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Strawberry Growth and Flowering

Article  in  International Journal of Fruit Science · January 2006

DOI: 10.1300/J492v05n01_04

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Strawberry Growth and Flowering:

An Architectural Model
G. Savini
D. Neri
F. Zucconi
N. Sugiyama

ABSTRACT. Strawberry is an herbaceous perennial that can be easily

propagated by stolons or by crown division. Seedlings have a few
months in the juvenile period. The growth is determined, with a terminal
inflorescence as a modified stem or cyme. Flower induction is sensitive
to thermo-photoperiod and to several agronomic and nutritional factors.
The strawberry plant is highly adapted to diverse temperate climates and
to production under forced artificial growing conditions. In the present
study, a dynamic architectural model of growth is described and dis-
cussed in order to reveal strawberry development under different culti-
vation systems. The aim is to improve the cultural knowledge and
understanding of the physiological control of axillary meristems, which
can enhance the strawberry fruit production. [Article copies available for a
fee from The Haworth Document Delivery Service: 1-800-HAWORTH. E-mail ad-
dress: <docdelivery@[Link]> Website: <[Link]
com> © 2005 by The Haworth Press, Inc. All rights reserved.]

G. Savini, D. Neri and F. Zucconi are Researchers in the Energetics Department,

Marche Polytechnics University, Via Brecce Bianche, 60100–Ancona, Italy (E-mail:
neri@[Link]).
N. Sugiyama is a Researcher at the Graduate School of Agriculture, University of To-
kyo, Yayoi, Bunkyo-ku, Tokyo 113, Japan (E-mail: anobuo@[Link]).
The authors would like to thank Dr. Y. Lim for the valuable help in editing the
manuscript, and the “Molari & Gatti nursery” from Cesena and Università Politecnica
delle Marche for the funding of G. Savini’s fellowship.
International Journal of Fruit Science, Vol. 5(1) 2005
[Link]
© 2005 by The Haworth Press, Inc. All rights reserved.
Digital Object Identifier: 10.1300/J492v05n01_04 27
28 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

KEYWORDS. Junebearing strawberry, everbearing strawberry, spring-

bearing strawberry, stolon, crown, flower induction

THE PLANT

The strawberry plant is an herbaceous perennial rosette. The stem or

crown has a central pitch core and can survive for several years because
of secondary cambium activity. The internodes are very short and a
number of long-petiole trifoliate leaves at various stages of develop-
ment are arranged spirally on the axis (White, 1927; Guttridge, 1955).
The base of well-developed crowns is protected by the decaying sheath-
ing stipules of dead leaves. Within the stipules of the last emerged leaf,
is the enclosed bud, consisting of 5-6 developing leaves and a leaf
primordia.
The axillary meristems may develop into stolons or branch crowns,
with a regular positioning along the axis, and in strict relation with
crown growth rate. Differentiation between the two organs probably oc-
curs very early during the formation of the first two internodes and may
be very long or extremely short. If the axillary meristem ceases growing
then a latent bud originates.
Under favorable environmental conditions floral induction occurs at
the apex of the crown. Extension growth of the crown continues along
the axis of the uppermost lateral meristem below the terminal inflores-
cence (extension crown axis), thus giving a sympodial structure to the
crown which is not apparent at first sight (Guttridge, 1952; Arney,
1953; Robertson and Wood, 1954; Dana, 1969; Nicoll and Galletta,
1987; Aspuria and Fujime, 1995; Yanagi and Oda, 1989). The exten-
sion crown axis typically forms three or four nodes before its vegetative
growth is terminated by the initiation of a further inflorescence.
The Above-Ground Organs. The description presented here of the
plant organs allows for the development of an architectural model of the
annual plant cycle under cultivated conditions. The study includes the
analysis of the hierarchies among organs in the plant and their spatial
evolution in a dynamic perspective.
a. Crown is a primary stem, originated by a seed or a stolon, in which
the leaves, roots and stolons are cyclically formed. Crowns have a vari-
able number of internodes and terminate with an inflorescence. Before
the emergence of the terminal inflorescence the stem can be called a
shoot. The part of the shoot included in the stipules of the last unfolded
leaf can be considered as a terminal bud.
 Savini et al. 29

