Rabbiteye and lowbush blueberries are also largely self-infertile and therefore cross pollination isrequired for the good fruit set and berry size. Bumble bees and wild bees pollinate blueberryflowers naturally. Cranberries are self-pollinated but insect or wind disturbance is needed for thepollen release (Rieger). Most wild Vaccinium species are cross pollinated by insects. Selfpollinationis possible in bilberry and lingonberry but it reduces seed production (Nuortila et al.2002). Fruit development. The fruit of the Vaccinium species is fleshy, round or oval by shape, theycontain several to many seeds and belong in the category of berries. More strictly, the Vacciniumfruits are classified as false or epigynous berries, in which the fruit is formed from an inferiorovary, but the floral tube ripens along with the ovary. In true berries the entire ovary wall ripensinto a soft pericarp (Fahn 1990). The fruits can grow clustered like blueberries and lingonberries orsingly in branch or leaf axis like bilberries or cranberries.Most Vaccinium species growing in the Northern hemisphere flower from May to June and produceberries from July to October. The fruit development of blueberries takes 45 to 90 days, and varies alot between different species, cultivars, and growth locations. The growth of most blueberries,especially later ripening cultivars, has been found to exhibit a double sigmoid pattern (Godoy et al.2008). Cranberry fruits mature in 60-120 days after fertilization, depending on the cultivar andweather. Among the cultivated blueberries highbush blueberries reach maturity faster (typically 45-75 days) compared to lowbush or rabbiteye blueberries (Rieger).The development of bilberries from flower to ripe fruit lasts usually 55 to 70 days, varying betweendifferent years. The different phases of bilberry fruit development and ripening are presented infigure 1. Typically, the beginning of fruit development involves divisions and expansions of thecells (stages 2 – 3). During the ripening (stages 4 – 6) the accumulation of anthocyanins, and othersecondary metabolites and sugars as well as the softening of the cell walls occur. Compared to thebilberry, lingonberry flowers and fruits ripen later during the season, approximately 78 – 84 daysafter full blossom (Gustavsson 2001).Phenolic compounds. The berries of the Vaccinium species are known especially for their highcontent of phenolic compounds that are strong antioxidants and possess health beneficialproperties. Vaccinium berries are among the best sources of anthocyanins and proanthocyanidins(Ovaskainen et al. 2008). Therefore, the accumulation of phenolic compounds has been one of thefocus areas in the study on the fruit development of the Vaccinium species. The composition offlavonoids and other phenolic compounds in fruit development and ripening has been analysed inbilberries, cranberries, and blueberries (Jaakola et al. 2002, Vvedenskaya and Vorsa 2004,Castrejon et al. 2008, Celik et al. 2008). The overall profile of phenolic compounds at theirdifferent stages of development shares similarities between the examined Vaccinium species.Figure 1. Fruit development and the ripening of bilberry from pollination to ripe fruit.At the beginning of fruit development proanthocyanidins and flavonols are the main flavonols inVaccinium fruits. At the onset of the ripening, the content of proanthocyanidins decreases at thesame time anthocyanins begin to accumulate. A high content of proanthocyanidins at the earlyphases of fruit development has also been detected from fruits other than the Vaccinium species.Proanthocyanidins have been suggested to provide protection against fungal pathogens and thepredation of unripe fruits (Harborne 1997).40
The levels of flavonols are more constant over the period of fruit development, although especiallyqurcetin glycosides are found to be in slightly higher level at the beginning of the fruitdevelopment. In addition to quercetin glycosides, many Vaccinium berries contain myricetinglycosides (Määttä-Riihinen et al. 2004). In the bilberry, myricetin glycosides were found toaccumulate during the ripening phase, along with anthocyanins (Jaakola et al. 2002). Of thephenolic compounds, in addition to flavonoids Vaccinium berries contain hydroxycinnamic acids.Blueberries contain high levels of caffeic acids (Määttä-Riihinen et al. 2004). The antioxidantactivity during fruit development has been shown to be associated with the total phenolic content atthe different developmental stages (Castrejon et al. 2008, Celic et al. 2008).Other nutraceuticals. There is a lot of information on the nutritional value of Vaccinium berries, butit is mainly focused on ripe fruits. Only a limited number of studies report the contents of thenutritional compounds during fruit development. Cano-Merdano and Darnell (1997) analysed sugaraccumulation during the development of rabbiteye blueberries. Glucose and fructose are the mainsugars in Vaccinium fruits, whereas the contents of sucrose are low (Cano-Merdano and Darnell1997, Viljakainen et al. 2002). According to studies of rabbiteye blueberries, the levels of sugarswere at their lowest about 20 days after bloom, after which the contents increased up to ten fold.The maximum levels were reached between 60 and 90 days after bloom, reaching the maximum inthe ripe fruits.Celic et al. (2008) analysed the content of organic acids at four different maturity stages ofcranberries. The most abundant organic acid in cranberries was citric acid (73 %) followed bymalic and ascorbic acid. The overall concentration of citric and malic acid increased over theripening, whereas