Flora 168, 263-27 5 ( 1979)

Developmental and Anatomical Studies in the Pericarp

of Capsicums

Y. S. DAvE, N.D. PATEL and K. S. RAo

Department of Biosciences, Sardar Patel University

Vallabh Vidyanagar, India

Summary
The epicarpic cell walls are beaded and enclose protein and/or oil m the lumens. The 2-4
layered outer hypodermal zone of the multilayered epicarp is either collenchymatous or chloren-
chymatous. The parenchymatous mesocarp is sometimes distinguished into outer and inner
regions of which the latter embed the vascular bundles. The inner hypodermis is a single layer
of giant cells which increase tremendously in their size towards the maturity of the fruit. The
number of parenchyma in a triangular-wedge shaped group with giant cell differs in the various
Capsicums. The solitary or small groups of lignified cells are present among the parenchyma of
endocarp. The placental cells towards the maturity of the fruit show the enlargement of their
vacuoles and intercellular spaces and finally their separation. They contain starch or insoluble
carbohydrates. The detailed study of C. jrutescens (var. Long Red) indicates that there is increase
in number of cell layers and cell size; it also reveals that there are differentiation and speciali·
zation of the cells during the development of the fruit from the ovary. The present paper deals
with the study of two varieties of Capsicum jrutescens and five varieties of C. annuum.

Introduction
It seems upto this date no adequate attention has been focused on the detailed
developmental and anatomical studies of the fruit and thus the information is scanty
in the literature (DAVE and PATEL 1975; KARAWYA et al. 1973; NEUBAUER 1971;
PATEL and DAVE 1976; PATEL et al. 1976; STERLING 1953; ZALA et al. 1976). In
contrast to that of seeds, fruit anatomy is not often used as a taxonomic character
(CuTTER 1971). Fruit studies have much to offer in aiding the anatomical knowledge
of different taxa. When most of the embryological studies have not achieved beyond
the structure of the ovary wall in its early stages before or after the fertilization, a
thorough investigation in the ovary wall after the fertilization and upto the maturity
and ripening of fruit is very important. Thus the present investigation on the de-
velopment and structure of the solanaceous fruits stands its value in the literature
and leads into an interesting avenue of morphological studies of the ovary and the
seed.
Materials and Methods
The materials of Capsicum were obtained from Pochas, Poona (Maharashtra) and Genetic
Division, I.A.R.I. New Delhi-12 and from the botanical garden of our University. The ovaries
from flower buds and open flowers and the fruits at their different developmental stages were
264 Y. S. DAVE, N.D. PATEL and K. S. RAo

Fig. 1

A. A whole fruit and transverdely cut portions of it from termin3l region to extreme tip region
(Capsicum annuum var. Hungarian wax), small white line indicate3 l em scale.
B. Cellular details of ovary wall form middle region (0. frutescens var. Lond Red). X 258.
C. T.S. of ovary form terminal region showing the parietal placentae (0. frutescens var. Long
Red). x40.
D. T.S. of ovary from extreme basal region showing basic vascular supply (C.frutescens var. Long
Red). X 40.
 Studies in the Pericarp of Capsicums 265

collected and measured in length and breadth. Then to facilitate the studies the developing fruits
were cut into basal, middle and terminal parts to fix in FAA (SASS 1958). The histochemical
techniques (JENSEN 1962) were employed for the detection of insoluble carbohydrates, lipids,
proteins (MAZIA et al. 1953), pectin, lignin etc.

