Rice Science, 2014, 21(3): 142149

Copyright 2014, China National Rice Research Institute

Published by Elsevier BV. All rights reserved
DOI: 10.1016/S1672-6308(13)60176-6

Structural and Histochemical Characterization of Developing

Rice Caryopsis
YU Xu-run, ZHOU Liang, XIONG Fei, WANG Zhong
(College of Bioscience and Biotechnology, Yangzhou University, Yangzhou 225009, China)

Abstract: The development of pericarp, seed coat, starchy endosperm and aleurone of the rice
caryopsis was investigated, histochemically and structurally, from the time of flowering to maturity. The
results showed that during its growth, the maximum length of the caryopsis was attained first, followed
by width and then thickness. Histochemical examination of the caryopsis showed that starch was mainly
accumulated in the endosperm, but the endosperm showed no metabolic activity, while embryo and
pericarp contained a few starch grains, and embryo and aleurone were strongly active. Aleuronic cells
contained many aleurone grains and spherosomes, and aleurone in the dorsal region developed earlier
and contained more layers of cells. Amyloplasts in endosperm contained many starch granules and
were spherical at early stages but polyhedric at late stages. The protein bodies appeared later than
amyloplasts, and the number of protein bodies in subaleurone was greater than those in the starchy
endosperm. The white-belly portion of endosperm might be relative to the status of amyloplast
development.
Key words: rice; caryopsis; structure; development; chalkiness

Rice (Oryza sativa) is the most widely consumed

staple food for a large part of the worlds population,
especially in developing countries (Sellappan et al,
2009; Yang and Zhang, 2010). The rice caryopsis
contains the caryopsis coat, aleurone, embryo and
starchy endosperm. Interior to the caryopsis coat is the
aleurone layer, which is tightly bound to the starchy
endosperm. The starchy endosperm, accounting for
more than 80% of the caryopsis, contains parenchyma
cells filled with storage components representing a
major source of food for humanity. The embryo
consists of the scutellum, embryonic axis and various
sheathing structures.
The structural development of the rice caryopsis is
very important in determining rice quality and filled
grain quantity (Wang et al, 1995). The size of the
caryopsis and the number of aleurone layers determine
the quantities of important micronutrients (e.g. iron
and zinc) in the grains (Sellappan et al, 2009). Ebenezer
et al (2001) reported that studies on the structure and
histochemistry of the rice caryopsis had identified
strategies required to enhance grain weight. Whitecore caryopses are larger than translucent ones, owing
Received: 21 June 2013; Accepted: 23 October 2013
Corresponding author: XIONG Fei (feixiong@yzu.edu.cn)

to vigorous initial growth at the ventral area, but with

incomplete packing of starch these kernels are
broken easily during polishing (Tamaki et al, 2009).
The study of the structure of the developing and
mature rice caryopsis has been well documented using
various methods, including histochemical staining and
scanning electron microscopy (SEM) (Wang et al, 2012).
The ultrastructure of the caryopsis coat and the aleurone
has been investigated (Bechtel and Pomeranz, 1977).
The histochemistry and ultrastructure of the rice
embryo have also been studied (Bechtel and Pomeranz,
1978a; Jones and Rost, 1989). The development, structural
changes and storage compounds of endosperm have
been the focus of research for many years (Bechtel
and Pomeranz, 1978b; Ellis et al, 1987; Krishnan et al,
2001; Krishnan and Dayanandan, 2003). Moreover,
caryopsis developments of two varieties with differences
in grain weight and protein content have been compared
(Harris and Juliano, 1977; Wang et al, 2004). However,
knowledge of the sequence of events in the process of
rice caryopsis development is incomplete. This study
describes the changes in caryopsis morphology and
structure during the course of seed development in
rice cultivar Yangdao 6. Based on these primary
measurements, the functional significance of the
caryopsis for seed development was further discussed.