Initially the crown is a short body and may later, by apical growth,
become cylindrical and eventually fork into two or more divisions. The
subterranean portion is covered with roots. Above ground, it is clothed
with the leaves bases, which closely encircle it for some three-fifths of
its circumference. These leaves bases are embricated and provide three
distinct vascular strands each.
The phyllotaxy is of the 2/5 type with a very much shortened spiral.
The leaf traces themselves do not leave wide gaps, but above the branch
trace of the axillary bud, which is present over each median leaf trace,
there develops an inflorescence or a stolon. The branch gap often be-
comes completely confluent with that of its subtending median leaf
trace (White, 1927).
b. Lateral branch is a stem originating from an axillary meristem or
lateral bud. Morphologically it is similar to primary shoot with leaves,
stolons and can differentiate into a terminal inflorescence or originate a
new higher order lateral crown.1 If in contact with soil, it produces roots.
Lateral branches can be called branch crowns (secondary crowns) and
they are generally formed when a primary crown is slowing down in
growth, and stolon formation is ending in late summer or mid-fall. After
winter rest, the new spring growth may induce development of several
lateral crowns per plant from latent buds depending on the weather and
cultural conditions (Mann, 1930).
The lateral crown formed by the axillary meristem just below the in-
florescence is called an extension crown, and it will continue the plant
expansion (Figure 1).
Branch and extension crowns develop fairly consistently until their
subtending leaves emerge, usually at the 4 nodes stage. At this stage
they are likely to form a terminal inflorescence, if initiation has com-
menced at the crown apex.
In Japan, forced culture with high density planting (70,000 plants per
hectare) allowed only the extension crown, in order to avoid an extra
production per plant, which could be detrimental to fruit quality and to
homogeneity and continuity of the production (Neri et al., 2000). On
the contrary, production under low chilling conditions with short-day
cultivars permits a continuous winter production with a reiterating and
alternating extension crown growth and inflorescence differentiation
(Savini, 2003). This originates a cyclic vegetative-reproductive alter-
nate growth within branches of higher order. Single crowned plants can
be produced throughout the winter season up to temperatures that are
compatible with growth. Under these conditions, the delay in ripening
of the fruits belonging to the same inflorescence is sufficient to support
30 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

FIGURE 1. Formation of the extension meristem which will generate the exten-
sion shoot. It is the last axillary primordium formed in crown apex, during flower
differentiation in ‘Tochiotome.’

a continuous cropping system with 0.8-1 kg of fruits per plant over 5-6
months.
With summer planting of cold stored bare-root plants of Junebearing
cultivars, two to four lateral branches per plant are formed during late
summer and early fall. Such spring production is high, up to more than 1
kg per plant, with a short production season of less than one month.
c. Stolon is a specialised true stem with long internodes and a vascular
structure, which is markedly different from that of the short internoded
crowns (Ball and Mann, 1927). These differences are closely correlated
with some basic functional variations. The stolon is commonly two
nodes in length and is not continuous beyond its first daughter crown,
produced at the second node, whereas the first node usually remains
dormant or develops another small stolon. Thus the stolon differs from a
branch crown principally in the extensive elongation of the first two
internodes. The comparison with a developing branch crown meristem,
 Savini et al. 31

at the two leaf primordia stage, shows that the first internode of a stolon
bud is longer than the first internode of a branch crown bud (Figure 2).
The stolon forms a chain by which the meristem from the axil of the
first leaf becomes the daughter plant. The meristem in the lateral bundle
traces of this leaf is placed so nearly in direct line with the vascular bun-
dles of the parent stolon, due to the bending up of the tip of the stolon
(Figure 3), that it is in a favored position for its supply of water and nutri-
ents. Hence its development is commonly far more rapid than the devel-
opment of any of the axillary buds forming leaves later on the same plant.
The young stolon emerges from the sheathing stipules of its subtend-
ing leaf as the leaf unfolds and expands. At this stage the apex usually
shows two scale leaves and two true leaf primordia. Elongation of the
first two internodes continues and as the stolon approaches full length
the third leaf (the first true leaf) emerges from the sheathing stipules of
the second scale leaf. As the third and fourth leaves (first and second
true leaves) unfold, roots begin to break trough the surface of the stem
near the second node.
d. Inflorescence develops terminally in the crown. Inflorescence ini-
tiation is considered to have taken place when the apical growing point