Observations
Except Capsicum frutescens (var. Long Red), the study of its other variety and
five more varieties of C. annuum for the sake of brevity are presented only in tabular
form.
The ovary of Capsicum is typically two loculed or 3 -A loculed by false septation
in basal and middle regions with axile placenta, but apically unilocular and with
parietal placenta (LAWRENCE 1960).
l. Capsicum frutescens (var. Long Red) is a very hot or pungent fruited variety,
about 10-12 em. long and 1.5-2 em. thick at base. The fruit is 2-4 locular and
without seeds in the terminal region and extreme tip is chamberless. The cells on
the periphery of the parenchymatous centre of the tip appear very peculiarly arranged
in concentric arcs of periclinally divided cells. Its hypodermal cells are also peri-
clinally divided.
Ovary wall: It is 13-16 cells thick without much difference from its base to
the tip. The outer epidermis is a single layer of isodiametric cells with their cuticle.
The epidermal cells divide anticlinally and the hypodermal cells in 3-4layers appear
periclinally divided. Below the latter there are 7-8 layers of large parenchyma em-
bedding a ring of vascular bundles which is placed more towards the inner hypodermis
made of largest cells. The largest cells are called "Riesenzellen" by AuGUSTIN (1907).
The cells of the outer epidermis, outer hypodermis, mesoderm and the giant cells
of the inner hypodermis show the presence of protein and insoluble carbohydrates
in the PAS reaction (JENSEN 1962). The cells of innermost layer of ovary wall appear
smaller than outer epidermal cells. About 4-6 inner epidermal cells line the inner
tangential wall of each giant hypodermal cell (Fig. 1 B). The inner epidermis and
hypodermis of giant cells continue in dissepiment upto some distance towards the
placental axis.
Placentae: The placental axis at the basal and middle regions is very broad,
towards the terminal region it is narrow and then it splits apart at a higher level,
so that the septae appear free and ovary uniloculed (Fig. 1 C). The tissue in the placen-

E. T.S. of fruit from extreme terminal region showing the closing of gap-Note cutinized cells
at arrow (C. frutescens var. Long Red). X 80.
F. Outer epidermal cell showing the cuticular net in the lumen and thick corrugated cuticle
on outer tangential walls (C.jrutescens var. Long Red). X 960.
(b, basal; m, middle; t, terminal; g, giant cell; e, epidermis; me, mesoderm; ov, ovule; dv, dorsa,l
vascular bundle; pb, placental bundle; vv, ventral vascular bundle).
 Fig. 2
A. Formation of ventilating cleft in outer epidermis (G. jrutescens var. Long Red). X 550.
B. Cellular details of fruit showing elongated inner mesocarpic cells (G. jrutescens var. Long
Red). X 24.
C. Small mound on outer epidermis (see arrow) (G. jrutescens var. Long Red). X 270.
D. Transversely cut portion of fruit G. jrutescens (var. Long Red). X 60.
E. Lignified inner epidermal cells in surface view (G.jrutescens var. Long Red). X 200.
F. Outer epidermis in surface view showing the formation of cleft (G. annuum var. acuminata).
X 120.
(me, mesocarp; pl, placentae; g, giant cell)
 Fig. 3
A. A portion enlarged from-T.S. of pericarp.
B. T.S. of pericarp stained with PAS showing the presence of insoluble carbohydrates (see
bracketed area) (0. annuum var. longum). X 240.
C. Cellular details of endocarp showing lignified cells and presence of starch in giant cell (C.
annuum var. longum). X 310.
D. Outer epidermis in surface view showing the cleft (C.jrutescens var. baccata). X 120.
E. Cellular details of epicarp showing cleft (see arrow) (0. annuum var. Hungarian wax). X 258.
F. T.S. of fruit at extreme tip (0. j1·utescens var. Long Red). X 38.
(ep, epidermis; om, outer mesocarp; im, inner mesocarp; en, endocarp; g, giant cell; s, starch;
sc, sclereid; p, parenchyma cell; c, central narrow chamber)
268 Y. S. DAVE, N.D. PATEL and K. S. RAo

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Fig. 4. Graph showing the mean area of the cells at its different developmental stage3.