YU Xu-run, et al. Structural and Histochemical Characterization of Developing Rice Caryopsis

MATERIALS AND METHODS

Rice materials
The experiment was conducted at the farm of Yangzhou
University, Yangzhou, China from May 2010 to
November 2012. Rice cultivars Yangdao 6 and
Xiangzaoxian 24 were provided by the Institute of
Yangzhou Agriculture Science, Yangzhou, China and
the Institute of Hunan Agiculture Science, Changsha,
China, respectively. Three seedlings were grown in a
27-cm pot, and pots were filled equally with paddy
field soil. The field soil was sandy loam (typic
fluvaquents), containing organic matter of 24.5 g/kg
and available nitrogen (N), phosphorus (P) and potassium
(K) of 105.0, 33.5 and 66.0 mg/kg, respectively.
Collection of caryopses at different developmental
stages
The flowering times of rice were accurately determined
by dotting the glumes and hanging time-tags on the
rice plants. The representative caryopses of different
stages of development were chosen for the experiment.
Determination of respiratory activity
Cross-sections of caryopses at different developmental
stages were stained by 1% 2, 3, 5-triphenyltetrazolium
chloride (TTC) for 1 h at 25 C. The red color
indicated the existence of dehydrogenase activity,
where respiration was vigorous, with the deeper color
indicating greater vigor.
Determination of starch accumulation

stages for LM specimen preparation. Specimens were

pre-fixed with 2% glutaraldehyde, 1% paraformaldehyde
and 0.05 mol/L sodium methyl mercaptide and then
fixed with 1% Osmium tetraoxide. Sample blocks
were washed and dehydrated via an ethanol series of
30%100%, and then embedded in Spurrs lowviscosity embedding medium. The semi-thin sections
of 1 m thickness were cut with a glass knife on a
Leica Ultrathin Microtome (EM UC6, Germany), and
stained with toluidine blue O for 3 min. The sections
were visualized and photographed with a Leica DM
LS microscopy (Leica, Wetzlar, Germany).

RESULTS
Morphological changes in developing rice
caryopses
The changes in size, shape and color of rice caryopses
from the time of flowering to 23 d after flowering
(DAF) are summarized in Fig. 1. During 17 DAF,
the ovary continually expanded, but developed much
faster in the longitudinal axis and attained full length
in the kernel at 7 DAF. The caryopsis appeared green
and the endosperm was packed with transparent liquid.
During 711 DAF, the caryopsis enlarged in the
transverse axis and attained the maximum width at 11
DAF; the caryopsis became greener and the endosperm
appeared milky-white. During 1317 DAF, the green
of the caryopsis faded slowly and became yellow, and
the endosperm appeared white. During 19 DAF to
maturity, the caryopsis became yellow and hard, and
the endosperm appeared transparent. In addition, the

One or two drops of I2-KI solution were added to a

caryopsis cross-section for 5 min, and the crosssection was then rinsed with water. The black-blue
color indicated the accumulation of starch, with
deeper color indicating more starch accumulations.
SEM observation of caryopsis structure
Mature rice caryopses were fractured by applying
slight pressure on the middle of the caryopsis with a
razor blade. Caryopses with the fractured surface
facing upwards were mounted on a specimen stub and
sputter-coated with gold before SEM viewing (XL30
ESEM, Philips, Holland) at 20 kV.
Light microscopy (LM) observation of caryopsis
structure
Caryopses were harvested at different developmental

143

Fig. 1. Morphological change of Yangdao 6 caryopsis.

Number means days after flowering. Bar = 2 mm.

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Rice Science, Vol. 21, No. 3, 2014

green color persisted longer in the dorsal region of the

caryopsis than in the other regions, indicating more
green cells in the dorsal region, stronger photosynthesis
and a longer duration of grain-filling.
Histochemical changes of starch in caryopsis
From 5 DAF to maturity, the endosperm was stained
increasingly by I2-KI (Fig. 2), which showed that
starch mainly accumulated in the endosperm. The
embryo was stained lightly, indicating that there were
some starch accumulations. The vascular bundle was
not stained, indicating almost no starch. Little starch
was observed in the pericarp at the early stage (15
DAF), maximum starch was observed at about 9 DAF,
less and less starch thereafter. No starch was found in
the mature pericarp.

Fig. 2. Vertical-section of Yangdao 6 caryopsis stained with I2-KI.

Number means days after flowering. Bar = 2 mm.