the ascorbic acid concentration decreased. Regulation of fruit development.Recent discoveries have begun to reveal the developmental cues that are responsible for primingand initiating the ripening in fruit bearing species (Giovannoni 2004, 2007). The tomato (Solanumlycopersicum) has been the model system of choice due to its genetic resources and the knowledgethat researchers have of its ripening physiology and biochemistry of ripening (Seymour et al.,2008). However, key information has also been obtained from the model plant Arabidopsis, and toa lesser extent from strawberries (Fragaria spp).The current state of the art is that a number of ripening related genes are known from these speciesincluding the ripening inhibitor (rin) and the Colourless non-ripening (Cnr) (Giovannoni, 2007).The gene at the tomato ripening inhibitor (rin) locus is a member of the MADS-box SEPALLATA(SEP) sub-family; LeMADS-RIN (Vrebalov et al., 2002). MADS-box genes were previouslyassociated with floral development, but LeMADS-RIN is necessary for ripening. The gene at theCnr locus encodes an SBP-box transcription factor (Manning et al., 2006), that is likely to interactwith the promoters of the SQUAMOSA (SQUA) sub-family of MADS-box genes (Lännenpää et al.,2004). These include TDR4, which shows enhanced expression during tomato fruit ripening(Eriksson et al., 2004), but has yet to be assigned a function. This regulatory network appears to beconserved across fruit bearing species.Nonetheless, links between the regulatory factors and the down stream effectors are poorlyunderstood and thus far the studies are limited almost exclusively to the tomato, despite the diverseripening behaviour of other important fruiting species.Studies clarifying the molecular basis of fruit development in the Vaccinium species have, untilrecently, been limited and nothing is known about the genes controlling the fruit ripening in theVaccinium species. Some structural genes encoding the key enzymes of the flavonoid biosyntheticpathway have been characterised from cranberry (Polaschock et al., 2002) and bilberry fruits(Jaakola et al., 2002). Just recently, two SQUA sub-family of MADS-box genes have been clonedfrom the bilberry (Jaakola et al., unpublished). The other of the two cloned transcription factorgenes was related by sequence homology with tomato TDR4 and Arabidopsis FRUITFULL (FUL)genes. The functional analyses revealed a hitherto unsuspected link between the SQUA MADS-boxgene and the production of secondary metabolites.ConclusionsThe genus Vaccinium includes several economically important berry species. Therefore, the studyon the quality characteristics over the development and ripening of the fruits can open newcommercially exploitable applications in the future. Understanding better the gene x environment41
- Page 3 and 4: Conference Organizing CommitteeChai
- Page 6 and 7: 15 Pormale J., Osvalde A. and Nolle
- Page 8 and 9: were established in 1985. Nowadays,
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- Page 45 and 46: Figure 1. A general scheme of the N
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the mean. Kisgyır 1 sample has the
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13. Porpáczy A. (1999) A húsos so
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was medium (0.014 - 0.017 g kg -1 s
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‘Salaspils Ražīgā’. Vigorous
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KopsavilkumsEiropas melleĦu (Vacci
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Figure 2. Chemometric PCA of 32 blu
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References1. Baloga D.W., Vorsa N.,
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obtained from fruits of black choke
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In our opinion, the best estimate a
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cuttings also varies markedly with
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shoots shorter than 10 mm were not
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14. Ostrolucka M.G., Gajdosova A, L
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„Metos RG-350” (http://www.meto
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500480Phenols,mg 100g -146044042040
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SHORT INFORMATION ABOUT THE HISTORY
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Evaluation of cultivars. After the
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the number of pistils (female clone
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Table 2. Number of flowers per harv
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ResultsFirst time upright dieback i
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grew rapidly on PDA at 20 - 24 o C.
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Figure 9. Conidia of Physalospora v
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References1. CABI, EPPO, (1997) Dia
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Results und DiscussionBerries were
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In literature Caruso eds. and Гop
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the total area under a cranberry ma
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Skilled works on development of the
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Tika atrastas dažas būtiskas ats
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appears to maintain a quite low lev
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8. Garkava - Gustavson L.,Persson H