FIGURE 2. Stolon formation starting from an axillary meristem in which there is

a very early consistent internode elongation.
32 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

FIGURE 3. Stolon growth and runner chain formation in ‘Tochiotome.’

has elongated and its top has become somewhat flattened. This stage
was often seen and probably is of short duration. Soon after this first in-
dication, the development of secondary flowers can be seen on the axis
of the inflorescence (Figure 4).
The branch structures of the strawberry inflorescence are defined as
“dichasial cymes” where individual trusses show great cultivar of
forms. The inflorescence structure is quite variable and many associated
physiological factors have been studied (Darrow, 1929; Waldo, 1930;
Robertson and Wood, 1954; Foster and Janick, 1969; Anderson and
Guttridge, 1982; Guttridge, 1985; Hancock, 1999). The axis of the in-
florescence, a principal peduncle, terminates with a primary flower and
emanates two lateral branches (secondary flowers). Similarly, two ter-
tiary ramifications come from secondary branches, each terminating in
a tertiary flower. Another ramification is also common and it originates
quaternary flowers (Figure 5).
The inflorescence structure is important in determining the timing
and quantity of fruit production (Webb et al., 1978). Ridout et al.(1999)
compared the branching structure in different commercial cultivars
with a simple stochastic approach modelling two parameters: number of
inflorescences per plant and flowers per each truss.
 Savini et al. 33

FIGURE 4. Primary flower with a deformed shape (it will likely originate a big
flat fruit) and two secondary flowers.

FIGURE 5. Inflorescence scheme of strawberry (redrawn from Jahn and Dana, 1970).

From the analysis of the literature it has been suggested that the
length of the inflorescence peduncle increases with the exposure to low
temperatures (chilling satisfaction), nevertheless this hypothesis re-
mains unclear. The inflorescences grown in spring (which are formed in
34 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

the preceding fall) are extremely compact even if they have been
exposed to the winter chilling.
An interesting alternative hypothesis on the control of the peduncle
length has been discussed by Faedi (1991). He emphasized that the
peduncle length of the inflorescence reveals the time of exposure of the
flower to induction conditions. This hypothesis was confirmed by
Savini (2003, Figure 6). The second internode did not change along the
experiment but the first one was strongly reduced by increasing the ex-
posure time (short day and temperature below 14°C) and not by the
chilling which followed the induction.
Finally, the inflorescence emerges from the protection of the sheath-
ing stipules of the leaf immediately below it, about the time the leaf
expands.
e. Bud–is originated by an axillary growing meristem which can be
identified at one and a half to two plastocrons from the inception of its
subtending leaf. Along the crown axis, the axillary meristems generally
originate two to four dormant buds at the base of young vegetative
plants (Neri et al., 2003). Then, when the growth is rapid under high

FIGURE 6. Different first node petiole length depending on time of flower in-
duction in the June-bearing cultivar ‘Paros’ (Savini 2003). Right: first truss in
late summer; Left: last truss formed in the autumn.
 Savini et al. 35

temperatures the axillary meristems develop into stolons, 4 to 15 per

plant in the nursery (Savini, 2003). Approaching growth arrest at
the end of the summer the axillary meristems produce buds again (Fig-
ure 7).
The axillary meristem immediately below an inflorescence on an es-
tablished plant has never been observed to develop into a stolon, al-
though it occasionally, develops into an inflorescence without further
leaf emergence. Only a sudden change from vegetative growth to
flower induction conditions may determine that the stolons growth is
immediately below the terminal inflorescences (Savini, 2003).
The buds along the plant axis may develop into branch crowns, dur-
ing or just after flower induction, in some cultivars or if the climatic
conditions are favorable (mild weather) (Guttridge, 1985). The rate of
leaf initiation on lateral buds was much less than on the main axis. A
similar slow growing condition was never observed in stolon growth.
Inflorescence initials were formed on the lateral buds after the forma-
tion of about three to four leaf primordia, as before, but these also were
small and developed slowly through the fall.