A. Epicarpic cells in surface view (measured middle lamella to middle lamella).

B. Epicarpic cells in T.S. (only lumens measured).
C. Outer mesocarpic cells in T.S.
D. Inner mesocarpic cells in T.S.
E. Giant cells in T.S.
F. Endocarpic cells in T.S.
G. Placental cells in T.S.

tae, septa and placental axis are homogenous parenchymatous and embed 2-3
large vascular bundles in the axis and 1-2 small vascular bundles in the septa.
Vascular bundles: There is a basic vascular ring of about 20-23 strands at
the base of the ovary (Fig. 1 D). Of these strands, two large strands after branching
from the ventrals move inward to form the placental axial bundles. Due to further
branching about 30-34 developing vascular strands are found in the basal and middle
regions but 10-16 in the terminal region and 7-8 at the tip of the ovary as the
strands are narrowing towards the tip (Fig. 1 C). From the placental bundles on
braching, the traces for ovules are supplied. The procambial strands in the ovary
wall are differenciating into xylem and phloem. The well developed vascular bundles
are conjoint and bicollateral with endarch xylem in the ovary wall, but placental
vascular bundles are with centrally placed xylem surrounded by phloem.
 Studies in the Pericarp of Cap3icums 269

Table 1. Mean areas of the cells of different zones of the fruit at its different developmental stages ( Capsi-
cum frutescens var. Long Red)

Different zones of the fruit Mean areas of the cells in ttm 2

Stage Stage Stage Stage
1 2 3 4

Epicarp cells 181.1 221.2 548.0 580.0

(only lumens measured)
Epicarp cells 128.8 264.3 456.8
(in surface view)
Outer mesocarpic cells 176.9 195.4 510.3 924.0
Inner mesocarpic cells 210.2 268.1 845.3 1064.0
Giant cells 1368.0 5012.0 15400.0 21640.0
Endocarpic cells 133.2 139.2 292.7 339.5
Placental cells 228.0 272.0 827.3 1010.0

Different zones of the fruit Mean areas of the cells in ttm 2

Stage Stage Stage Stage
5 6 7 8

Epicarp cells 925.0 812.5 790.0 630.0

(only lumens measured)
Epicarp cells 576.9 973.8 1340.0 1380.0
(in surface view)
Outer mesocarpic cells 1269.0 1752.0 2577.0 2906.0
Inner mesocarpic cells 1437.0 2536.0 3262.0 4020.0
Giant cells 31340.0 39500.0 42780.0 44710.0
Endocarpic cells 357.8 385.7 402.8 415.0
Placental cells 1349.0 2313.0 2447.0 2516.0

Developing pericarp: The developing pericarp at stage two is 18-22 cells

thick with considerable enlargement of giant cells. The pericarp is varying in its
thickness from 15-35 cell layers during its development. The whole pericarp some-
times is irregularly lobed and infolded at the tip to form a pore lined by the same
epidermal cells (Fig. 1 E).
Epicarp: The epicarpic cells in stage three are much cutinized and divided obli-
quely or periclinally to form a raised structure (Fig. 20). At the fifth stage of their
development they are slightly more vacuolated and large polygonal or rectangular
in shape. Sometimes the cuticle of epicarp cell is seen splitted. The epicarpic cells
at stage five are much thickened and with narrow lumens. The outer cuticle of epi-
carpic cell is corrugated (Fig. l F) or smooth at some places. At sixth stage the epi-
carpic cells are much vacuolated, enlarged and with thick pitted walls. The discon-
tinuation of cutin indicates the splitting apart of the epidermal cells. At times simul-
taneously with the development of thick walls, this splitting apart of middle lamella
causes the formation of pores (Fig. 2A).

