Annals of Botany 80 : 419–425, 1997

Cell Number and Cell Size in Parthenocarpic vs. Pollinated Blueberry

(Vaccinium ashei) Fruits
R A Q U E L C A N O-M E D R A N O* and R E B E C C A L. D A R N E LL‡
Horticultural Sciences Department, UniŠersity of Florida, GainesŠille, FL 32611, USA

Received : 26 February 1997 Accepted : 1 May 1997

Gibberellic acid (GA ) promotes parthenocarpic fruit development and is used commercially to increase fruit set in
$
many crops. However, fruit size is usually smaller than that of pollinated fruit. The purpose of this work was to
determine the anatomical basis for differences in fruit size between pollinated and GA -induced parthenocarpic
$
blueberry (Vaccinium ashei Reade) fruits. Fresh weights at ripening averaged 1±6 and 2±5 g for GA -treated Šs.
$
pollinated fruits, respectively. In both pollinated and GA -treated fruits, mesocarp cell number comprised about 75 %
$
of the total pericarp cell number, and increased from C 7000 cells per cross-sectional area at bloom to C 9000 at
harvest. The duration of the cell division period in pollinated and GA -treated fruits was similar, with the majority
$
of cell division ceasing by 24 d after bloom (DAB). Cell size in both middle and inner mesocarp of ripe pollinated
fruits was significantly larger than in ripe GA -treated fruits (31 000 Šs. 22 000 µm#). Differences in final fruit size
$
between pollinated and GA -induced parthenocarpic blueberry fruit are due to differences in cell enlargement rather
than cell number.
$ # 1997 Annals of Botany Company

Key words : Blueberry, cell number, cell size, gibberellic acid, parthenocarpy, Vaccinium ashei.

esculentum Mill.) fruit, exogenous applications of auxins to

INTRODUCTION
In blueberry (Vaccinium spp.), parthenocarpic fruit set and
development is induced by exogenous applications of
gibberellic acid (GA ) ; however, final fruit size of partheno-
$
carpic fruits is smaller than that of pollinated fruits
(Williamson et al., 1995). The smaller fruit size of GA -
$
induced parthenocarpic fruits has also been observed in
other crops, including grape (Vitis Šinifera L.) (Iwahori,
Weaver and Pool, 1968), cranberry (Vaccinium macrocarpon
Ait.) (Devlin and Demoranville, 1967), peach (Prunus persica
L.) (Stembridge and Gambrell, 1972), and citrus (Citrus
reticulata Blanco) (Garcı! a-Martı! nez and Garcı! a-Papı! , 1979).
Cell number at anthesis, the length of the cell division
period after anthesis, and the extent of cell enlargement
determine final fruit size in a number of fruits (Coombe,
1976). Plant growth regulators may induce parthenocarpic
fruit set and development by directly affecting cell division
and}or cell enlargement. Cytokinin-induced parthenocarpic
fruit development in cucumber (Cucumis satiŠus L.) was
accompanied by increased cell division in the pericarp
compared to the non-pollinated fruits (Takeno et al., 1992).
Exogenous auxin application to non-pollinated watermelon
(Citrullus lanatus (Thunb.) Matsum. & Nakai) flowers
increased both cell division and cell enlargement in pericarp
tissue over that of the non-pollinated control (Sedgley,
Newbury and Possingham, 1977). In both instances,
hormone-induced parthenocarpic fruits were similar in size
to their pollinated counterparts. In tomato (Lycopersicon
emasculated flowers transiently increased the rate of cell
division compared to the pollinated control, and resulted in
fruit that was similar in size to the seeded fruit (Bunger-
Kibler and Bangerth, 1983). On the other hand, application
of GA increased cell enlargement, but decreased cell
$
division throughout tomato fruit development compared to
the pollinated control, and resulted in fruit that was
significantly smaller than the seeded fruit. In pea (Pisum
satiŠum L.) ovaries, GA applied at anthesis induced
$
mesocarp cell division and enlargement (Vercher et al.,
1984), enhanced cell division in the endocarp (Vercher,
Molowny and Carbonell, 1987), and promoted partheno-
carpic fruit development compared to the non-pollinated
ovaries. However, GA -treated ovaries were significantly
$
smaller than pollinated ovaries. It is unclear from this work
if the smaller size of GA -treated compared to pollinated
$
ovaries was due to a decrease in cell size, cell number, or
both.
Although GA -induced parthenocarpic fruit development
$
occurs in a variety of crops, and development is significantly
enhanced compared to non-pollinated fruits, growth is
usually less than that observed in pollinated fruits. The
smaller final fruit size of GA -induced parthenocarpic
$
blueberry fruit compared to pollinated fruit may be a result
of decreased cell number and}or decreased cell size. The
objective of this study was to determine the anatomical
basis for differences in fruit size in pollinated, GA -induced
$
parthenocarpic, and non-pollinated blueberry fruits.