FIGURE 7. Axillary buds with different shape, and likely fate, in ‘Tochiotome.’
After their complete formation, they may remain dormant (latent) or develop
into a lateral shoot but rarely into a stolon.
36 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

If the apex is artificially excised during the growing season, axillary

meristems originate only as lateral branches or new crowns but never
stolons (Neri et al., 2003). The axillary meristems, which were already
growing as stolon but not emerged yet, developed into a 10-cm long sto-
lon similar to a lateral branch (Figure 8).
f. Leaf has a typical dicot structure, trifoliate with a long petiole and
well-developed and foliaceous basal stipules. The leaf petiole at its base
is semicircular in cross section. Petiole length is very susceptible to
thermo-photoperiod, short days and low temperatures can drastically
shorten it. Gibberellins can be also be used to counteract the action of
short days, inducing a programmed elongation in out-of-season produc-
tion (Guttridge and Thompson, 1959; 1963).
Petiole length has frequently been used as a parameter for dormancy
studies; being short when a dormant bud is moved into optimal growing
conditions without sufficient chilling requirement (Guttridge, 1958,
Jonkers, 1965; Gosselink and Smith, 1966, Le Mière et al., 1996; Rob-
ert et al., 1999). Petioles of newly emerging leaves are normal if the
chilling requirement is sufficiently met.
Chilling requirements for cultivated cultivars reaching the optimal
production are different in relation to planting date, growing media,

FIGURE 8. Shoot formation after artificial apex removal. Without the crown
apex lateral buds are able to develop into shoots or short stolons. The stolon
internodes (both primary and secondary) did not normally elongate.
 Savini et al. 37

nutrition, physiological stage (vegetative, reproductive), and genetic

background.

THE ANNUAL CYCLE AND FLOWER INDUCTION

In Junebearing cultivars inflorescence initials are present during the

first two weeks of September. Similar data for stolon plants cannot be so
easily determined. Although inconclusive, some authors indicate that
stolon plants rooted before September have inflorescence initials form
about the same time as their mother plants. Stolons, rooted during Sep-
tember, form initials around the time of rooting. Later rooting plants
commence initiation before rooting but later than the older plants.
Mother plants and older stolon plants were usually found to have sec-
ond order inflorescences by the time of dormancy, and a few crowns
had third order inflorescences (Robertson and Wood, 1954).
Inflorescence initiation does not normally occur on branch crown
buds before it has taken place on the primary axis. If inflorescence initi-
ation has taken place on the primary axis of the crown, the terminal in-
florescences commonly develop on branch crown buds at this stage or
soon after.
The presence of the terminal inflorescence on a crown is associated
with a change in the rate of extension growth of the three or four axillary
buds immediately below. In any cultivar under field conditions, these
buds were all found to be branch crowns. Other cultivars in the field and
under experimental condition in the greenhouse sometimes continue
stolon elongation, for a time at least, while inflorescence formation was
taking place (Guttridge, 1955). This can be explained by the delay be-
tween formation and elongation of the stolon. Some stolons might be in-
duced late in the season when the flower induction is starting because of
vegetative growth reduction; therefore they will apparently appear
when the flower induction is completed.

DESCRIPTION OF SPATIAL ORGANIZATION OF STOLONS

AND CROWNS

Axillary meristems can be seen on the young stolon before it has

emerged from the parent crown. At the time the stolon emerges, axillary
meristems are present at the first and second scale leaves and each has
one or two leaf primordia, the second being no more than a ridge on the
38 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

