New Zealand Journal of Botany

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Structure and development of fruit and seeds in

Chinese gooseberry (Actinidia chinensis Planch.)

M. E. Hopping

To cite this article: M. E. Hopping (1976) Structure and development of fruit and seeds in
Chinese gooseberry (Actinidia chinensis Planch.), New Zealand Journal of Botany, 14:1, 63-68,
DOI: 10.1080/0028825X.1976.10428651

To link to this article: https://doi.org/10.1080/0028825X.1976.10428651

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New Zealand lournal of Botany. 1976. Vol. 14: 63-8. 63

Structure and development of fruit and seeds in Chinese gooseberry

(Actinidia chinensis Planch.)

M. E. HOPPING

Plant Diseases Division, DSIR, Private Bag, Auckland, New Zealand

"

(Received 15 July 1975)

ABSTRACT

The development of fruit and seed tissues in Chinese gooseberry (Actinidia chinensis
Planch. cv. 'Monty') was examined at intervals after flowering. Cell division in all fruit tissues
commenced immediately after flowering and persisted in the outer pericarp, inner pericarp,
and central core for 23, 33 and 111 days, respectively.
The fruit growth curve was double sigmoid owing to an initial period of cell enlarge-
ment in all tissues (Stage I, 0-58 days) followed by a period of retarded enlargement
(Stage H. 58-76 days)' and a further period of enlargement in the inner pericarp (Stage III.
76-160 days after flowering).
The structure of the seed was unusual in that the nucellus possessed a well dcfined
hypostase, and the endosperm and embryo were surrounded by an endothelium. Only one,
uniseriate, integument was present which subsequently developed into a thickened testa. The
cellular nucellus and endosperm developed concomitantly until the end of Stage II and were
absorbed, in part, during Stages II and III by the developing embryo.