* Current address : Colegio de Postgraduados en Ciencias Agri-

colas, Monticell-Texcoco, 56230 Edomex, Mexico.
‡ For correspondence. Fax ­352 392 6479

0305-7364}97}100419­07 $25.00}0 bo970462 # 1997 Annals of Botany Company

420 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes
MATERIALS AND METHODS
Softwood cuttings of Vaccinium ashei Reade cv. Beckyblue
were rooted in spring 1990 and grown outdoors in 22-l pots
in a 1 : 1 peat : pine bark mix. Plants were watered every
other day and fertilized with 20N-5±6P-11K water soluble
fertilizer. In December 1991, nine uniform plants were
transferred to a dark cooler and chilled at 7³1 °C for 30 d.
After chilling, plants were placed in a glasshouse with
average day}night temperatures of 27}16 °C to force
budbreak. Flower clusters were thinned to four to five
florets per cluster at bloom, removing the most developed
and least developed florets. The total number of florets per
plant ranged from 200 to 215. Flowers were either hand
pollinated at bloom with pollen from a cross-compatible
cultivar, or sprayed at bloom and again at 7 d after bloom
(DAB) with 0±7 mGA (Pro Gibb 4 %, Abbott Labora-
$
tories, Chicago, ILL, USA, prepared in McIllvaine buffer
pH 3±5, and 0±01 % Tween-80). Flowers remaining non-
pollinated were sprayed at bloom with buffer and surfactant.
Plants subjected to these treatments were arranged in a
F. 1. Median cross section of a ‘ Beckyblue ’ blueberry fruit at 0 days
after bloom. Bar ¯ 150 µm. ep, Epidermis ; hp, hypodermis ; om, outer
mesocarp ; mm, middle mesocarp ; im, inner mesocarp ; en, endocarp ;
vb, vascular bundles ; lc, locule ; pl, placental tissue ; ov, ovule.
randomized design with three single plant replications per
treatment.
Three fruit sub-samples were taken at 0, 3, 19, and 24
DAB for all treatments. Subsequent to the abscission of
non-pollinated fruits (between 24 and 45 DAB), sub-
samples were taken at 45 DAB and at ripening for pollinated
and GA -treated fruits. Fresh weight, length, and diameter
$
were measured for each harvested fruit.
At each sampling date, harvested fruit were fixed
individually in formalin-acetic acid-ethanol mixture (FAA).
Fruit median cross sections were dehydrated in an ethanol-
tertiary butyl alcohol series, and embedded in paraplast.
Fruit were sectioned at 8 to 20 µm, attached to slides with
albumen fixative and stained with safranin and fast green.
Sections were mounted in Permount after staining.
Cell number and cell size were determined from micro-
photographs taken of the median fruit cross section. A
Lasico planimeter (Model gL10, series 81191, Los Angeles
Scientific Instrument Co. Inc. Los Angeles, CA, USA) was
used to measure the total area of the amplified photograph
as well as the cell area of 10 cells considered representative
of each of the following regions : epidermis, hypodermis,
outer mesocarp, middle mesocarp, inner mesocarp and
endocarp (Fig. 1). The outer mesocarp was defined as the
region between the epicarp and the first large vascular
bundle, the middle mesocarp was the region between the
two large vascular bundles, and the inner mesocarp was
the region between the second large vascular bundle and the
endocarp. Vascular bundle areas as well as the locular area
were subtracted to obtain total mesocarp area. Values were
converted to µm# using a reference area of 1 mm# from a
haemocytometer chamber that was microphotographed
each time new film was used. Cell number in a given cross
sectional area was determined by following the equations
used by Scorza et al. (1991). Briefly, the area of the whole
fruit cross section was determined by measuring ovary}fruit
diameter and using the equation for the area of an ellipse.
Cell number in a whole fruit cross section was calculated by
using the equation x}a ¯ x"}a", where x is the cell number
in a given sample cross section, a is the area of the given
cross section, x" is the cell number in the whole fruit cross
section, and a" is the area of the whole fruit cross section.
RESULTS
Fruit growth
Growth of GA -treated fruits followed a similar pattern as
$
that of pollinated fruits ; however, fruit weight was signifi-
cantly lower in the GA -treated fruits from 45 DAB to
$
ripening (Fig. 2). Non-pollinated fruits abscised between 24
and 45 DAB and weighed significantly less than both GA
$
and pollinated fruits at this time. The fruit development
period averaged 72 d for pollinated fruits and 87 d for GA -
$
treated fruits.
The histological composition of a blueberry ovary at 0
DAB in cross section is depicted in Fig. 1. The individual
pericarp tissues were comprised of a single epidermal and
single hypodermal layer (which together formed the epi-
carp) ; several layers of mesocarp ; and the endocarp, a single
 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes 421
3.0
15 GA3 Endocarp Mesocarp Epicarp
GA3 POLL NP A A
B
10 C BC
2.5