meristem. Both these axillary meristems differ from meristems devel-

oping in the axils of later, true leaves.
The meristem at the first node of a stolon may either remain as dor-
mant bud or develop as stolon, forming a branch on the stolon axis (Fig-
ure 3). The last stolon to develop in any one season sometimes fails to
attain full length but remains some five or ten centimetres long and is
thick and fleshy. This type of modification was seen more frequently on
experimental plants in the glasshouse than on plants in the field.
The meristem in the axil of the second scale leaf, which is at the base
of the young stolon plant, develops very rapidly and in this respect dif-
fers from later formed axillary buds. This is the meristem that forms the
continuation stolon (Robertson and Wood, 1954) in a stolon chain. The
stolon emerges from the protection of the sheathing stipules of its sub-
tending scale leaf as the primary stolon attains full length and before
root initials appear.
When environmental conditions are unfavorable for the differentia-
tion of the axillary buds into stolons (while favorable for flower induc-
tion), the axillary meristems develop into branch crown buds. Leaf
primordia develop on the axillary bud at about the same rate as on the
apical meristem of the main axis, so that by the time the subtending leaf
unfolds the axillary bud has about four leaf primordia. These buds differ
from stolon buds in the absence of appreciable stem tissue between the
nodes, also in the less precocious development of the bud in the axil of
the second leaf primordium.
The first (lowest) leaf primordium on the branch crown bud develops
only a small leaf blade and may be almost alaminate, although the stip-
ules are large and sheathing. Similarly, the blade of the second leaf is re-
duced, although not so markedly as the first. The final size of the lamina
of the first two leaves depends on the position of the branch crown in re-
lation to terminal inflorescences on the main axis. The nearer a branch
crown is to a morphologically higher inflorescence, the larger the
laminas of the first two leaves. The first two leaves on the axillary bud
immediately below the terminal inflorescences of the crown are nearly
normal in shape and size. It can be hypothesised that when the bud is
formed the stipules are big and leaf lamina small. In the stolon there are
not stipules which protect it when it is formed. In the inflorescence
when the petiole has big stipules it indicates a sudden arrest in growth.
Stipule dimensions can be used as a parameter for indicating growth
stasis and to reconstruct the history of growth cycles and arrest of the
single meristems on the whole plant.
 Savini et al. 39

THE ARCHITECTURAL MODEL

The features of the growth habit so far discussed are variable between
cultivars or crowns, or between plants of different ages or growing un-
der differing environments. The season of flower initiation and the posi-
tion of branch crown formation are heavily dependent on environment,
size of plant, and the cultivar.
The position on the axis of the lateral meristems, which develop into
stolons in spring, depends on the position of inflorescence meristems. It
was observed that the meristems subtended by the two lowest leaves on
the extension crown axis above the last-formed inflorescence com-
monly developed into the first stolon of the season.
It is also reported that stolon differentiation commences early in the
year and that probably the first axillary meristem, which is formed in the
spring, becomes the first stolon. This happens only in the absence of
flower and fruit growth, when vegetative plants with high growth rates
mainly originate from stolons by axillary meristems. On the contrary if
the growth is poor or flowers and fruits are abundant, the first axillary
meristems become buds. Stolons show a sylleptic2 behaviour (Champagnat,
1961; Neri et al., 2001) starting the growth immediately after their for-
mation as axillary meristems without a rest phase such as that of a bud
covered by leaf stipules.3
The stolons always develop from the axil of expanding or unfold
leaves, while lateral crowns originate at the axil of leaves, independ-
ently from leaf growth, confirming their sylleptic origin (Dana, 1974;
Savini 2003). Moreover a lateral crown may be initiated in the autumn
and grow in the following spring while a stolon is formed and grows in
the same season. Therefore the lateral crowns are to be considered
proleptic, being formed by axillary buds, which arrest the growth before
elongation.
The plant behavior in the field and under various experimental indoor
environments suggests that the development of buds into branch crowns,
does not depend on quantitative relationships between the plant and its
environment. Crown multiplication appears to be influenced by such
factors as plant size, which is in relation to its environmental and
growth conditions. The formation and maintenance of young stolon
plants during the summer affects crown multiplication by cutting off the
supply of newly formed branch crown buds, probably also by the com-
petitive suppression of growth of buds already present on the crown.
It is concluded that the buds will originate from the branch crowns
more or less immediately, if condition are favorable to an increase in
40 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

plant size during the time when newly formed branch crown buds are
present. This time is limited, in vigorously growing plants, to the period
between the emergence of the last stolon from the bud in late summer or
autumn and the emergence of the first stolon in spring, which approxi-
mately coincides with flowering. Axis development ends with an apical
flower bud. This crown proliferation is also dependent on the flower ini-
tiation of the terminal meristem. This fact determines the possibility of
growth only in newly formed lateral branch crowns, the number of
which is dependent on crown size.