2. Capsicum annuum var. longum
Fmit Frmt broad at base, tappermg
at distal end; flesh is thick.
Epicarp Uniseriate; cells tabular or
columnar; smooth cuticle on the outer
tangential and radial walls;
cell walls appear wavy or beaded;
ventilating clefts absent.
Outer 2-4 layered collenchymatous;
Hypodermis collenchyma are lamellar and
with or without chloroplasts.
Mesocarp 18-24 celled thick. Outer mesocarp
with chloroplasts and form less
intercellular spaces and the inner
mesocarp is with more intercellular
spaces, vascular bundles are more
in basal and middle regions but less
in terminal and tip regions.
Inner Giant cells are bounded on their
Hypodermis radial walls by triangular wedge
shaped groups of 8-10 parenchyma
(Fig. 3B).
Endocarp Thinly cutinized parenchyma alternates
with groups of 2-14 sclereids (Fig. 3C).
Parenchyma walls are sinuous or wavy
contain starch and oil globules.
Sclereid walls are pitted.
Placentae Epidermal cells are polygonal, thin
walled and contain starch. 5-6 vascular
bundles in basal, 2-3 in middle anrl
terminal regions. Placental chambers
large in basal and middle regions but
no chamber at extreme tip.
Table 2
~
3. Capsicum annu~tm var. acuminata 4. Capsicun~ annuurn var. grossa --1
0
Fruits longer than broad with Fruit large, angular and depressect
slightly curved tips; flesh is thin. at base; flesh IS thick.
Uniseriate; smooth cuticle on Uniseriate; cells tabular or
outer tangential walls; cell walls rectangular, striated cuticle on outer
appear sinuous and pitted; tangential walls;
ventilating clefts thick (Fig. 2F). cell walls wavy or pitted;
ventilating clefts thick and irregular.
3-4layered, parenchymatous with- 2-3 layers of lamellar parenchyma.
out intercellular spaces. The cells :<
at basal region of fruit are compressed rn
and in terminal region are polygonal. lj
;...
<
t'l
8-10 celled thick, without distinct Many celled (15-16) thick
zones. Intercellular spaces, chloroplasts in basal and middle regions. ~
and starch present. Vascular Vascular bundle3 are more in the b:13'1l lj
bundles more in the basal and middle and middle regions. "::1
;...
regions but one or none in the extreme >-3
t'l
tip. t-<
~
::l
~
Groups of 2-7 parenchyma. Groups of 4-10 parenchyma.
?i
rn
~
0
Its parenchyma alternates with Its parenchyma alternates with
groups of 3-13 sinuous and pitted groups 2-14 sinuous and pitted
sclereids. Starch is present. sclereids.
Parenchyma thin walled with Thin walled parenchyma embedding
intercellular spaces. Placental 6-10 vascular bundles.
out core of thin walled tabular
or columnar cells. Solitary sclereids
arc rare.
 5. Capsicum jrutescens var. baccata
Fruit Fruits short, red or yellow when
ripened; berry is full of seeds.
Epicarp Uniseriate; cells tabular or rectangular,
cuticle on outer tangential wall.
Cell walls sinous and/or pitted in
terminal and middle region,
but striated and without pits in
basal region; ventilating clefts only
in basal region (Fig. 3D).
Outer I or 2 layered, lamellar collenchymatous
Hypodermis with less thick end walls, chloroplasts
present.
Meso carp 13-14 celled thick in basal and middle
regions. Outer and inner zones
are not distinct.
Inner Groups of 2-10 parenchyma.
Hypodermis
Endocarp Its parenchyma alternate with
groups of 3-14 sinous and pitted
sclereids.
Placentae Placental parenchyma large with
intercellular spaces. Placental
out core with tabular or columnar cells.
6. Capsicum annuum var. abbreviata 7. Capsicum annuum var. Hungarian
wax.
Fruit broad at base, tappering Fruit is conical in shape; waxy
at distal end; yellow or red when yellow to bright red when ripe,
ripe, berry full of seeds. berry is unilocular without seeds.
Uni or multiseriate; irregularly 1-3 layered; outer tangential
cutinized and compressed, cuticle walls cutinized; cuticle extends
smooth and thick on outer tangential on the walls of the inner cell where
walls, cell wall striated and ventilating clefts formed (Fig. 3E);
non-pitted. Ventilating clefts cell walls are straight and without pits. rn
present. ~
0..
(0"
w
4- 5 layered; thinly lamellar 3-4 layered thinly lamellar s·
d-
collenchymatous without chloro- collenchymatous with chloroplasts. ::s--
Cl>
plasts.
"d
Cl>
9-10 celled thick. No differenciation 20-24 celled thick, largely vacuolated ;:!.
Q
go
of outer and inner zones. with intercellular spaces.
~
Insoluble carbohydrates present. Insoluble carbohydrates present. 0
...,
Groups of 9-12 parenchyma. Giant cells are not distinct at som3
~
places of fruit. Triangular wedge
shaped groups of 3-4 parenchyma
"'~-
".::
are few. §
Its parenchyma alternate with Completely parenchymatous.
groups of 4-17 pitted sclereids. Polygonal parenchyma are with
Starch is present. straight walls. No sclereids (Fig. 3F).
Thin walled parenchymatous Placental axis broad at base,
with scanty cytoplasm and narrow in the middle and parietal
large intercellular spaces. at the terminal region. Placenta
Outer layer with tabular or absent at extreme tip (Fig. IA).
columnar cells lined by cuticle. Epidermal cells polygonal and thin
walled in surface view. Intercellular
t:-:l
spaces are present. -:J
,_.
272 Y. S. DAVE, N.D. PATEL and K. S. RAo