INTRODUCTION both local and export demand. The important com-

The genus Actinidia, which contains 36 species
and several varicties (Li 1952), extends from Japan
through north-western Asia to Western China and
as far south as Malaysia. All members of the genus
are climbing dioecious vines which grow in thickets
~Ind along forest margins. Van Tieghem (1899)
trected the family Actinidiaceae on the basis of
ovule structure and anatomy. The family rank of
Actinidiaceae has been recently confirmed by Vi jay-
arabhavan (1963) and assigned to the Theales rather
than the Dilleniales (Johri 1963).
At least two species, A. arguta Sieb. & ZUCCo and
A. chinensis Planch., are of economic importance
because of their large, edible fruit although several
other species, A. arisanensis Hayata, A. eriantha
Benth., and A. [ali/olia Merr.• have fruit that are
comparable in diameter (Li 1952).
In New Zealand, A. chinensis, the Chinese goose-
berry or kiwifruit, has been established as a fruit
crop (Fletcher 1971) and in recent years the acreage
under cultivation has rapidly been increased to meet
mercial characteristics of fruit shape and size depend
on cell multiplication and enlargement in the fruit.
Although the structure of the seed tissues has been
recorded (van Tieghem 1899, Crete 1944, Johri 1963)
no similar studies on the structure of the fruit tissues
have been pUblishd. This paper reports a morpho-
logical study of both seed and fruit development
with emphasis on the correlations between the de-
velopment of tissues in the seed and the duration of
cell division and enlargement in the fruit.
MATERIALS AND METHODS
A representative sample of Chinese gooseberry
(Actinidia c?inensis Planch. cv. 'Monty') fruits was
collected at Intervals commencing at flowering (75%
of flowers open) from mature vines growing at the
Oratia Research Orchard, Auckland. Twenty fruits
from each sample were immediately weighed and
then oven d.ried to constant weight. The remainder
were fixed In formalin: acetic acid: ethyl alcohol
(60%), (1: 1 :20).
64
Fixed fruits were later cut in half transversely,
and the thicknesses of the central core and of the
inner and outer pericarp were measured along a radial
line from core to epidermis using a dissecting micro-
scope fitted with a micrometer eyepiece. Median
transverse sections (IS /Lm) were cut from portions
of these fruits on a freezing microtome, stained with
haematoxylin, and the cell number in each of the
three tissues was counted in a radial direction. Mean
cell size was calculated for each tissue from tissue
width : cell number determinations. All measure-
ments were repeated on at least 10 fruits.
To further study the structure of the fruit, perm-
anent mounts of fruit tissues at selected stages of
development were prepared by first infiltrating tissue
segments with 20% gelatin (Galigher & Kozloff
1964) for 3 hr at 40·c and then hardening them in
4% formalin for 24 hr. Sections (freezing micro-
tome. 15/LM) were dehydrated in an ethyl alcohol
serie~ and stained with fast green.
Seeds were removed from the inner pericarp under
a dissecting microscope, cut in half longitudinally,
and the length of the seed tissues (nucellus, endo-
sperm. and embryo) was measured as an index of
seed development. At least one seed from each of
10 fruits was measured. Portions of the inner peri-
carp were dehydrated (tertiary butyl alcohol). em-
bedded in paraffin (melting range 52-54·c) and
median transverse sections (13I'm) were cut on a
rotary microtome. Sections were affixed to slides with
Haupts' adhesive and 2% formalin. and stained with
safranin and fast green.
RESULTS
Fruit Itrueture
The Chinese gooscberry fruit developed from a
superior, multicarpellate ovary (26-38 carpels/fruit,
Fil. lA). Each carpel contained numerous OVUles
(Fig. tA, t8), borne in distinct locules (axile placen-
tation) that became filled with thin-wa11ed placental
cells. 8ecause the entire ground tissue developed into
a fte.hy tissue the fruit is a berry (Esau 1965).
At flOWering the ovary was bounded by a uniseri-
ate epidennal layer that subsequently developed into
a rnu1t1aeriate epidennis (Fia IC). A cuticle covered
the outermost layer of cells; the inner layers (2-4 cells
wide) had thickened cell walls and extensive tannin
depoaits. At irregular intervals, multi-celJular hairs
protruded from the outer epidermal layer. Many of
theie hairs were broken off durina fruit development
to leave areas of disrupted cells that did not possess
the characteristic phellolen of lenticels. The pericarp
New Zealand Journal of Botany 14, 1976
consisted of an outer layer of thin-walled parenchy-
matous cells which extended from the epidermis to
the outer bundles (outer pericarp, Fig. lA, ID), and
an inner layer of elongated septum cells which ex-
tended from the outer bundles to the core and sur-
rounded the locules (inner pericarp, Fig. I A, I B) .
The outer pericarp could be further divided into an
outer hypodermal layer, 10-15 cells wide, that
extended inwards from the epidermis (Fig. ID), and
an inner layer of elliptical cells that extended from
the hypodermal layer to the outer bundles.
During pericarp expansion, cells of the hypodermal
layer elongated in the tangential plane whereas those
of the inner layer increased several fold in diameter
and became essentially spherical. However, some cells
in both layers did not enlarge appreciably and, at
maturity, filled the spaces between cells that had
increased in size. Enlargement of septum cells. which
enclose the locules and collectively make up the
inner pericarp (Fig. IE), was almost entirely in the
radial direction although some tangential enlarge-
ment occurred in the zone immediately adjacent to
the outer pericarp. The cells of both pericarp layers
contained chloroplasts that persisted until maturity.
During maturation chlorophyll was lost from the
inner pericarp but not from the outer pericarp.
Two "rings" of vascular bundles were observed in
fruit tissues. The outer vascular bundles were located
between locules in the transition zone between the
inner and outer pericarp (Fig. lA, 18, ID) and
branched in the radial plane towards the epidermis.
The other series of vascular bundles, the inner
bundles, occurred at the base of each locule (Fig.
lA, 18, IF). Vascular traces from these bundles
extended into each locule and terminated at the
micropyle of each seed (Fig. 38). No branches from
the inner bundles extended into the cells of the
septum or into the parenchymatous central core.
The locule wall. which enclosed the developing
seeds, was bounded by a uniseriate epidennis possess-
ing a thin cuticle. At flowering, no cellular contents
other than the seeds were observed inside the locules
but as the locules enlarged they became progressively
filled with large placental cells. During maturation
the placental cells degenerated and produced a muci-
laginous matrix which supported the seeds.
Fruit development
The growth curve of developing Chinese goose-
berry fruit, a.s measured by increases in fresh weight,
was double SIgmoid (Fig. 2A) - similar to that shown
by drupes (Crane 1964). This growth curve can be
divided into three stages: Stage I (O-S8 days after
flowering), in which fruit weight and volume increase
rapidly, was followed by a period of slow growth
Hopping-Structure &. development of fruit &. seeds in Chinese gooseberry 6'
J1&. 1 Radial transverse sec-
tioJU through Chinese loose-
A
berry fruits - A, whole fruit; D,
diasramatic representation of a
IODJitudinal section throu,h a
mature fruit; C. epidermis and
outer pericarp 93 days after
ftowerin,; D. outer pericarp 23
days after flowering; E. inner
poricalP 12 dar after flower-
mat F. centra core 93 da}'ll
after ftowerina.
C - c:entia1 core, B - epi-
cIermia. H - bair, IB - inner
bundle, JP - inner pericarp.
L - locole. 08 - outer bundle,
OP - outer pe#carp, P - pIa-
ClIOtae, S - Heel, SE - aeptwn.