5 cb cb ba a a
2.0

Cell number/x.s. (thousands)

0
Fresh weight (g)

15 POLL
1.5 A A

10 C
B B

1.0 5 cb c c ba a

0
0.5 15 NP

B BC A
10 BC
0 20 40 60 80
Days after bloom 5 cb cb cb cb
F. 2. Developmental changes in fresh weight of GA -treated (GA ),
$
pollinated (POLL), and non-pollinated (NP) ‘ Beckyblue ’ blueberry
$
fruits. Values are means³s.e., n ¯ 3 ; s.e. bars present only when larger 0
0 3 10 24 72 87
than symbol. Non-pollinated fruit abscised between 24 and 45 DAB.
Days after bloom

F. 3. Cell number changes per median cross sectional area (x.s.) in
developing pericarp of GA -treated (GA ), pollinated (POLL), and
$ $
non-pollinated (NP) blueberry fruits. Mean separation across treatment
layer of cells lining the locules. Throughout development,
mesocarp tissue in all treatments comprised 90 to 98 % of and time by LSMeans, P ¯ 0±05.
the total pericarp tissue.

However, there were no differences in cell size between GA -

$
Epicarp cell number and size treated (C 1860 µm#) and pollinated fruits (C 2140 µm#) at
ripening. The overall increase in hypodermal cell size was
Epicarp cell number was similar among the three
about 3±5 times from 0 DAB to ripening.
treatments throughout development. Cell number at an-
Both epidermal and hypodermal cells were rectangular-
thesis averaged 1170 cells per cross sectional area (Fig. 3),
shaped, but hypodermal cells were larger than epidermal
increased gradually for all treatments, and at ripening,
cells. At 0 DAB only one layer of each tissue was easily
averaged 1800 and 2450 cells per cross sectional area for
distinguished (Fig. 1), but as development progressed, two
GA -treated and pollinated fruits, respectively. Epicarp cell
$ or three cell tiers of hypodermal cells were observed (Fig. 4).
number increased 1±4-fold in GA -treated fruits and two-
$ Pigment accumulation (appearing as darkened regions) was
fold in pollinated fruits throughout development. There was
observed in hypodermal cells at 24 DAB, and a massive
a 1±7-fold increase in epicarp cell number in non-pollinated
accumulation of pigment was observed at ripening in both
fruits prior to abscission.
the epidermis and hypodermis (Fig. 5).
Cell size in epidermal tissue did not change significantly
from 0 to 10 DAB in any treatment, averaging C 400 µm#.
By 24 DAB, epidermal cells of GA -treated fruits were
$ Endocarp cell number and size
significantly larger (C 600 µm#) than pollinated and non-
pollinated fruits (C 470 µm#). However, by ripening, epi- Endocarp cell number was similar among treatments
dermal cell size in both pollinated and GA -treated fruits throughout development (Fig. 3). Endocarp cell size
$
were similar, averaging 1175 µm#. increased about 6±5-fold from bloom to ripening in both
Increases in hypodermal cell size followed a similar GA -treated and pollinated fruits, with no difference
$
pattern, and averaged C 500 µm# at 10 DAB for all between treatments. At ripening, cell size averaged 1130 µm#
treatments. Differences among treatments were not evident for both treatments. Cell size in non-pollinated fruits was
until 24 DAB, when cell size in GA -treated fruits was about significantly less than that of GA -treated and pollinated
$ $
1±6-fold greater than in pollinated and non-pollinated fruits. fruits by 24 DAB (C 350 µm# Šs. C 500 µm#).
422 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes

