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Flower Induction and Flower Bud Development in Apple and Sweet
Flower Induction and Flower Bud Development in Apple and Sweet
To cite this article: Nikolaos Koutinas, Gueno Pepelyankov & Valentin Lichev (2010) Flower
Induction and Flower Bud Development in Apple and Sweet Cherry, Biotechnology &
Biotechnological Equipment, 24:1, 1549-1558, DOI: 10.2478/V10133-010-0003-9
ABSTRACT
As a result of studies conducted in the last 20-30 years, new information has been obtained about the flower bud formation in
apple and cherry, including some data about the presence of genes determining the reproductive organs of the apple. Some
issues about the flower induction, histological transformation of the apical meristem, morphological differentiation of the flower
buds, factors and conditions influencing the flower bud formation and the quality of the reproductive organs are discussed in this
survey. Some attention has been paid to the tree pruning, summer pruning in particular, the fertilization, irrigation and treatment
with growth regulators, by means of which processes of flower bud formation can be regulated.
Keywords: apple, cherry, flower induction, histological flowers in given genotypes can be accelerated by application of
transformation, morphological differentiation, agricultural different methods. It seems that the most effective is stimulation
practices of the vegetative growth, so that the seedlings can reach a large
size as soon as possible. The onset of the flower bud formation
Biotechnol. & Biotechnol. Eq. 2010, 24(1), 1549-1558
in the seedlings of the apple can be accelerated by means of
grafting on the dwarfing rootstock M 9; the growing conditions
Introduction are of decisive importance, however (116). The more vigorous
Numerous studies on flower bud formation of temperate fruit the growth, the smaller the difference is, in flowering between
plants have been conducted in many countries (16, 17, 64, 67).
the grafted seedlings, and those grown on their own roots. The
In spite of that, a great part of the most important characteristics
transition from the juvenile into that of maturity phase may
of flower bud formation remains insufficiently clarified both
be due to various mechanisms, including the hormonal control
by theoretical biology and applied horticulture (16). Special
over the distribution of assimilates in the zone of the apical
attention deserves the clarification of the morphogenetic
meristems (52).
changes that occur between the vegetative state of the buds
and the initiation of flower organs (39). In order to reduce the duration of the unproductive period
in the grafted young trees grafting on dwarfing or semi-
The presence of relative dormancy between the initiation
dwarfing rootstocks is applied on a mass scale in apple and pear
of the flowers and anthesis is characteristic of most fruit plants
(64). In apple, for example, the cycle of flower development growing in particularly. However, the weakening of growth is
often lasts from 9 to 10 months, but only a short lapse of not always desirable, particularly in the first one and second
time is due to an active passing of this process, namely at the year after planting (25). In this respect, the characteristic of
appearance and formation of the floral primordia in summer the cultivars to form feathers and the conditions created for a
and at the final formation of flower parts in spring (67). The vigorous growth, are of special importance. As considerably
period of relative dormancy permits the buds to pass a complete more fruit buds are initiated for the third year in pear cultivars
cycle of their development in two successive calendar years. that form more feathers in the first than in second year, the
This provides an opportunity of decreasing the competition authors considered that the best prediction for the duration
for nutrients between the developing flowers and vegetative of the unproductive period can be done on the basis of the
parts, thus presenting a great advantage for the perennial fruit difference in the number of feathers per tree in the first two
tree species, including apple and sweet cherry, in comparison years. The formation of feathers is usually described as a type
with annual plants (64). The ability for a change in the of branching due to the decreased apical dominance (22).
predetermination of the buds, in the case when conditions for The flower bud formation in the mature trees is
the transition from the vegetative into the reproductive state fundamentally determined by the presence of hormones (16).
become unfavorable, may be considered another advantage. The lack of initiation of flower buds in the cultivars with
According to Zimmerman (132), duration of the juvenile alternate bearing, was related in the past directly to the effect
period, characteristic of the tree plants propagated by seeds, of exhausting assimilates and the reserve substances by the
is genetically controlled. The time of appearance of the first fruits. Now it has been attributed to the action of hormones
Biotechnol. & Biotechnol. Eq. 24/2010/1 1549
and particularly to gibberellins, produced in the young fruitlets histological transformation and morphological differentiation
in the so called ‘on year’ (80). (17).