Change of dehydrogenase activity of developing

caryopsis
Dehydrogenase can be stained by TTC, generating a
red color, which is generally used to represent the
activity of respiration (Wang et al, 2004). The response
to TTC varied in different tissues of the caryopsis (Fig.
3). From 1 DAF to maturity, the embryo was most
deeply stained, showing that it had high activity and
vigorous respiration. The aleurone, lying on the
peripheral endosperm, was stained red throughout,
which also showed that aleurone cells were alive. The
starch endosperm was not stained, showing that its
cells were dead (Wang et al, 2004). The dehydrogenase
activity of the dorsal vascular bundle increased after
flowering, attained the maximum at 12 DAF, and
reduced gradually thereafter. The dehydrogenase activity
ceased immediately when grain filling stopped, which
showed that grain filling was closely related to the
activity of the dorsal vascular bundle (Gu et al, 2001).
Development of pericarp
The caryopsis coat, composed of the pericarp, seed
coat and nucellar layers, surrounds the rice grain
(Bechtel and Pomeranz, 1977). Rice pericarp, the
outermost layer, developed from the ovary wall and
successively divided, moving inward, into exocarp,
mesocarp and endocarp (Fig. 4).
Exocarp is the outermost layer of the pericarp. With
the extension and expansion of the ovary, exocarp
cells elongated longitudinally (Fig. 4-A and -D). At
maturity, exocarp cells were rectangular with thick
walls (Fig. 4-E and -F). Mesocarp is the main part of
the pericarp, and contains 56 layers of parenchyma

Fig. 3. Vertical-section of Yangdao 6 caryopsis stained with

1% 2, 3, 5-triphenyltetrazolium chloride (TTC).
Number means days after flowering. Bar = 2 mm.

cells. By 35 DAF, many starch grains had been

accumulated in mesocarp cells. At 7 DAF, the volume
of mesocarp cells increased; however, at the same
time the mesocarp cells and the starch grains within
began to degrade due to programmed cell death (Fig.
4-C) (Domnguez et al, 2001; Li et al, 2004; Zhou et al,
2009). The spaces, formed by degeneration of cells,
continuously extended. After about 15 DAF, the
starch grains had completely disappeared and the
mesocarp contained only one layer of cells (Fig. 4-E
and -F). Endocarp develops from the innermost
epidermis of the pericarp and consists of cross cells
and tube cells. During caryopsis development, some
inner epidermal cells did not divide and only
elongated longitudinally and so formed the tube cells,
whereas some inner epidermal cells also elongated at
right angles to the long axis of the tube cells and
formed the cross cells (Fig. 4-F).

YU Xu-run, et al. Structural and Histochemical Characterization of Developing Rice Caryopsis

145

number and gathered around aleuronic grains, and the

cell walls became thicker. By 915 DAF, the number
of vacuoles decreased, the numbers of aleurone grains
and spherosomes increased significantly and the cell
walls greatly thickened (Fig. 5-D to -F). After 15 DAF,
aleurone was developed completely.
In addition, cross-sections showed quantitatively
that aleurone varied in dorsal and ventral locations
(Fig. 6). The dorsal aleurone developed earlier and
contained 35 layers of cells, with large cells of
irregular shape (Fig. 6-A); whereas the ventral aleurone
developed slowly and contained only one layer of
cells, with small cells of square shape (Fig. 6-B).
Development of amyloplasts and protein bodies in
starch endosperm

Fig. 4. Micrographs of Yangdao 6 pericarp and seed coat beside

the dorsal vascular bundle.
A, 3 d after flowering (DAF); B, 5 DAF; C, 7 DAF; D, 9 DAF;
E, 15 DAF; F, Beside the lateral vascular bundle at 15 DAF.
DV, Dorsal vascular bundle; Am, Amyloplast; Ep, Exocarp; Me,
Mesocarp; In, Integument; NE, Nucellus epidermis; TC, Tube cell;
CC, Cross cells. Bars = 30 m.

Rice has multiple granular amyloplasts, meaning that

one rice amyloplast contains several starch granules.
At 3 DAF, the cell wall of starchy endosperm was
formed, but no starch was observed in the cells (Fig.
7-A). At 5 DAF, small spherical or elliptical amyloplasts
were formed (Fig. 7-B). With the development of
starchy endosperm, the number and size of amyloplasts
increased. At 15 DAF, the amyloplasts appeared