F. 4. Median cross section of (A) GA -treated, (B) non-pollinated, and (C) pollinated ‘ Beckyblue ’ blueberry fruit at 24 days after bloom.
$
Bar ¯ 150 µm. ep, Epidermis ; hp, hypodermis ; om, outer mesocarp ; mm, middle mesocarp ; im, inner mesocarp ; en, endocarp ; vb, vascular
bundles. Note differences in mesocarp cell size between non-pollinated and GA -treated or pollinated fruits.
$

significantly smaller than in the other two treatments

Mesocarp cell number and size
Although mesocarp cell number in both pollinated and
GA -treated fruits increased between bloom and ripening,
$
C 80 % of the cells in ripe fruit were already present prior
to bloom. Cell number increased from C 6900 to C 8900
cells per cross sectional area between bloom and ripening,
with no consistent differences between treatments within a
given sampling date or phenological stage (Fig. 3). Cell
number in non-pollinated fruits did not increase between
anthesis and fruit abscission.
There were no differences in cell size in the outer mesocarp
of GA -treated and pollinated fruits throughout fruit
$ $
development (Fig. 6). Cell size increased from C 900 µm# at
0 DAB to C 16 270 µm# at ripening. At 24 DAB, cell size in
the outer mesocarp tissue of non-pollinated fruits was
(4035 µm# Šs. 5400 µm#).
Cell size in the middle and inner mesocarp tissue averaged
C 900 µm# at bloom. By 24 DAB, cell size in both tissues was
significantly larger in pollinated fruits than in GA -treated
$
fruits (Fig. 6). Enlargement also occurred in non-pollinated
fruits, but by 24 DAB average cell size was significantly
smaller than that of pollinated and GA -treated fruits. At
$
ripening, cell size in the middle and inner mesocarp of
pollinated fruits averaged 40–45 % more than cell size in
GA -treated fruits (C 31 250 Šs. C 21 950 µm#). Throughout
$
development, cell size in middle and inner mesocarp tissues
increased 25-fold in GA -treated fruits and 33-fold in
pollinated fruits.
There was a high correlation between mesocarp cell size
and fruit fresh weight across treatments (r ¯ 0±93, P %
 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes 423
20
Outer mesocarp
16

12

8
GA3 POLL NP
4 *

0
Middle mesocarp *

Cell size (ím2 × 1000)

28

*
20

12 *
*
*
4
0
*
Inner mesocarp
28
*
20
*
12 *
*
4

0 10 20 30 40 50 60 70 80 90
Days after bloom
F. 6. Increases in cell size in developing mesocarp tissues of GA -
treated (GA ), pollinated (POLL), and non-pollinated (NP) blueberry
$
$
fruits. Values are means³s.e., n ¯ 3 ; s.e. bars present only when larger
than symbol. Asterisks indicate means that are significantly different
among treatments within a given phenological stage by LSMeans,
P ¯ 0±05.