THE SCHEME DRAWING

The architecture of the strawberry plant as extended axes was first

drawn by Guttridge (1955). In the present work, which conveniently
follow his general model, some variations are introduced. Plant organs
are presented with different colors i.e. dead leaves in yellow. Leaves are
trifoliate when they represent expanded leaves and linear when they are
still expanding or included in a bud within the stipules of the subtending
leaf. Cyclic growth is represented by a different color of the axis. The
crown growth is determined as well as any cycle that terminates with
flower differentiation and the physiological arrest of axis elongation
(Figure 9). Axillary meristems may be inactive for months and they
can break out in the next growing season after a chill period. The hi-
erarchy of bud growth is dependent on meristems correlation to-
gether with organs development. The strawberry plants show a high
phenotypic plasticity and interaction with environment and cultural
techniques. Breeding has selected particular habits well adapted to
forced culture and with potential high production and large fruits. Easy
stolonization is also desirable together with the capability to have lateral
branch formation in Junebearing cultivars.
The new cultivars are classified as Junebearing (facultative short
day), everbearing, and day-neutral types. Several studies reported er-
ratic response of plants to day length, and a new classification was
adopted: strong day-neutral; weak day-neutral; older everbearing; very
weak day-neutral, Junebearing (Nicoll and Galletta, 1987). Some re-
searchers (Darrow, 1966; Sonsteby and Nes, 1998; Savini, 2003) have
shown that this classification is not necessary because the cultivars are a
continuum between “Junebearing” and “everbearing” and the differ-
ences are only qualitative.
 Savini et al. 41

FIGURE 9. Schematic drawing of strawberry plant organs.

The fundamental diversity is due to the potential to make one or sev-

eral cycles during the same growing season. This character is linked
with stolon making capability. Junebearing cultivars have only one
growing flush while the everbearers have more cycles and they are less
inclined to produce stolons but they have more lateral branches. It
seems that vegetative growth is contrasted by flower differentiation and
vice-versa.
The physiological response of different genotypes to environmental
and cultural conditions can be described by the drawing of the architec-
ture of the plant in a dynamic analysis of the cultural cycle. As a case
study, the cultivated cultivar ‘Paros’ was compared in Northern and
Southern Italy, using the suggested architectural model (Figures 10 and
11). Besides, the architectural model of the same Junebearing cultivar
was compared with an everbearing cultivar in Northern Italy (Figure 12).
From both these comparisons with Junebearing and everbearing
cultivars it emerges that the architectural model is a useful tool to repre-
sent the vegetative plant behavior. It is worth noting that the growth
and reproductive cycle of the two cultivars is similar. The reproduc-
tive differentiation happens after a continuous growth of the new
shoot formed just below the terminal inflorescence of the higher or-
42 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

FIGURE 10. Schematic representation of the Junebearing cultivar ‘Paros’ at

the end of autumn. Plants were cultivated in northern Italy using a summer
planting of frigo plants (in grey) that experienced growth secession in Novem-
ber. 1°, 2° represent the order of the inflorescences on the new buds; the desic-
cated leaves are colored in orange.

der crown, leaving the old buds quiescent. Differences are evident
in the length of the vegetative cycle before flower formation, being
much shorter in the everbearing cultivars than in Junebearing ones.
Moreover the everbearing cultivars form more inflorescences per
plant.
The analysis of plant architecture is able to show the spatial relation-
ship between vegetative and reproductive organs, reporting the fate of
all the meristems, which are present in a plant along a selected period of
time. Thus it is possible to dynamically study the response of the plants
in the nursery, in the field, or under forced culture. Whereas in the litera-
ture there are only few papers of such analyses reported, and the under-
standing of plant behavior is often difficult and misunderstood because
the results are reported in a static way.
The signal response by the plant is different in relation to its physio-
logical phase; therefore, the same cultural techniques may give a very
different result depending on the plant quality from the nursery, and on
the interaction with environmental conditions. In the case of ‘Gariguette’
 Savini et al. 43

FIGURE 11. Schematic representation of the Junebearing cultivar ‘Paros’ at

the end of the autumn. Cultivated in southern Italy with summer planting of frigo
plants (in grey) and mild temperatures which induced continuous growth. The
desiccated leaves are colored in orange.

