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POLLINATION IN JAPANESE PLUM

Article  in  Acta horticulturae · September 2010

DOI: 10.17660/ActaHortic.2010.874.28

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Pollination in Japanese Plum
V. Nencetti, E. Giordani and E. Bellini S. Radice
Horticulture Department Centro de Estudios Farmacológicos y
University of Florence Botánicos
Viale delle Idee, 30 CEFJBO-CONICET-UBA
Sesto Florentino Buenos Aires
Florence Argentina
Italy

Keywords: Prunus salicina, anthesis, pollen germinability, self-incompatibility,

pollinating insects

Abstract
Efficient pollination is fundamental for guaranteeing adequate productivity
in Japanese plum. Total gametophytic self-incompatibility distinguishes all cultivars
of this pomological group and makes it necessary to plant suitable pollinizers able to
guarantee an adequate production of fertile pollen during anthesis. Based on studies
carried out over more than a decade at the Horticulture Department, University of
Florence (DOFI), observations of numerous cultivars and genotypes obtained from
crosses have revealed the main characteristics required for a good pollinizer include
elevated production of flowers, long anthesis period which is stable from year to
year, scalar flowering, abundant pollen production, high viability and fertility of
pollen grains corresponding to high germinability. It is also important that the good
pollinizers show interfertility with the majority of widespread cultivars and that
their flowers are very attractive to pollinating insects.

INTRODUCTION
Productivity of Japanese plum depends on both the fertility of an adequate number
of flowers on a given tree and the successful fertilization of these with pollen from other
compatible plum genotypes. All Japanese plum cultivars commonly planted in Italy are
completely self-incompatible (factorial sterility) (Bellini and Bini, 1978; Arora and Singh,
1990; Palara et al., 1990; Bellini et al., 1996). As is true for all drupes, Japanese plum is
not able to produce mature parthenocarpic or apomictic fruits.
Considering the above, it is clear that plantings must include suitable pollinizers or
consist of compatibly interfertile cultivars (Costa and Grandi, 1982; Richards et al.,
1992). Often, however, productivity of Japanese plum orchards is low, which is generally
blamed on the inefficiency of inadequate pollinizers. This inefficiency can be caused by
numerous factors, most importantly the inter-incompatibility of genotypes used. There are
other causes which can be responsible for decreasing or negating the efficiency of
pollinizers (Bellini et al., 1995).
Among these other causes, the role of the pollinating insects which transport
pollen from the flowers of the pollinizer to those of the cultivar is very important
(Delaplane, 2000). The pollination of plum, being insect-mediated, can fail if factors such
as weather are not favorable (Loreti and Pisani, 1991). The presence of bee hives during
bloom has not resolved the problem, since with respect to other pollinators such as
bumblebees, honey bees are less active in the unfavorable weather conditions typical of
late winter which can coincide with the bloom period of Japanese plum (Delaplane,
2000). The broad genetic variability in this pomological group explains the different
attractions of various genotypes to pollinators. Dimensions, color and arrangment of floral
organs, and the amount of nectar available could be the cause of this phenomenon (Radice
et al., 2008 a). It is certain that the preferences of various species of insects do have an
important role in the success of pollination.
Another important reason for unsuccessful pollination can be the lack of overlap
in bloom between pollinizer and the cultivar to be pollinated. Genotypes which are