F. 5. Median cross section of (A) GA -treated and (B) pollinated
$
‘ Beckyblue ’ blueberry fruit at ripening (87 and 72 DAB, respectively).
Bar ¯ 150 µm. ep, Epidermis ; hp, hypodermis ; om, outer mesocarp ;
mm, middle mesocarp ; im, inner mesocarp ; en, endocarp ; vb, vascular
bundles ; sc, sclereids. Note differences in middle and inner mesocarp
cell size between GA -treated and pollinated fruits.
$
0±01), while cell number and fresh weight were poorly
correlated (r ¯ 0±38, P % 0±16). Increases in mesocarp cell
size closely paralleled fruit fresh weight increases throughout
development in all treatments (Fig. 7).
DISCUSSION
The fruit cell division period in epicarp and mesocarp
tissues of pollinated and GA -treated blueberry fruits
$
occurred during the first 24 DAB. Cell division also occurred
in the epicarp of non-pollinated fruits, but no division was
apparent in the mesocarp tissue of these fruits. These results
are consistent with those reported for other pollinated fruits
such as peach (Jackson, 1968 ; Scorza et al., 1991), grapes
(Harris, Kriedemann and Possingham, 1968), apricot
(Prunus armeniaca L.) (Jackson and Coombe, 1966), and
sour cherry (Prunus cerasus L.) (Tukey and Young, 1939),
where cell division occurs primarily during the first stage of
growth in fruits exhibiting a double-sigmoid growth curve.
Edwards (1970) reported increases of two to three-fold in
mesocarp cell number at the end of the first growth stage in
some rabbiteye blueberry clones. However, only a 1±2-fold
increase in mesocarp cell number was observed in our study.
These differences may be due to the different counting
methods ; in the work by Edwards, mesocarp cell number
was determined as the average distance from the endocarp
to the epicarp. It was unclear, however, whether intralocular
shape was taken into account in his cell counts as it was in
ours.
424 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes
GA3 Cell size Fresh weight Cell number
25
20
15
Cell number (thousands) and cell size ( ím2 × 1000)
10
5 was similar among treatments at 10 DAB. By 24 DAB,
0 3
POLL
25
20
15
10
5 $
0 3
NP
25
20
15
10
5 according to the crop. In apple, fruit size appears to be
0 20 40 60 80
Days after anthesis
F. 7. Developmental changes in mesocarp cell size, fruit fresh weight,
and pericarp cell number in GA -treated (GA ), pollinated (POLL),
$ $
and non-pollinated (NP) blueberry fruits.
Cell counts may also be affected by fruit shape, and in
some cases, elongated fruits have resulted from GA
$
applications (Weaver and McCune, 1960). However, in our
study, the fruit length : diameter ratio was similar among
treatments throughout development ; thus we assumed cell
counts were not influenced by treatment differences in fruit
shape.
Although cell division occurred in mesocarp tissue of
both pollinated and GA -treated fruits after anthesis, the
$
majority of cells present at ripening were formed pre-
anthesis. In many fruits, cell division slows markedly after
anthesis (Coombe, 1976). For example, in apple (Malus
domestica Borkh.), only three to four cell generations are
formed after anthesis, compared to C 20 before anthesis
(Coombe, 1976 ; Goffinet, Robinson and Lakso, 1995). Yet,
actual cell numbers formed after anthesis significantly
exceed the number formed prior to anthesis. This appears to
be true in a wide variety of fruits other than apple, such as
strawberry (Fragaria¬ananassa Duch.) (Cheng and Breen,
1992), peach (Scorza et al., 1991), and apricot (Harris et al.,
1968), where cell numbers formed after anthesis comprise 80
to 97 % of the total cell number. The situation in blueberry
is more analogous to that in cucumber in which C 70 %
of the total pericarp cell number is formed before anthesis
(Marcelis and Hofman-Eijer, 1993). Certain Rubus and
Ribes species are also reported to complete pericarp cell
division prior to anthesis (Coombe, 1976). Thus, factors
affecting cell division during flower bud development in
3 these crops, including blueberry, would have a much greater
impact on final fruit size than factors affecting cell division
during fruit development.
2 On the other hand, factors affecting cell enlargement