Outer Hypodermis: At stage two the outer hypodermal cells are anticlinally
and periclinally divided. They are dense with cytoplasm and nuclei and contain
chloroplasts. At stage six 2-4 layers of outer hypodermis are distinctly collenchyma-
tous.
Mesocarp: The inner mesocarpic cells show the presence of insoluble carbo-
hydrates. At the fifth stage the mesocarpic cells have become much vacuolated and
enlarged. Figure B (Fig. 2) shows the abnormally elongated inner mesocarpic paren-
chyma. They appear radially elongated that is at right angle to the axis of the fruit
in its sixth stage. All the branches of basic vascular ring fall in a transversely cut
ring of vascular bundles indicating their vertical extension through the mesocarpic
tissue from base to the terminal region of the fruit.
Inner Hypodermis: The inner hypodermis is made-up of giant parenchyma
cells which are much vacuolated and are longest among the rest of the pericarpic
tissues. They contain insoluble carbohydrates and protein. At the fifth stage they
are enlarged and are empty or with scanty cytoplasm and degenerated nucleus. In
mature pericarp the giant cells measure 4- 6 mm in length and 0.6-0.8 mm in breadth.
These giant cells are bounded on their radial walls by a triangular wedge shaped
group of parenchyma. The numerous bristles seen on the inner surface of the pericarp
are due to the presence of these giant cells. They are characteristic of all the Capsicum
fruits (Fig. 2D).
Endocarp: It develops from the inner epidermis of ovary wall. Its cells are poly-
gonal, straight walled and contain starch, protein and oil. In transection of the peri-
carp small groupt't of 3-6 endocarpic cells, with lignified walls and persistant cyto-
plasm up to the fifth and sixth stage, lie on the lower tangential walls of the giant
cells (Fig. 2E). Later the small groups of 3-12 lignified endocarpic cells called scle-
reids with sinuous and beaded walls, are observed alternately arranged with the rest
of the parenchyma in the endocarp.
Developing Placentae: The cells of the placental epidermis are very small,
vacuolated and polygonal. The solitary, or scattered sclereids are also found in the
epidermis. At the fourth stage the epidermal cells are more vacuolated and elongated.
In the sixth stage endocarp continues on the dissepiment. The placental parenchyma
are with large air spaces and contain insoluble carbohydrates. There are four vascular
bundles in the basal placental axis, which branch and supply to the ovule. The two
of these placental bundles extend into the placental axis reaching the extreme tip
of the fruit.
Growth Pattern: The growth patterns of this fruit are of the simple sigmoid
type for the cells of (i) the pericarp; (ii) the outer meso carp; (iii) the inner meso carp
and (iv) the inner hypodermis (Table 1). When only the lumens of the epicarpic cells
are measured and a graph is plotted, it shows that their lumens increase in size due
to the enlargement of the cells upto the fifth stage of their development. After the
fifth stage the growth curve goes downward indicating the narrowing of the lumens
due to the increment in thickness of their walls (Fig. 4B). The growth curve for the
cells of the placenta is somewhat complicated and appear to be double sigmoid
 Studies in the Pericarp of Capsicums 273