Proc. 9th IS on Plum & Prune Genetics, 203

Breeding & Pomology
Ed.: F. Sottile
Acta Hort. 874, ISHS 2010
described as having simultaneous bloom may not actually overlap bloom periods
depending on the environment and on weather conditions in a given year. The complex
mechanisms of chilling requirements (Malgarejo, 1996) and subsequent heat unit
accumulation, which are not well studied in Japanese plum, may affect flowering date and
duration of anthesis differently in different cultivars. In southern Italy, this shift of
flowering time is greater than in northern Italy where flowering is generally more
synchronized (Nencetti et al., 2006). This can be explained by the fact that the north
experiences more severe winters, satisfying cold requirements for all cultivars and leaving
them ready to accumulate heat units in order to initiate bloom. In many areas of the south
of Italy, the flowering period of the different Japanese plum cultivars is wider and can
cause lack of overlap in bloom between cultivar and pollinizer with severe reduction of
productivity. Thus, a good pollinizer for the north may be considered inadequate in many
southern areas.
The ability to prolong anthesis with a continuous cascade of opening flowers is
another characteristic of a good pollinizer. This feature of a pollinizer can reduce or
eliminate the problem of lack of overlap.
Another fundamental characteristic of a good pollinizer is the high level of
flowering every year (potential fertility). Additionally the pollinizers must produce viable
and fertile pollen, with high germinability of pollen grains.
A breeding program for Japanese plum was initiated in 1989 at DOFI with the
objective of obtaining early ripening cultivars with good fruit characteristics suitable to
climatic conditions in nothern Italy.
In this paper the data from the DO-FI breeding program are used to elucidate the
complex mechanisms of pollination in Japanese plum.

MATERIALS AND METHODS

Trees of 55 cultivars and 14 selections were grown in Emilia Romagna at Vignola
(MO). Three trees per genotype were studied and trained to delayed open-centre trees
planted at 4 x 3.5 m spacing. Trees were grafted on Myrobalan B. Cultivars were
observed and evaluated over four fruiting years. In addition to evaluation of fruit
characteristics, other observations concerned the timing of developmental stages and the
general characterisitics of the tree (Table 1), including ripening date, vigor, growth habit,
bloom date and productivity. Some of the cultivars of the collection were used for
crosses. The crossings were performed over several years with many combinations of
cultivars. The level of interfertility for each crossing was evaluated (Table 2). Data was
discarded from years when late frost during bloom damaged the crosses.
Progeny from the first crosses were evaluated in an adjacent field while progeny
from the later crosses were evaluated in Tuscany at Azienda Montepaldi of the University
of Florence at San Casciano Val di Pesa (FI). These progeny were evaluated in the same
manner as the cultivars in the collection. Particularly, with regard to bloom, the following
parameters were evaluated for all genotypes: sites of floral differentiation, flower
abundance, duration, date of beginning and end of anthesis.
Among the seedlings grown at Azienda di Montepaldi, 10 eight-year-old seedlings
(S1, S2, S3, S4, S5, S7, S8, S9, S10, S12) were chosen to study floral biology (Padula et
al., 2004; Ontivero et al., 2005, 2006; Radice et al., 2008b).
More recently, additional work was carried out on another group of seedlings
grown at Azienda di Montepaldi, to study the attractiveness of the flowers of different
genotypes to insect pollenators. Overall, observations were taken of about 900 seedlings
(Radice et al., 2008a).

RESULTS AND DISCUSSION

Data gathered during four years of observation on the cultivars in the Vignola
collection show that 15 of the cultivars present had very low average yields, 12 had low
average yields, 10 medium average yields, and 18 had high average yields (Table 1). A
positive correlation between yield and open growth habit of the tree was observed