the suspension of the application of mineral nutrients in potted stolon

plants induced flower differentiation with only a reduction in the num-
ber of the internodes of the principal axis. Therefore if the architectural
analysis is properly applied, it can help in the determination of plant
quality in the nursery based on grower preferences.
Cultivation has modified notably the ability of the plant to form new
crowns along the principal axis. In fact, the frigo plants, before planting,
have only one terminal bud with an inflorescence, differentiated in the
preceding year before cold storage period. After transplanting, the di-
verse period of growth induced the formation of a different number of
lateral buds: three in central and northern (Jesi and Cesena) and one in
southern Italy. On the contrary, the total number of the inflorescences
per plant increases toward the south (Figures 13 and 14).
44 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

FIGURE 12. Schematic representation of the everbearing cultivar ‘Seascape’

at the end of the autumn. Cultivated in northern Italy with summer planting of
frigo plants (in grey), 1°, 2°, 3° represent the order of the inflorescences on the
new buds developed after planting; the desiccated leaves are colored in or-
ange.
3°

2° 3° 3°
2°
3°
3°

1°
1°

2°

2°

3°
3°

This behavior could be due to the period of planting, which is delayed

by about 30 days in the south. When the planting was carried out in
Cesena and Jesi (in the north at the end of July) climatic conditions fit
with the vegetative development and are favorable for branch crown
growth. Therefore the differentiation of terminal buds of these branches
may happen only subsequently, in September. After that differentiation
period the temperature drops close to 0° C in October and there is no
possibility to have 3° inflorescences, and only few 2° order develop.
Contrarily, in the south (Metaponto), later planting occurs in tem-
perature and photoperiod conditions that are favorable to the flower
differentiation, and this produces low vegetative growth with only one
extension crown per plant. The bulk of the flower differentiation occurs
in the buds of 2° and 3° order on the extension crown because mild
 Savini et al. 45

FIGURE 13. Comparisons between the crown growth and the number of inflo-
rescences per crown at the three different locations. Different letters indicate
statistical differences: Newman-Keuls test, p < 0.05.

6 b
5 ab
n./ plant

4 d
a
d
3
2
c
1
n. inf./crown
0
n. crown
Cesena
Jesi
Metaponto

FIGURE 14. Comparison among the number of flowers in the primary and sec-
ondary inflorescences at three different locations. Different letters indicate sta-
tistical differences: Newman-Keuls test, p < 0.05.

weather during autumn allows for a prolonged period favorable to dif-

ferentiation. In this condition, vegetative growth may easily be resumed
using artificial lighting or gibberellins application.
Moreover in northern Italy, the inflorescence in the primary bud has
induced a very high number of flowers (12-13 per inflorescence) in
comparison to the south (10 per inflorescence). This behavior could be
due to the prolonged duration of differentiation that the plant received in
the north during the growth of the primary buds. This favorable time is
46 INTERNATIONAL JOURNAL OF FRUIT SCIENCE

drastically reduced during secondary buds differentiation (Figure 14).

While in the south, the tendency is to sustain a constant differentiation
for the whole autumn, which both orders of inflorescence have a more
similar number of flowers.

CONCLUSION
The architectural analysis proved to be a useful tool for evaluating
the vegetative and reproductive organization of the strawberry plant. It
is particularly useful for evaluation of nursery material in respect to
their production potential. Additionally, it is a useful way for monitor-
ing the efficiency of the best cultivation techniques in each climatic
condition for every genotype.
The analysis is particularly interesting for the study of phenotypic
adaptation of similar genotypes in two different environments of cultivation
(northern and southern Italy). Clearly identifying flower differentiation
patterns is one of the reasons for the prolonged production (about 40
days longer) in the south.
It has been noticed that side ramification happens only after the for-
mation of the terminal inflorescences on the main axis. This supposes a
sort of vegetative control of this meristem in the growth of the lateral
buds and causes the typical spatial organization of the strawberry plant.
Perhaps in the future we will be able to evaluate the architectural
model in the cultivar evolution and permit a better programming of the
total production and increase in fruit quality.

NOTES
1. Similar to fractals in mathematics, simple architectural models can be reiterated
to different levels.
2. Sylleptic shoots develop from axillary meristems on a growing shoot without any
rest period after their induction, and with a contemporaneous growth with the main axis
from which they are formed.
3. In the axillary bud the leaves may have a reduced petiole and no lamina so that
the stipules develop as bracts covering completely the bud.

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