(Fig. 4G). It seems that it has two periods of rapid growth with a period of slow
growth in between i. e.; between the third and the fouth stages. The epicarpic cells
show more increment in their size from stage 5-7 (Fig. 4A). But their lumens be-
come much broader at their third and fifth stages indicating that during the develop-
ment from stages 2-3 and 4-5 there is no much thickening of their cell walls. It
is seen from graph B that the thickening of their walls is increasing from the sixth
stage. The cells of the inner and outer mesocarp show constant increment in their
size. It is more in stage 5-7 (Fig. 4C, D). The cells of the endocarp do not show a
significant increment in their size from their developing stages 3-8 (Fig. 4F). The
major increment in size is during stages 2-3. The placental cells appear considerably
enlarged during the stages 5-6 (Fig. 4G). The giant cells have increased more from
stages 2-3 (Fig. 4E).
The study of C. frutescens, var. baccata and five varieties of C. annuum is tabulated
as follows: (Table 2).

Discussion
A thin cuticle on the outer tangential walls of the outer epidermis of ovary wall
later thickens as the fruit matures. According to JACKSoN and SNoWDON (1968)
some fragments of fruit shmv the strongly striated cuticle but on others striations
are not visible. In seeds a cuticle can develop between the testa and the remains of
the nucellus or endosperm (MARTIN and JuNIPER 1970). The cuticle of the epicarpic
cells of chilli sometimes extends inbetwcen the outer epidermis and subepidermal
layers and even occlude the lumen with cuticular subtance or form a cuticular net.
The epicarp shows beaded cell walls and presence of protein and/or oil in its cells.
JACKSoN and SNoWDoN (1968) reported that the fragments from near the base of
the fruit showed the epicarpic cell walls considerably thickened and distinct pits.
The multilayered collenchymatous or chlorenchymatous outer hypodermis is the
result of frequent anticlinal and periclinal divisions during its development.
The mesocarp rem'1ins parenchymatous throughout the development. Sometimes
the outer and inner mesocarpic regiom can be distinguished. The outer mesocarp
is made up of small parenchyma with or without chloroplasts and the inner meso-
carpi& made up of larger parenchyma embedding the vascular bundles placed more
towards the hypodermal layer of giant cells. The giant cells increase considerably
in their size towards the maturity of the pericarp. According to KAISER (1935) these
interesting giant cells called "Riesenzellen" (AUGUSTIN 1907) begin as meristematic
cells equal in size and similar in shape to the others but grow at a tremendously acce-
lerated rate, so that in the ripe fruits they attain a size several thousand times their
original volume. In our observation the greatest increment in size of the giant cells
of C. frutescens (var. Long Red) is found between stage 2 and stage 3. The total
increment in the area of giant cells at the mature stage is about 33 times than that
at the stage 1. They are bounded on their radial wall regions by the triangular wedge
shaped groups of parenchyma upto certain depth. The giant cells are in direct touch
18 Flora, Bd. 168
274 Y. S. DAVE, N.D. PATEL and K. S. RAo