Flower bud induction
Sites of flower bud formation
According to Dolega et al. (32), induction gives an impulse to
Most of the commercially significant apple cultivars bear fruit
the sufficiently developed and ‘susceptible’ buds to transition
mainly on spurs (37). The main leaf area of the trees from the
from vegetative to generative phase. It is still not clear which
‘green apex’ phase to the complete bloom is formed on these
are the fundamental reasons for induction (16, 76). It could be
spurs. Some of the leaves also develop on them later; the total
spur leaf area is formed up to one month after flowering (38). said that this is a qualitative change, the final result of which
By and large, during this period the preliminary induction of can be programming of the strategically disposed parts from
the flower bud formation on the spurs occurs (17). Leaves on the meristem to forming flowers (17). The induction can also
the spurs in the spur-type apple cultivars constitute more than be viewed as a process during which previously repressed
60% of the total leaf area of the tree; hence they are essential information is being transformed to form a new structure,
for the fruit-bearing (104). In the spur cultivars of ‘Delicious’ namely the flower bud. The genes of the type of AFL (Apple
the leaf number and leaf area per spur are larger in the spur Floricaula/Leafy) are supposed to be included in the induction
type strains of ‘Delicious’ than in the standard strains (119). (68). According to Luckwill (78) the induction is connected
The presence of a sufficiently large leaf area per spur in pear with changes in the hormonal balance, and according to Sachs
is a significant factor favoring flower bud initiation (63). In (106) with changes in the distribution of the assimilates in the
apple, every vegetative spur needs 100 to 150 cm2 leaf area apical meristem. Link (76) shares the point of view that the
in the last 90 days of the growing season in order to initiate a flower bud induction, as well as the fruit set and the fall of fruits
‘vigorous’ flower bud. Therefore, proper agricultural measures or of primordia of different organs of the plant, depend on the
are necessary to ensure development of such area (104). At a interaction in space and time of its own growth substances and
young age and under special conditions, trees of some apple those retaining growth. Together with the well-known method
cultivars initiate flower buds on the tops of the long shoots and of defoliation for determining the critical period of the flower
laterally in the axils of their leaves (37). The flower buds in bud induction in apple, it is possible to use spraying by GA3
apple, apical or lateral, are mixed and they contain primordia within the supposed time of the induction (75).
of vegetative and reproductive organs. Flower buds of sweet Histological transformation
cherry are initiated laterally, in fascicles and are unmixed When the apex of the still vegetative bud receives a signal
(simple) (124). They are disposed laterally on short spurs or for differentiating as a flower bud, a sequence of events takes
near to the base of longer shoots. place (17). The mitotic activity becomes total for the whole
apex changing its histological structure. The central meristem
Development of the flower buds is more unfolded and is now under the subdermatogen. By this
Flower bud formation is a complex phenomenon. Considering rearranging, but without any morphological changes in the
the flower bud formation genes to be a basic factor in its apex, the histological transformation is carried out, and from
occurrence, and formation of gametes in the time of meiosis that moment on the initiation of the flowers is irreversible.
as a final event, Wellensiek (121) formulated the statement that
the initial reasons for formation of flowers can be defined as Morphological differentiation
‘something happening between the flower forming genes and Apple is a fruit-tree species of the temperate climatic zone, on
the meiosis’. This statement is still valid (16). For the flower which the greatest number of studies is conducted, referring to
initiation it is necessary to stop the retaining of the genes the initiation of flower buds. The studies on the sweet cherry
responsible for the flower bud formation and in this sense the are limited.
induction of flower bud formation can mean something similar The apple flower bud is a shortened axis, usually carrying
to de-blocking. 21 formations, inserted in the spiral sequence (1). It consists
The identification and characterization of a number of of bud scales, transitional leaves, true leaves and bracts (a leaf
homeotic genes connected with the flower bud formation in form, lamina and stipules of which are totally depressed). The
some model angiosperm species, such as Antirrhinum majus L. axis ends in primordium of the ‘king flower’ and the primordia
and Arabidopsis thaliana (L) Heynh., have allowed, in the last of the lateral flowers are formed in the axils of the three bracts
ten-year period, to seek their homologues in the fruit species, and the three upper leaves. Under the whorl of the flowers in
and more specific in the apple, which is the most important the axils of one or two of leaves, a primordium of a vegetative
representative of the species from the temperate climatic zone bud is located. Out of this bud, during the vegetation, a bourse
(39). Such genes are isolated from flower buds in the still initial shoot is formed which can, in its turn, form a new flower bud.
phases of their development, from flower organs and fruits of The morphological differentiation occurs only if the
some apple cultivars (68, 110, 111, 118, 129). structure of a vegetative bud is complete (16). The differentiation
The flower bud formation in the apple passes in the of flower primordia in apple can start when the primordia of
following succession: induction of flower bud formation, the appendages described above have already been initiated.