204
(P<0.01) (Table 1).
Studying the progeny obtained allowed the idenitification of crosses with the
greatest interfertility (Table 2). Some combinations had no yield (‘Andys Pride ×
Burmosa’, ‘Andys Pride × Santa Rosa’, ‘Angeleno × Blackamber’, ‘’Black Star × Santa
Rosa’, ‘Blackamber × Angeleno’, ‘Burmosa × Laroda’, ‘Red Beaut × Burmosa’, ‘Santa
Rosa × Burmosa’). Some of the reciprocal crosses were interfertile (for example
‘Angeleno’ and ‘Santa Rosa’) while others were not (for example ‘Angeleno’ and
‘Blackamber’), and some were partially interfertile (for example ‘Burmosa’ and ‘Laroda’)
(Table 3).
Some seedling progeny of ‘T.C Sun × Shiro’ and ‘Black Diamond × Morettini
355’ had the ability to interrupt early bloom (10−15% of flowers open) due to sudden
lowering of minimum temperatures (-2°C); blooming recommenced after 5−6 days when
temperatures rose again.
Studies of floral biology carried out on the 10 seedlings showed large variability in
the parameters analyzed. From microsporogenesis studies, it was observed that some (S2)
were earlier than others. During microsporogenesis, alterations in the process are possible
such as the dynamic degradation of the carpel. Such alterations compromised the quality
of pollen (S3, S4) (Ontivero et al., 2005). Germinability of pollen grains was also
different, ranging from 0% to intermediate (30%), to high (90%). Following pollination,
differences in pollen development were observed. With artificial self-pollination, there
was no development of the pollen grain. In some cases, both using pollen from other
genotypes and with free pollination, the pollen tube grew normally along the style
(Ontivero et al., 2006).
Studies of floral biology have shown four possible anther forms present in the
various genotypes (white, yellow-orange, brown, purple) which produce a greatly variable
quantity of pollen. There were no significant differences in the presence of nectar in the
flowers except at the end of bloom. The same was true for the scent intensity, though the
type of fragrance (grassy, floral or sweet) varied. Diptera and hymenoptera visited the
flowers during the whole period of bloom. Bees were predominant compared to
bumblebees and other insects. Fruit set was very low and in some progeny more than 60%
of the trees did not produce fruits.

CONCLUSIONS
The very low yield of some cultivars studied within the collection and of the other
genotypes studied, despite the availability of many pollen types during bloom, suggests
that pollination is necessary but not sufficient to guarantee good yield. Some cultivars can
not express their yield potential because of adverse climatic conditions or intrinsic genetic
problems.
Results from the crossings indicate that interfertility is specific to individual
combinations of cultivars or genotypes: a pollinizer that is good for one cultivar may not
be suitable for others. Another important problem is the overlapping of bloom, which is
difficult to study and control because it is strongly affected by climatic conditions which
change from year to year. For both of the above reasons, the use of several pollenizer
genotypes is recommended. In addition to the preceding requirements, a good pollinizer
must have abundant flowers, continuous flowering opening, and a prolonged bloom
period. Additionally, pollen should be plentiful, vital, and with high germinabilty. Studies
of floral biology indiate that anther color, the abundance of pollen, the presence of nectar
up to the end of bloom, and the fragrance of the scent released can affect the
attractiveness of the flowers to insects: more insects visit certain progeny.

Literature Cited
Arora, R.L. and Singh, R. 1990. Genetics of incompatibility in plum (Prunus salicina
Lindl.). Ind. J. Hort. 47:1−11.
Bellini, E. and Bini, G. 1978. La fertilità nel susino. Riv. Ortoflorofrutticoltura Italiana.
p.403−422.

