In grapes of Cabernet Sauvignon, berry growth, content of anthocyanin and hormonal composition affected by seed number were investigated.Gibberellin-like substance content showed highest values 45 d after anthesis, while abscisic acid content (on a per-berry basis) had two peaks, the first one just after berry set, the second coinciding with ripening phenomenon.Seed number was positively correlated with the cöncentrations of gibberellinlike substances and abscisic acid (on a per-berry basis). One-seeded berries, however, had a higher amount of anthocyanin than two or three-seeded ones.Beziehungen zwischen Anzahl der Samen sowie Gibberellin- und Abscisinsäuregehalt und Reifeverlauf der Traubenbeeren bei Cabernet SauvignonBei den Trauben von Cabernet Sauvignon wurden das Beerenwachstum, der Anthocyangehalt und die Zusammensetzung der Phytohormone in Beziehung zur Anzahl der Samen untersucht.Die Gibberellinaktivität erreichte 45 d nach der Anthese ihren höchsten Stand, während der Abscisinsäuregehalt (je Beere) zwei Maxima zeigte; das erste trat kurz nach dem Beerenansatz auf, das zweite fiel mit dem Beginn der Beerenreife zusammen. Die Anzahl der Samen war positiv mit der Gibberellinaktivität und der Abscisinsäurekonzentration (je Beere) korreliert. Einsamige Beeren. wiesen jedoch einen höheren Anthocyangehalt als zwei- oder dreisamige Beeren auf

Fregoni, M.

Miravalle, R.

Scienza, A.

Visai, C.

English

JKI Open Journal Systems (Julius Kühn-Institut)