during blueberry fruit development would be expected to
have a much greater impact on final fruit size than factors
1 affecting cell division. In blueberry mesocarp tissue, cell size
mesocarp cell size and overall fruit weight were significantly
smaller in non-pollinated fruits compared to pollinated and
GA -treated fruits, indicating that cell enlargement and
Fresh weight (g)
$
2 fruit growth is dependent on the stimulus provided by
pollination or GA applications. This stimulus was only
$
partially provided by exogenous GA applications, however,
$
1 since pollinated fruits exhibited a marked increase in
mesocarp cell size compared to GA -treated fruits between
45 DAB and ripening. This result could possibly be
attributed to an exhausted GA supply, however, previous
$
work indicated that additional applications of exogenous
GA at 21 and 42 DAB were unsuccessful in stimulating
2 $
further pericarp growth compared to the applications used
in the present study (Cano-Medrano, 1994).
1 The relative contributions of cell size and cell enlargement
to final fruit size are not always easy to determine, and differ
0 determined primarily by the extent of cell division after
pollination (Goffinet et al., 1995 ; Pearson and Robertson,
1952). Final fruit size in peach and strawberry is dependent
on the extent of cell division both before and after pollination
(Cheng and Breen, 1992 ; Scorza et al., 1991). On the other
hand, final fruit size in grape and cucumber appears to be
determined primarily by the extent of cell enlargement
(Harris et al., 1968 ; Marcelis, 1993).
In our study, differences in fruit size among pollinated,
GA -treated, and non-pollinated blueberry fruits were due
$
primarily to differences in cell size, with cell number playing
only a minor role. In mesocarp tissue, cell number in both
pollinated and GA -treated fruits increased 1±2-fold, while
$
cell size increased 25 to 30-fold throughout development.
The importance of cell size in determining fruit weight in
blueberry is supported by a high correlation (across all
treatments) between cell size and fruit fresh weight, while
the correlation between cell number and fresh weight was
insignificant. At ripening, GA -treated and pollinated fruits
$
had similar mesocarp cell number, but both mesocarp cell
size and fresh weight of GA -treated fruits were reduced by
$
about 67 % compared to pollinated fruits.
Cell enlargement has been described in physical terms as
a function of cell wall extensibility, osmotic potential of the
cell, and}or turgor pressure (Cosgrove, 1993). Gibberellins
are believed to be involved in both cell wall extensibility and
osmotic potential. Exogenous applications of gibberellins
have promoted enlargement by increasing cell wall ex-
tensibility in AŠena satiŠa L. stem segments (Adams et al.,
1975), Phaseolus Šulgaris L. leaves (Brock and Cleland,
1990), and wheat (Triticum aestiŠum L.) leaves (Keyes,
Sorrels and Setter, 1990). In cucumber hypocotyls, however,
GA induced enlargement by decreasing the osmotic
$
potential rather than increasing extensibility (Katsumi and
Kazama, 1978).
 Cano-Medrano and Darnell—Blueberry Fruit Cell Numbers and Sizes
In a previous study, GA -treated and pollinated fruits
$ of grape berry development. Vitis 7 : 106–119.
began accumulating considerable amounts of sugars at the
same time (24 DAB) and in similar concentrations (Cano-
Medrano and Darnell, 1997), but the increase in solute
concentration was not sufficient to elicit cell enlargement in
GA -treated fruits to the same extent as in pollinated fruits.
$
This suggests that decreased osmotic potential in pollinated
compared to GA -treated fruits was not the basis for
$ various tree factors. Australian Journal of Agricultural Research
differences in cell enlargement. Decreased wall extensibility
may be responsible for the decreased mesocarp cell
enlargement rate and resultant decrease in cell size in GA -
$ cucumber hypocotyl sections. Botanical Magazine. Tokyo Special
treated fruits compared to pollinated fruits, although this
remains to be determined.
A C K N O W L E D G E M E N TS
University of Florida Journal Series no. R-05698.
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