205
Bellini, E., Nencetti, V. and Sabbatini, I. 1995. La fertilità del susino cino-giapponese.
L’Informatore Agrario 22:57−64.
Bellini, E., Nencetti, V., Giordani, E., Sabbatini, I. and Caruso, S. 1996. Osservazioni sul
comportamento di nuove cultivar di susino. L’Informatore Agrario 27:53−60.
Costa, G. and Grandi, M. 1982. Contributo alla conoscenza delle esigenze di
impollinazione di nuove cultivar di susino. Atti Incontro frutticolo SOI “La coltura del
susino” Ferrara, 19 febbraio. Rivista di Frutticoltura e di ortofloricoltura 12:74−76
Delaplane, K.S. 2000. Crop Pollination by Bees. Cambridge, MA, USA: CABI
Publishing p.344.
Loreti, F. and Pisani, P.L. 1991. L’impollinazione nel susino. Frutticoltura, 53(4):21−26.
Melgarejo, P.M. 1996. El frìo invernal, factor limitante para el cultivo frutal. Modelos y
métodos para determinar la acumulaciòn de frìo y de calor en frutales. A. Madrid
Vicente, Ediciones 127:166.
Nencetti, V., Bellini, E., Natarelli, L., Pirazzini, P.and Insero, O. 2006. Fruttiferi 2006
Liste varietali: Susino. Supplemento L'Informatore Agrario 23:63−66.
Ontivero, M., Radice, S., Giordani, E. and Bellini, E. 2005. Preliminary studies on
microsporogenesis in Prunus salicina Lindl. Int. J. Hort. Sci. Biotech. 80:599−604.
Ontivero, M.R., Radice, S., Giordani, E. and Bellini, E. 2006. Effects of different
pollination treatments in genotypes of Purnus salicina Lindl. Intl. J. Hort. Sci.
12:141−146.
Palara U., Passerini V. and Stecchetti B. 1990. Biologia fiorale e caratterizzazione di
alcune cultivar americane di susino cino-giapponese della serie “Black”. Rivista di
Frutticoltura e di rtofloricoltura 52(6):39−43.
Padula, G., Bellini, E, Radice, S. Giordani, E. and Nencetti, V. 2004. Fertilità in susino
cino-giapponese (Prunus salicina Lindl.): prime indagini morfologiche e fisiologiche
su polline. Italus Hortus p.52−53.
Radice, S., Giordani, E., Nencetti, V. and Bellini, E. 2008a. Phenological expression in
Prunus salicina Lindl. Genotypes and its relation with insect attraction and
pollination. IX ISHS Int. Symp. Plum and Prune Genetics, Breeding and Pomology.
Palermo.
Radice, S., Ontivero, M., Giordani, E. and Bellini, E. 2008b. Anatomical differences on
development of fertile and sterile pollen grains of Prunus salicina Lindl. Plant Syst.
Evol. (in press).
Richards, G.D., Porter, G.W., Rodriguez, A.J. and Sherman, W.B. 1992. Pollen
production and cross compatibility in low-chill Japanese-type plum. Proc. Fla. State
Hort. Soc. 105:302−304.

206
Tables

Table 1. Main characteristics of Japanese plum cultivars present in the Vignola (MO)
field collection.

Cultivar Ripening Vigor Growth habit Bloom period Productivity

date
Au-Amber -30 h intermediate me l
Red Beaut® -30 h upright me vl
Early Wilson -29 mh intermediate e h
Durado® -27 m intermediate me l
Burmosa -22 mh open e vl
Obilnaja -19 h open me h
Morettini 355 -17 h open m h
Royal Garnet® -14 ml upright e vl
Black Beaut® -8 m intermediate m vl
Robusto -8 h open m l
Robusto Segundo -7 h open m l
Au-Roadside -7 h intermediate e l
Au-Rubrum -7 h intermediate m m
Gulf Ruby -5 m intermediate e vl
July Sun® -4 m intermediate me l
Ozark Premier -4 m open me h
Au Cherr y -3 h intermediate me m
Del Rey Sun® -2 m intermediate m h
Dolly® -2 ml intermediate m vl
Shiro 0 h open m h
Sangue di Drago 0 m open me h
Blackamber 0 ml upright me vl
Black Star® 0 m upright m m
Santa Rosa +1 m upright m vl
Byrongold +1 m upright m vl
Beaut Sun® +5 m upright me vl
Black Gold® +10 m intermediate m h
Yellow Sun® +15 m upright m vl
Black Diamond® +18 mh upright m h
Catalina +20 m upright e l
June Beaut +20 mh upright ml l
Laroda +21 mh upright m h
11 P 600® +25 m upright l vl
Green Sun® +27 mh intermediate m h
Fortune +28 ml upright m m
Zanzi Sun® +29 l upright m l
Sterling +30 mh upright l l
Salad +30 mh upright m l
Original Sun® +33 mh intermediate m m
Midnight Sun® +33 m upright m m
Bella di Barbiano +35 m intermediate me m
Tardiva di Scanzano +35 m upright m vl
Tracy Sun® +35 l upright me l
Globe Sun® +37 m upright m vl
TC Sun® +37 mh intermediate m h
Eric Sun® +41 m intermediate m vl
Howard Sun® +50 l intermediate m m
Larry Ann® +50 h upright me h
Royal Diamond® +53 ml intermediate l h
Ruby Blood +55 m intermediate m m
Januaria +55 ml open m m
Centenaria +55 ml intermediate m h
Autumn Giant® +60 m intermediate m h
Angeleno® +60 h upright m h
October Sun +61 m intermediate l h
Legend: vigor and productivity (vl=very low, l=low, ml=medium-low, m=medium, mh=medium-high,
h=high); bloom period (e=early, me=medium-early, m=medium, ml=medium-late, l=late); ripening
date ± days relative to ‘Shiro’, which matures in mid-July in central Italy.