Vitis 17, 361-368 (1978) Department of Viticulture, Catholic University, Piacenza, Italy Relationships betwe~n seed number, gibberellin and abscisic acid levels and ripening in Cabernet Sauvignon grape berries1) by A. ScrnNzA2), R. MrnAVALLE, C. V1sA18) and M. FREGONI Beziehungen zwischen Anzahl der Samen sowie Gibberellin- und Abscisinsäure-gehalt und Reifeverlauf der Traubenbeeren bei Cabernet Sauvignon Z u s a m m e n f a s s u n g . - Bei den Trauben von Cabernet Sauvignon wurden das Beerenwachstum, der Anthocyangehalt und die Zusammensetzung der Phytohormone in Beziehung zur Anzahl der Samen untersucht. Die Gibberellinaktivität erreichte 45 d nach der Anthese ihren höchsten Stand, wäh-rend der Abscisinsäuregehalt (je Beere) zwei Maxima zeigte; das erste trat kurz nach dem Beerenansatz auf, das zweite fiel mit dem Beginn der Beerenreife zusammen. Die Anzahl der Samen war positiv mit der Gibberellinaktivität und der Abscisinsäu-rekonzentration (je Beere) korreliert. Einsamige Beeren. wiesen jedoch einen höheren Anthocyangehalt als zwei- oder dreisamige Beeren auf. Introduction Considerable research has proved a definite link between seed number, berry development and its sugar and acid content (W1NKLER and WILLIAMS 1936, Oum 1946, GÄRTEL 1954, PAsTENA 1964, ScHUMANN 1973, RAPP and KtENERT 1974). Such relationship has been attributed to hormones, such as auxins, gibberellins, cytokinins, formed in the seeds and spread into the pulp, thus promoting cell division and extension and changing pulp composition (CooMBE 1960, NITscH et ai. 1960, HALE 1968, lwAHORI et al. 1968, ALI.EWELDT and HIFNY 1972, HAtE and CooMBE 1974, ALLEWELDT et ai. 1975, lNABA et ai. 1976). Berry growth pattern follows a double-sigmoid curve in which four phases were identified by NITscH et ai. (1960). This pattern may be divided in two separate sigmoid curves: the first one is associated with strong mitotic cell activity and development of berry seeds, and is assumed to be under growth promoter control, while, during the second part, cell extension takes place and ripening processes occur; in this phase abscisic acid and small amounts of ethylene were detected in grape berries. Our knowledge about gibberellin-like substances in grape berries is not ex-haustive, but the results of CooMBE (1960) and lNABA et ai. 0976) indicate that a maximum content is reached during phases I and II of berry growth. In addition, Irn et al. (1969) and FARMAHAN and PANDEY (1976) found higher levels of gibberellin-like substances in seeded than in seedless berries. 1) Research performed under C.N.R. programme "Biology of Reproduction". 'l Professor at Piacenza and Assistant at the University of Milano. ') Asslstant at the Institute of Pomology of Milano. 362 A. ScrnNzA, R. MIRAVALLE, C. V1sA1 and M. FRBGONJ Abscisic acid attains a high level at the beginning of phase IV, which cor-responds with the initiation of the ripening processes in the berries (DüRING 1974, HALB and CooMBE 1974, CooMBB 1976, L1wv and ANGBLOVA 1977). After ABA treat-ments HALB and CooMBB (1974) found an earlier maturation and a higher sugar content in the berries, while, phase II, ethrel application increased the ABA content with a corresponding hastening of maturation. Abscisic acid is supposed to be produced not only in the seeds (Lon 1968, ALLB-wllLoT et al. 1975, CooMBB 1976) but seems also to be transported from the leaves to the berries (DüRING et al. 1978). Material and methods In Mr. ERNBSTO V1cBvAN1's vineyard at Ancarano di Rivergaro in the province of Piacenza berries were picked every 8-10 days starting at fruit set until ripeness, following the technique described by HucuN and JmLLIARo (1959), using Cabernet Sauvignon vines on Kober 5 BB rootstocks. Berries were classified in the laboratory according to their seed number. Berries containing 1, 2 or 3 seeds were analyzed as follows: Determination of gib b e r e 11 in - l i k e s u b s t a n c e activity by barley en-dosperm bioassay. Extraction, purification and chromatography were carried out following EccHBR et al. (1976). Chromatograms were tested as described by CooMBE et al. (1967), modified by RYUco and INTRIERJ (1972). Free a b s c i s i c a c i d determination by high pressure liquid chromatography (HPLC). Extraction by meth!lnol and ether was carried out following the method of DüRING and ScrnNzA (1975), while the identificartion and quantitative determination was performed as follows, modifying the method of DüRING and BAcHMANN (1975): Colurnn: Lisorb AMM (25 X 0.21 cm); mobi:le phase: buffer phosphate 0.02 M, pH 5; flow rate: 60 ml/h at 2100 psi; column temperature: 25 °C; detector: UV 262 nm; inj ection: 5 ftl. Sugar determination was carried out by refractometer and a c i d i t y ex-pressed in tartaric acid titrated by 0.1 n NaOH, using centrifugated grape juice. Tartaric acid contents were determined by REBBLEJN's method (1961) modified by LIPKA and TANNER (1974), while malic acid was determined by the enzymatic method described by BERGMEYER (1970). The an t hoc y an in content was determined following the method of EwART and KuEWBR (1977). All results were elaborated according to analysis of variance and L.S.D. calcula-tion by TuKBY's test (1956). Results Gibberellin-like substances in berries The gibberellin-like activity of extracted berries showed two peaks in phases II and III of the growth curve. The second peak showed particularly high levels (up to 0,14-1,15 p.g/100 g fresh weight), according to seed number (Fig. 1). During the final part of phase III a decline of gibberellin-like substances was detected, while, during phase IV, gibberellin activity was very low. In phase III, Seed number, gibberellin, abscisic acid levels; ripening of berries 363 three-seeded berries contained higher levels of gibberellin-like substances than one or two-seeded berries. ~ 1,15 "' 1,10 ~ 1,05 "i 1,00 ~ 0,95 "' ijl 0,90 ~ 0,20 _, z 0,15 _, ~ 0,10 °' ~ 0,05 (3 0 ~ 40 .;; 35 "' 0 30 ~ -Cl 25 ::i. 0 u 20 <{ 15 u in u 10 fJ) a> <{ 5 0 1.60 "' 1,4 1-1,20 J: Q 1,00 w s: 1.80 > "" 1.60 "" w 1,40 "' 1.20 *•* 11 19 1 31 JUNE 45 J U L Y * 61 79 AUGUST I LSD p.0,05 94 days • te< 108 ant hflsis SEPTEMBER Fig. 1: Gibberellin-like substance and ABA contents (µg/100 g fresh w eight) in b erries w ith different number of seeds in r elation to the development phases of the berries. Gibberellin- und Abscisinsäuregehalt (l•g/100 g Frischgewicht) von Beeren m it unter-schiedlicher Anzahl von Samen in Beziehung zu den Entwicklungsphasen der Beeren. 