agronomic, genetics and production research objective. The concept of quality is crucial for fruit produced for the fresh fruit market. In citrus, which are non-climacteric fruit, fruit quality traits in the tree is set to trigger the harvest time that must precede fruit abscission. However, this fruit drop is sometimes very close to the stage of maturity and therefore causes losses because of shorter time to ensure the harvest. This is usually observed in orange grown in Spain and Tunisia. Furthermore, the quality of a citrus fruit is often defined by the skin color, fruit size, lack of seeds, aroma and juice content, sugar and acidity. If seedlessness can be obtained by induced mutations or triploidy, the control of the change in other characters is based on many factors: variety, interaction with the rootstock, environmental impact and cultivation techniques. Among them, the varietal genotype, which means the genetic structure inherited from the parents, is a key factor to achieve the expected level of improvement in breeding. In citrus, obtaining a genetic structure adapted to production targets is not easy because the characteristics associated with reproduction such as polyembryony, juvenility or gamete self-incompatibility. In the present work, we aimed to develop knowledge on the variation of fruit quality traits by including abscission, heritability and traits inheritance to facilitate breeding programs and breeding.We first studied the variation of the abscission in several varieties of orange on 3 sites, Tunisia, Spain and Corsica. This study showed that for the same varieties, the process of abscission measured by investigating the decrease the fruit detachment force (FDF) required to separate the fruit from the calyx, was very dependent on the environment, but not on the fruit quality traits. Interestingly, Corsica does not seem favorable to the expression of this trait. Among the environmental factors described on the three locations, the increase of the number of days that favor growth (average temperature ≥ 13 °C) in late winter seems to be the cause of the sudden and massive fruit drop. Nevertheless, in Corsica, the FDF can also decrease during maturation in other citrus than oranges.The analysis of inheritance of fruit quality traits and abscission was achieved by investigating the segregation of QTLs in a population of 116 hybrids resulting from a backcross (clementine × mandarin), clementine being itself a hybrid (mandarin × orange). Analyzes were replicated two consecutive years at several maturity dates which were bounded by maturity dates of both parents. Most traits presented a significant variation and a transgressive inheritance arising from the high heterozygosity of parental genomes. Three genetic maps have been developed (parental and consensus) using SSR and SNP markers covering about 75% of the reference genome. After estimating the random effect on the variance of the traits (BLUP), QTLs of each trait were detected (1 QTL for acidity, citrate, juice content, TSS, FDF as well as 5 QTLs for the a* color index). Most of them were detected at a single date of maturation.Interannual and inter-population stability of these QTLs will be checked before any possible use of the linked markers in breeding programs

Abscission Study during Citrus Fruit Maturation in Corsica: Unfavorable Environmental Conditions for Fruit Shedding

International audienceCitrus fruit development and ripening are complex processes involving physiological and biochemical changes that are under hormonal, nutritional and environmental control. One of the most evident phenomena in late maturation is shedding of ripe fruit. A study made in Spain on sweet oranges showed that fruit shedding was related to the increase of sugars content of fruit pulp juice. To investigate this potential relationship between fruit abscission and internal maturity parameters we investigated the fruit maturity process of 10 clementine x mandarin hybrids and 9 commercial sweet orange varieties with different maturity period. Morphological and biochemical analyses (acidity, total soluble solids, fruit weight thickness of the flavedo, thickness of the peduncle, external colo) were carried out on fruit sets of the different varieties during the maturation process, from December to June. The abscission initiation was evaluated by measuring the force required to detach the fruit of the peduncle. Analysis showed that under the local conditions, abscission was generally not expressed even through fruit maturation was evolving. Therefore, we suppose that even though the local conditions have an important effect, abscission of citrus fruit results also from environmental-genotype interactions. Moreover, evolution of abscission and fruit parameters are not correlated suggesting the independence in the process of maturation and abscission under the environmental conditions of Corsica

Abscission of orange fruit (Citrus sinensis (L.) Osb.) in the Mediterranean basin depends more on environmental conditions than on fruit ripeness

Orange fruit abscission usually occurs at the final stage of fruit maturation but in some areas of citrus production, in advance of the usual harvest period, and sometimes suddenly and intensely. The reasons for this precocious citrus fruit abscission remains unclear. Therefore, the aim of this study was to try to clarify what the determinants of this phenomenon are. A multi-site experimentation was carried out on six orange cultivars, in Corsica, Spain and Tunisia where the phenomenon of early massive fruit drop varies. Climatic parameters, fruit maturity parameters and fruit detachment force (FDF) were recorded along the fruit maturation period. Respectively to the fruit drop, the FDF decreased in Tunisia and in Spain until the fruit falls, whereas in Corsica, it remained relatively constant throughout fruit maturation. Although data on fruit maturity parameters (rind color, acidity and total soluble solids) differed at the three sites, their evolution was similar during the period of maturation. FDF was not related to changes in any fruit maturity parameters, and more likely depended on changes in temperatures on days when the mean temperature was above 13 °C. Massive fruit abscission could be linked to the earlier more rapid restart of vegetative growth in Tunisia and Spain than in Corsica