with one another at the rest of their radial wall regions. Each giant cell is in contact
with the inner epidermis or endocarp on its side facing the chamber of the fruit.
To the side of the giant cell facing the interior of the pericarp, there are attached
parenchyma of mesocarp. FRIDVALSZKY and NAGY (1966) have recognized and de-
scribed that the structure of the giant cells- considering mainly its being foveolated-
is different on the various sides of the cell depending on which tissue and cell type,
respectively, it is in contact with.
KONECSNI (1971) examined two kinds of endocarp cells, the pitted thick walled
sclereids and the parenchyma. The endocarpic parenchyma containing protein,
starch and oil appear wavy and elongated. The endocarpic sclereids with sinuous
and beaded walls are mostly absent at the tip of the fruit, but are found to be com-
pletely absent in C. annuum (var. Hungarian Wax).
The study of growth patterns in the fruit of C. frutescens (var. Long Red) for the
cells of its epicarp, the outer mesocarp, the inner mesocarp and the giant cells reveal
that upto some stages the cells either increase in area andfor in thickness. The pla-
cental cells show two periods of rapid growth increment with a period of slow growth
in between them.

References

ATJGUSTIN, B.: Historisch-kritische und anatomisch-entwicklungsgeschichtliche Untersuchungen

iiber den Paprika. Nemetbogsan, Hungary 1907.
CuTTER, E. G.: Plant Anatomy. Edward Arnold, London 1971.
DAVE, Y. S., and PATEL, N.D.: Anatomy of the fruit wall of Luffa cylindrica (L.) Roem. Vidya
18, 37-48 (1975).
FmDVALSZKY, L., and NAGY, J.: The differentiation, microscopic and submicroscopic structure
of giant cell wall. Acta Agron. Acad. Sci. Hung. 15,69-78 (1966).
JACKSON, B. P., and SNOWDON, D. W.: Powdered Vegetable Drugs. J. & A. Churchill, London
1968.
JENSO::<r, W. A.: Botanical Histochemistry. Freeman, London 1962.
KAISER, S.: The inheritance of a geotropic response in Capsicum fruits. Bull. Torrey Bot. Club
62, 75-80 (1935).
KARAWYA, M. S., HILAL, S. H., and ToHAMY, S. F.: Macro- and micromorphology of fruit of
Nicotiana rustica L. Egypt. J. Pharm. Sci. 14, 163-173 (1973).
KoNECSNI, I.: Contribution to the anatomy of Capsicum annum L. III. Comparative study of
endocarp. Acta Agron. Acad. Sci. Hung. 20, 3-15 (1971).
LAWRENCE, G. H. M.: Taxonomy of Vascular Plants. Macmillan Co., New York 1960.
MARTIN, J. T., and JuNIPER, B. E.: The Cuticles of Plants. Edward Arnold, Edinburg 1970.
MAZIA, D., BREWER, P. A., and ALFERT, M.: The cytochemical staining and measurement of
protein with mercuric bromphenol blue. Biol. Bull. 104, 57-67 (1953).
NEUBAUER, B. F.: The development of the achene of Polygonum pensylvanicum: embryo, endo-
sperm and pericarp. Am. J. Bot. 58, 655-664 (1971).
PATEL, B. R., PATEL, N.D., and DAVE, Y. S.: Pericarpial study in the developing fruit of Cassia
occidentalis L. Flora 165, 295-322 ( 1976).
 Studies in the Pericarp of Capsicums 275

PATEL, N. D., and DAVE, Y. S.: Stomata in the pericarp of Datura innoxia Mill., D. metal L.
and ventilating pores of Physalis minimaL. Flora 165, 61-64 (1976).
SASS, J. E.: Botanical Microtechnique. Iowa State College Press, Ames 1958.
STERLING, C.: Developmental anatomy of the fruit of Prunus domestica L. Torrey Bot. Club
Bull. 80,457-477 (1953).
ZALA, J. V., PATEL, N.D., and DAVE, Y. S.: Structure of the developing and dry pericarp of
Oatharanthus roseeus ( Apocynaceae). Flora 165, 355-360 (1976).

Received May 6, 1978.

Author's Address: Dr. Y. S. DAVE, Department of Bio.'wiences, Sardar Patel University, Vallabh
Vidyanagar 388120, Gujarat, India.

18*