1550 Biotechnol. & Biotechnol. Eq. 24/2010/1
The total appendages formed before the appearance of the of the bottom of the cave sinks and five swellings of the carpels
first indication for the transition of the buds from vegetative appear. The lateral flowers are formed like the apical one.
into reproductive state represents a ‘critical number of nodes’. The appearance of the primordia of the separate flowers
This number is varying depending on cultivar, from 16 to 20 in sweet cherry, within the framework of the same bud, is
(55, 57, 62, 80). The total number of appendages formed by nearly simultaneous (131). Certain differences in the phases of
the appearance of the sepals of the apical flower in the cluster development of flowers in a given bud of sour cherry can be
is also specific for the cultivars and can be influenced by the observed up to the initiation of the anthers and carpels; after
ecological conditions and the ways of growing (83). that, all flowers look similar (30).
For the formation of bracts and the initiation of flowers With the onset of the morphological differentiation in
in their bases, it is necessary for the plastochrone (the time the month of July (6, 31, 66, 67, 86) carpel primordia in the
between the forming of the primordia of two neighboring apple appear in September to October. The further formation
organs) to be shortened to 7 days (42). In a series of publications of flower parts occurs till the transition of plants into a state
(40, 41, 42) raised an idea of a system of development based on of winter dormancy (67). Toward the end of autumn all
the inhibition of given organs by the neighboring ones. flower primordia, except those of the sepals, are found in an
The widening and slight swelling of the bud apices, undifferentiated state. The carpels usually form seed cases but
when they become dome-like in shape, is usually accepted are not yet present in form of a pistil.
as the first visible indication of the transition from vegetative The microsporogenesis and the macrosporogenesis in the
to reproductive state (1, 50, 57, 61, 62, 67, 81, 130). For a apple occur in the following calendar year (71, 89). From the
greater precision in determining the moment of transition to a initiation of the anthers in the autumn till the full formation
reproductive state in the apple, the onset of the bract initiation of pollen in spring more than half a year passes under the
could be taken into consideration (42, 70). However, when condition of Moldova (71).
observed under a stereoscopic microscope, the bracts are not The rate of development of flower buds during the
visible at the moment of their appearance but a little later. It different phases is not constant, particularly under unfavorable
would be better to use an electronic microscope enabling the meteorological conditions (62). The development can be
observation of the separate parts of the developing flower buds retarded for a certain period of time. Consequently, it is difficult
at their mere initiation, as it can be concluded from the results to establish how long a given phase lasts.
of some studies on apple and sweet cherry (1, 13, 14, 30, 39,
In sweet cherry the rate of development of buds is
50). In connection with the clarification of the morphogenetic
accelerated. The pistil primordia can be observed at the
events between the vegetative development and the onset of the
beginning of September under the conditions of Bulgaria
floral organ initiation, Foster et al. (39) suggested the concept,
(45) and toward the end of the same month in Germany
according to which the widening of the bud apex preceding
(32) or Switzerland (14). Under identical conditions of the
the doming could be accepted as the first visible sign of the
environment for growing, pistil stigmas can be observed in
transition to floral development in the apple.
December in sweet cherry and in March of the following year
There is a certain order in the onset and the rate of in apple (131).
morphological differentiation in some fruit tree species, which
can be put in the following order of succession: sweet cherry, Factors and conditions for flower bud formation
apple, pear, peach, quince (131). Our studies (unpublished
The development of flower buds, from induction to the
data) reveal the same order as far as sweet cherry and apple
anthesis, is subjected to the influence of different factors and
are concerned.
conditions, which at a certain moment are responsible for
According to Kolomiets (67), Bergh (13), Huang (62) and flower bud formation.
Foster et al. (39) the morphological differentiation in apple is
expressed in the following order. After induction of the flower Cultivar
bud formation and the histological transformation of the apical The onset of the histological (15) and morphological (20,
meristem, laterally to the growing apex a swelling is formed 61, 66, 95, 130) differentiation in apple may depend on the
and this is a primordium of a bract. In the axil of the bract a characteristics of a cultivar. The cultivars can exert an influence
lateral flower is initiated. Simultaneously the primordium of also with respect to the further development of flower buds
the following bract is formed and in its axil the next lateral (89).
flower is formed, etc. It is followed by the primordium of the
Rootstock
apical flower, which overtakes in its development the lateral
flowers. The five sepals on the apical flower do not appear Rootstocks, as components of the grafted fruit trees, may
simultaneously and at the beginning they are not of the same influence the time of initiation of flower buds. According to
size. Later on, on the inner side of the sepals, at the point of Buban (15) some apple rootstocks exert this influence only on
their juncture, petal primordia appear. After that, the stamen specific cultivars. The same was noted by Elek (33) in sour
primordia are formed, in shape of a ring, under the petals, with cherry. Apparently an interaction between the two components
a second ring of stamens below the first one. Finally the center of a tree takes place, leading to a change in the time of onset