364 A. ScrnNzA, R. M1RAvALLE, C. V1sA1 and M. FREGONI 29 • 27 ß 25 23 il: 21 19 17 15 13 11 .... 1 phase n 1 phase nr 1 11 19 31 45 1 UNE J u l y phase JV 61 79 AUGUST t ACIDITY %o *~!~: 94 SEPTEMBER 108 days alter anthesis Fig. 2: Behaviour of sugar content and acids in the berries in relation to the number of seeds. Z~cker- und Säureverlauf in den Beeren in Beziehung zur Anzahl der Samen. Abscisic acid content in berries The highest levels of abscisic acid in berries were detected in the period im-mediately after fruit set (40,7 pg/100 g fresh weight) (Fig. 1). A decline of the abscisic acid level was noticed during phases II and III of the growth curve with a content of 3-7,6 ,ug/100 g fresh weight depending on seed number. In phase IV, the abscisic acid concentrations raised up to 26-32 pg/100 g fresh weight, and then declined to low values. While there were no significant differences in abscisic acid values to a fresh-weight basis detectable between the one, two and three-seeded berries except one date (31st d after anthesis), abscisic acid on a per-berry basis shows higher amounts in three-seeded than in two or one-seeded berries (Fig. 4). Abscisic acid content in rachis Abscisic acid levels in rachis developed independently from those of the berries; i.e. ABA concentration increased greatly during the first phases of rachis develop-ment. The level kept constant in the period of berry colour change and a second rise was observed during the final phase of ripening. Seed number, gibberellin, abscisic acid levels; ripening of berries 365 Sugar content, titrable acidity, tartaric and malic acid content There is a small (but only at the 61st d after anthesis) significant difference in the increase of sugar percentage between the one, two and three-seeded berries showing a more rapid sugar increase in one and two-seeded berries (Fig. 2). The increase of sugars on a per-berry basis shows that this increase is more rapid in three-seeded than in two or one-seeded berries (Fig. 4). The number of seeds per berry had no significant influence on the course of concentrations of titrable acids (Fig. 2). Tartaric acid content decreased steadily from fruit set to ripeness, being always higher in one-seeded berries, but these differences are not significant. Malle acid increased and subsequently decreased with progressing maturity. The malic acid maximum of three-seeded berries is reached 16 d later than that of one or two-seeded berries. Anthocyanin content The appearance of anthocyanins was associated with berry colour change; they accumulate in the skin progressively during the ripening process, especially in August. One-seeded berries had a statistically higher amount of anthocyanin than two or three-seeded berries, in particular during the final phase of ripening in Septem-ber (Fig. 3). 1 u '-.. O> E z z ~ u 0 J: ,_ z <( 0.20 0 .18 0 .16 0 .14 0 .12 0 .10 O.Ö8 0.06 0.04 ..... 0.02 61 79 94 AUGUST I LSD p• 0,05 S EPTEMBER 108 days alter anthesis Fig. 3: Influence of seed number on the anthocyanins (mg/cm2) of the skins. Einfluß der Samenzahl auf den Anthocyangehalt der Beerenhaut (mg/cm2). 366 190 170 150 VI ... °' "' 130 ... CD § 110 ' "' ::1. 0 90 ü < 70 u in ü VI 50 CD < 30 10 260 220 VI ... °' "' ... 180 CD 8 Z.140 "' "' 100 < C> :;) VI 60 20 A. ScrnNzA, R. MrnAvALLE, C. V1sA1 and M. FREGONI * ·~ • ."" ·• \. * ~ ••• „d. :~/„.~ ~·1 •• „c1 ................ * • ·--~ *--=:: ••• * ·-· 3 „.c1. ~·-•2mdo */ „ /.--.„~ dt/. .....-::•=-• -:::t~ ~·""'Ph.ii hase III 1 phase rv 11 19 IUNE 31 45 1 61 79 IULY AUGUST 94 108 SEPTEMBER days after anthesis Fig. 4: lnfluence of seed number on ABA content (µg/1000 berries) and sugar content (g/1000 berries) in berries. Einfluß der Samenzahl auf den Abscisinsäuregehalt (µg/1000 Beeren) und den Zuckerge-halt (g/1000 Beeren) von Traubenbeeren. Discussion From the above mentioned results it can be concluded that grape-berry develop-ment and changes in their chemical composition are eorrelated to their number of seeds. The content of gibberellin-like substances was at its highest at the end of phase III. Especially in this period the results show a positive correlation between the content of gibberellin-like substances and the number of seeds, while no signifi-cant correlation was registered between the abscisic acid content of berries (to a Seed number, gibberellin, abscisic acid levels; ripening of berries 367 fresh weight basis) and the number of seeds. Therefore, the gibberellin : abscisic acid ratio is higher in the more developed, i.e. three-seeded berries. This observation agrees in some respect with the results of CoNSIDJNI! and CooMBI! (1972) who found that GA3 application was inefficient to stimulate the berry growth of two to three-seeded berries, while application of CCC reduced berry size. The course of the content of abscisic acid in the berries during their develop-ment (on a per-berry basis) agrees well with earlier results of CooMBI! and HALi! (1973), DüRING et al. (1978) and INABA et al. (1976) who also found high amounts of abscisic acid in berries after anthesis and a second increase which was correlated to the inception of the ripening processes, e.g. sugar increase. In our results the content of abscisic acid as well as sugars increased more rapidly in three-seeded berries than in two or one-seeded berries. Summary In grapes of Cabernet Sauvignon, berry growth, content of anthocyanin and hormonal composition affected by seed number were investigated. Gibberellin-like substance content showed highest values 45 d after anthesis, while abscisic acid content (on a per-berry basis) had two peaks, the first one just after berry set, the second coinciding with ripening phenomenon. Seed number was positively correlated with the cöncentrations of gibberellin-like substances and abscisic acid (on a per-berry basis). One-seeded berries, how-ever, had a higher amount of anthocyanin than two or three-seeded ones. 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Relationships between seed number, gibberellin and abscisic acid levels and ripening in Cabernet Sauvignon grape berries

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Relationships between seed number, gibberellin and abscisic acid levels and ripening in Cabernet Sauvignon grape berries

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