B
 A
60
Wt.
40

20
DR, Wt.

o
OUT ER -:E RIC AR P
44 B

36

28
CENTRAL CORE

INNER
I
PERICARP

20

30 c
w _
N
_ N
0 20
V')
OUTER \ERICARP

CENTRAL CORE

o
2.4
e
-....
e
I:
1.6

C) ENDOSPERM
Z 0.8
EMBRYO
w
-'
o
o 40 80 120 160
DAYS AFTER FLOWERING
PI,. % Fruit and seed devclopnlcnt in Chinese 8ooseberry. A, cumulative increases in fresh
and dry weight of fruit; B, cell number of the central core and the inner and outer pericarp;
Ct mean cell size of the central core and the inner and outer pericarp; D, cumulative
increase. in the length of the seed, nucellua. endosperm, and embryo.
Hopping-Structure & development of fruit & seeds in Chinese gooseberry 67

(Stage II, 58-76 days after ~owe~ng) and. a further

period of rapid growth until fruIt matunty (Stage
111, 7tH60 days after Rowering). No similar division
A
of the: dry-weight growth curve could be demon-
strated.

Fruit growth during Stage 1 was initially by cell

division in the central core and the inner and outer
pericarp, followed by cell enlargement in aU ti~ue5
(Fig. 2B, IC). Cell division in the outer and Inner
pericarp ceased afler 23 and 33 days respectively, but
continued in the central core, albeit at a slow rate,
until ItO days after flowering. Cell enlargement com-
menced in aU three tissues immediately after flower-
ing anI! continued until the onset of Stage II (44 days
alter lIowering) (Fig. 2C). During this time cells of
the inner pericarp underwent a five-fold increase in
len,th.

Stage II of fruit growth was characterised by a

period of reduced fresh-weight gain and this can be
atCribed to an abrupt slowing of cell enlargement in
both inner pericarp and central core. Subsequently,
fruit wei,ht increased and cell enlargement of the
inner pericarp and central core tis'Iues continued
(Stale III) until the fruit reached maturity. How-
ever, mean cell size of the outer pericarp decreased
slightly during this time.
Faa. 3 Median transverse section!! throup the innct
pericarp of Olineae aooseberry fruit - A. embryo ~8
days after flowering; and B, embryo 93 days after
ftowcril1l.
EM - embryo. EN - endosperm, END - endo-
thelium, H - hypostase, 1 - intelument, M - micro-
pyle, N - nucellus, T .. testa, VT - vaac:utar trace.