207
Table 2. Degree of interfertility between Japanese plum cultivars.

Interfertility Crosses
High “Laroda x Burmosa”, “Santa Rosa x Angeleno”, “Laroda x Ozark Premier”,
“Black Gold x Burmosa”, “Black Diamond x Santa Rosa”, “Santa Rosa x
Black Star”, “TC Sun x Golden Plum”, “Santa Rosa x Laroda”, “Black
Diamond x Burmosa”, “Laroda x Queen Rosa”, “Laroda x Black Gold”

Medium “TC Sun x Byrongold”, “TC Sun x Shiro”, “Angeleno x Santa Rosa”,
“Angeleno x Burmosa”, “Black Diamond x Black Star”, “Angeleno x
Frontier”, “Burmosa x Santa Rosa”, “Black Star x Black Gold”, “Ozark
Premier x Burmosa”, “Black Diamond x Black Gold”, “Black Gold x Black
Diamond”, “Simka x Blackamber”, “Blackamber x Queen Rosa”, “Black
Gold x Laroda”
Low “Blackamber x Simka”, “Black Diamond x Morettini 355”, “Santa Rosa x
Red Beaut”, “Burmosa x Red Beaut”, “Blackamber x Santa Rosa”, “Black
Star x Friar”, “Burmosa x Ozark Premier”, “Burmosa x Andys Pride”,
“Laroda x Morettini 355”, “Blackamber x Burmosa”, “Black Star x Calita”,
“Black Star x Frontier”, “Blackamber x Red Beaut”
Absent “Andys Pride x Burmosa”, “Andys Pride x Santa Rosa”, “Angeleno x
Blackamber”, “Black Star x Santa Rosa”, “Blackamber x Angeleno”,
“Burmosa x Laroda”, “Red Beaut x Burmosa”, “Santa Rosa x Burmosa”

Table 3. Reciprocal interfertility in crosses of Japanese plum cultivars.

Degree of Reciprocal
Crosses
interfertility interfertility
“Angeleno x Santa Rosa” medium
good
“ Santa Rosa x Angeleno” high
“Black Gold x Laroda” medium
good
“Laroda x Black Gold” high
“Black Diamond x Black Gold” medium
good
“Black Gold x Black Diamond ” medium
“Blackamber x Simka” low
moderate
“Simka x Blackamber” medium
“Burmosa x Ozark Premier” low
moderate
“Ozark Premier x Burmosa” medium
“Burmosa x Laroda” absent
partial
“Laroda x Burmosa” high
“Black Star x Santa Rosa” absent
partial
“ Santa Rosa x Black Star” high
“Burmosa x Santa Rosa” medium
partial
“Santa Rosa x Burmosa” absent
“Burmosa x Red Beaut” low
partial
“Red Beaut x Burmosa” absent
“Andys Pride x Burmosa” absent
partial
“Burmosa x Andys Pride” low
“Angeleno x Blackamber” absent
absent
“Blackamber x Angeleno” absent

208
Figures

Fig. 1. ‘Santa Rosa’ and ‘Angeleno’, two inter-fertile cultivars in full blooming.

Fig. 2. Bee visiting Japanese plum flower.

209
Fig. 3. Anthers of different colours, bees prefer the range of colours yellowish-red.

Fig. 4. Japanese plum flowers with pink nectaries.

210
Fig. 5. Scalar and high production of flowers.

Fig. 6. Characters of the good pollinizer: elevated production of flowers, long anthesis
period, scalar flowering, abundant and fertile pollen production.

211
 Fig. 8. Fertile anthers observed using an electron microscope.

212

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