Abscission Study during Citrus Fruit Maturation in Corsica: Unfavorable Environmental Conditions for Fruit Shedding

International audienceCitrus fruit development and ripening are complex processes involving physiological and biochemical changes that are under hormonal, nutritional and environmental control. One of the most evident phenomena in late maturation is shedding of ripe fruit. A study made in Spain on sweet oranges showed that fruit shedding was related to the increase of sugars content of fruit pulp juice. To investigate this potential relationship between fruit abscission and internal maturity parameters we investigated the fruit maturity process of 10 clementine x mandarin hybrids and 9 commercial sweet orange varieties with different maturity period. Morphological and biochemical analyses (acidity, total soluble solids, fruit weight thickness of the flavedo, thickness of the peduncle, external colo) were carried out on fruit sets of the different varieties during the maturation process, from December to June. The abscission initiation was evaluated by measuring the force required to detach the fruit of the peduncle. Analysis showed that under the local conditions, abscission was generally not expressed even through fruit maturation was evolving. Therefore, we suppose that even though the local conditions have an important effect, abscission of citrus fruit results also from environmental-genotype interactions. Moreover, evolution of abscission and fruit parameters are not correlated suggesting the independence in the process of maturation and abscission under the environmental conditions of Corsica

Effect of environment on citrus fruit abscission and maturation

International audienceFruit shedding is an important cause of citrus fruit loss in different citrus-producing areas in the world. This physiological process can occur in certain citrus cultivars during the final stage of fruit maturation. The fruit maturation process interacts with the environment and more precisely with climatic factors such as temperature. The maturity parameters that induce fruit drop and the interaction between citrus fruit abscission and climate are not known. To evaluate the effect of environment on fruit maturation and abscission, a multi-site study was carried out in Spain, Tunisia and Corsica on six orange cultivars. The examined parameters were fruit detachment force (FDF), weight, diameter, colour, total soluble sugar (TSS) and titratable acidity (TA). We observed that FDF did not evolve in the same way in the three sites. While in Tunisia and in Spain it decreased, in Corsica, it remained constant throughout the fruit maturation process whatever the orange cultivar. Variation of temperature and radiation measured between the three sites cannot be the direct factors controlling this fruit process. Moreover, FDF seems not to be dependent on the evolution of any fruit quality parameters. Considering these results, fruit abscission and fruit maturation could be two independent phenomena at least under the environmental conditions in Corsica

Mapping of QTLs for citrus quality traits throughout the fruit maturation process on clementine (Citrus reticulata × C. sinensis) and mandarin (C. reticulata Blanco) genetic maps

International audienceCitrus fruit quality is defined as the combination of physical and chemical traits; some of which may change during the ripening phase, e.g., acidity and sugar content. A clear understanding of their genetic control would be very helpful for marker-assisted breeding programs especially with regard to the juvenile phase and some reproductive features that hamper the selection of improved hybrids. A genetic study was thus performed on the heredity of quality traits and QTL detection based on segregation in a progeny generated from a cross between clementine cv “Commun” ( Citrus × reticulata cv clementine ) and mandarin cv “Willow leaf” ( C. reticulata Blanco). Parental and consensus genetic linkage maps were constructed using 645 SNP and SSR markers. These maps were represented by 10 linkage groups in clementine and 12 linkage groups in mandarin, representing 75% and 58% respectively of the previously published clementine reference map. A total of 16 traits, including fruit mass, equatorial diameter, juice percentage, total soluble solids, acidity, pH, glucose, fructose, sucrose, and citric and malic acid concentrations were evaluated at three maturation dates. High variations indicating transgressive segregation were found for all traits, with normal or close to normal distributions. QTL analysis performed using the multiple QTL model allowed the detection of 34 QTLs on the three maps. QTLs were distributed in different linkage groups and generally detected at only one date of the ripening phase. The percentage of total variation explained ranged from 12 to 37% per QTL. Major QTLs ( R 2 ≥ 30%) were detected for equatorial diameter, glucose, and fructose (expressed in percentage dry matter) on linkage groups 8 and 9. Co-localization of QTLs controlling correlated and uncorrelated traits were mainly found on linkage groups 2, 4, 8, and 9, particularly between fruit mass and acidity