At dowerina the anatropous ovule consisted of a Seed deveiopmeat

matul'C embryo sac embedded in a we)J-defincd
nucellua which was bounded by a uniscriate integu-
ment. The cell waUs of the integument became pro-
peuively thickened, especially the inner anticlinal
waU, and the cells became ftlled with dark-stainin&
deposits (Fig. 3A). A cuticle waS present on the
outaide of the integument. The nuceUus eniatrJed by
cen multiplication, and a hypostasc (C~t6 1944,
10hn '963) was present during early seed develop-
ment. The hypostasc. which was more apparent in
~ons of fresh material than in material that had
been embedded in paraffin, consisted of a grid-like
artaIIgCment of elongated nucellar cells. The layer
of celt, surrounding the embryo sac enlal'lcd and
thickened to form a conspicuous endothelium (Fil.
3A). Inside the endothelium a cellular endosperm
developed tlllt eventually underwent multiple divi-
sions to produce I mass of small, thin-walled cells.
After fertilisation the embryo, which wu loc:ated at
the micropvlar end of the embryo sac (Fig. 3A),
diYided and then remained quiescent at the two-
c:eUecf staRe for at least 60 days. Between 60 and 110
days af~r t\oweting the embryo developed (Fig. 38)
to its maximum size and had two conspicuous coty-
ledons which did not absorb aU the enciotpenn.
Seed development. as measured by lrowth in
length. beaan immediately after fertiliaation flnd
~ntinued for nearly 80 days (Fig. 20). During this
time the nuc:eUw reached its maximum size and was,
in turn. replaced almost entirely by the endosperm
and endothelium. 1n contraat to many other fruits
(Crane 1964) the embryo remained in the two-celled
stale during most of the endo&pcrm and nuceUus
growth. About 60 days after ftowering the two-c:eJled
embryo divided to form a spheroid and then de-
veloped rapidly to reach its finalaize at I to days.
DISCUSSION
The double aiamoid arowth curve of Chinese
800seberry fruit ia typical of that shown by drupes
such as the peach (Pr"nus persleD, Conners 1919)
and the sour cherry (PTIInUS emu"" Tu~ey t 934)
and other berries such as the lrape (Yiris vlnl/era.
Coombe 1960). Pratt" R.eid (1974) reported that
the Cbineae 800acberry cv. 'Bruno' had a triple ail-
maid srowth curvo with periods of retarded fruit
68
growth between 63-84 and 119-147 days after
anthesis. The latter period of retarded growth was
evident from fresh weight. length, breadth. and
volume measurements. I found that the onset and
duration of the first period of reduced growth (Fig.
2A, 2C) was similar to that reported by Pratt &
Reid (1974) but I obtained no evidence from either
fresh weight or pericarp cell size for a second period
of reduced growth just before maturity. Possibly the
sampling interval was too great to enable a small
reduction in fruit weight and pericarp cell size to be
detected.
Fruit growth during Stage I was due to an initial
period (0-33 days) of cell division in the pericarp
followed by a period of cell enlargement (Fig. 2B,
2C). This pattern of cell division followed by cell
enlargement is typical of the early development of
drupes such as the sour cherry (Tukey 1934). Cell
division and enlargement in the central core, how-
ever, commenced at flowering and continued until
mid-Stage III. During this time the rate of cell
enlargement, but not of cell division, followed the
double sigmoid pattern of fruit weight gain. Stage III
of fruit growth was almost entirely due to elonga-
tion of the septum cells of the inner pericarp (Fig.
2C). No apparent cell enlargement occurred in the
outer pericarp over this period.
The development of seed tissues in Prunus sp. can
be correlated with macroscopic changes in the fruit
in that the development of the nuceUus reaches a
maximum by the end of Stage I. Subsequently, the
nucellus is absorbed by the developing endosperm
and embryo during Stage II (Tukey 1934) . No
further growth of the seed occurs during Stage III.
In contrast, nucellus and endosperm development in
the Chinese gooseberry occurred concomitantly
during both Stage I and II and although embryo
growth commenced during Stage II it continued un-
til the middle of Stage III.
REFERENCES
CoNNERS. C. H. 1919: Growth of fruits of the peach.
New lersey Agricultural Experiment Station
Annual Report 4(): 82-a.
New Zealand Journal of Botany 14, 1976
COOMBE, B. G. 1960: Relationship of growth and
development to changes in sugars, auxins, and
gibberellins in fruit of seeded and seedless
varieties of Vitis vinifera. Plant Physiology 35:
241-50.
CRANE, JULIAN C. 1964: Growth substances in fruit
setting and development. Annual Review of
Plant Physiology 15: 303-26.
CRETE, P. 1944: Recherches anatomiques sur la semi-
nogenese de I'Actinidia chinensis Planch.
Affinites des Actinidiacees. Bulletin de la
Societe Botanique de France 91: 153-60.
ESAU, K. 1965: "Plant Anatomy". 2nd ed. John
Wiley and Sons, New York. 767 pp.
FLETCHER, W. A. 1971: Growing Chinese goose-
berries. New Zealand Department of Agricul-
ture Bulletin No. 349: 1-38.
GALIGHER, A. E.; KOZl.OFF, E. N. 1964: "Essentials
of Practical Microtechnique'. Henry Kimpton,
London. 484 pp.
lOHRl, B. M. 1963: Embryology and taxonomy. In
Maheshwari, P. (Ed.) "Recent Advances in
the Embryology of Angiosperms". Pp. 395-
444. University of Delhi, India.
LI, HUI-LIN 1952: A taxonomic review of the genus
Actinidia. lournal of the Arnold Arboretum
33: 1-61.
PRATT, H. K.; REID, M. S. 1974: Chinese gooseberry:
seasonal patterns in fruit growth and matura-
tion, ripening, respiration and the role of
ethylene. Journal of the Science of Food and
Agriculture 25: 747-57.
TUKEY, H. B. 1934: Growth of the embryo, seed and
pericarp of the sour cherry (Prunus cerasus)
in relation to season of fruit ripening. Pro-
ceedings of the American Society for Horti-
cultural Science 31: 125-44.
VAN TIEGHEM, P. 1899: Sur les genres Actinidie et
Sauravie consideres comme types d'une farnille
nouvelle les Actinidiacees. Journal de Botan-
ique 13: 170.
VIJAYARABHAVAN, M. R. 1963: Embryology of Actin-
idia polygama Franch. and Sav. and the sys-
tematic position of the family Actinidiaceae.
Proceedings of the 50th Indian Science Con-
gress (Delhi). Pt. III. Abstract p. 393.