Development of seed coat

The seed coat, interior to the pericarp, develops from
integument and nucellar epidermis. At 3 DAF, the
integument consisted of 34 layers of cells (Fig. 4-A);
however, at 5 DAF, some layers of cells were
absorbed by the nucellus and only a layer of
integument persisted (Fig. 4-B). At 15 DAF, the
integument and nucellar epidermis were squeezed
together due to the expansion of endosperm and
disappearance of inclusions in ovule and nucellar
epidermis cells (Fig. 4-E and -F), and finally formed
the seed coat. The pericarp and the seed coat were
closely linked and difficult to separate from each other.
Development of aleurone
Interior to the nucellus is the aleurone layer. At 5 DAF,
surface endosperm cells were highly vacuolated and
many spherosomes appeared around vacuoles, and
plastids also initiated (Fig. 5-B). By 7 DAF, the small
vacuoles in aleurone cells were filled with materials
and differentiated into aleuronic grains (Fig. 5-C). At
the same time, spherosomes continued to increase in

Fig. 5. Micrographs of aleurone of Yangdao 6.

A, 3 d after flowering (DAF); B, 5 DAF; C, 7 DAF; D, 9 DAF; E,
11 DAF; F, 15 DAF.
Am, Amyloplast; Al, Aleurone; NE, Nucellus epidermis; PB, Protein
body; Pe, Pericarp; Sp, Spherosomes; Va, Vacuole. Bars = 30 m.

Rice Science, Vol. 21, No. 3, 2014

146

polyhedric and were fully packed with starch grains

(Fig. 7-F).
Protein bodies were initiated at about 7 DAF (Fig.
7-C), started to accumulate at 9 DAF (Fig. 7-D) and
reached the maximum numbers by 11 DAF (Fig. 7-E).
In mature grain, the protein bodies were distorted in
the spaces between amyloplasts, probably owing to
pressure of the enlarged amyloplasts. In addition, the
subaleurone had more protein bodies than the other
regions.
Ultrastructure of mature rice grains
In the mature grains, the morphology and arrangement
patterns of amyloplasts varied in dorsal, ventral and
central regions of the endosperm (Figs. 8 and 9).
Amyloplasts in the dorsal region were well developed,
the starch grains were more polygonal in shape and
compactly arranged, and spaces between granules
were small (Figs. 8-A, 9-B and 9-C). Similar observation
was noted in the central region of endosperm (Fig. 8C). Conversely, many amyloplasts in the ventral
region of endosperm were spherical and arranged
loosely with large spaces between amyloplasts (Figs.
8-E, 9-D and 9-E). In comparison to Yangdao 6, the
starch grains in Xiangzaoxian 24 were more loosely
arranged, and spaces between granules were larger,
resulting in increased chalkiness (Fig. 9).

Fig. 7. Micrographs of starchy endosperm cells of Yangdao 6.

A, 3 d after flowering (DAF); B, 5 DAF; C, 7 DAF; D, 9 DAF;
E, 11 DAF; F, 15 DAF.
Am, Amyloplast; Nu, Nucleus; PB, Protein body. Bars = 30 m.

DISCUSSION
Relationship between structural development of
caryopsis and grain weight
Rice grain weight is one of determining agronomic
traits of crop yield and is relevant to the structural
development of the caryopsis (Takai et al, 2005). In
the present study, compared with Xiangzaoxian 24,
Yangdao 6 had large grains (30 mg/grain) and high
potential yield, possibly due to its stronger physiological
activity and grain-filling, and well-developed pericarp

Fig. 6. Micrographs of aleurone of Yangdao 6 on the dorsal region

(A) and ventral region (B) at 9 d after flowering.
Al, Aleurone; NE, Nucellus epidermis. Bars = 30 m.

Fig. 8. Scanning electron microscopy (SEM) images of dorsal (A

and B), central (C and D) and ventral (E and F) endosperm
in mature caryopsis of Yangdao 6.
Am, Amyloplast; SG, Starch grain.

YU Xu-run, et al. Structural and Histochemical Characterization of Developing Rice Caryopsis

Fig. 9. Scanning electron microscopy (SEM) images of cross-section

(A), dorsal endosperm (B and C) and ventral endosperm (D
and E) of Xiangzaoxian 24.
DE, Dorsal endosperm; VE, Ventral endosperm; Wb, White-belly.

and vascular bundle. Based on caryopsis staining with

I2-KI and TTC, Yangdao 6 had a longer period of
grain-filling and greater metabolic activity or ability to
transport assimilates, which resulted in improved grain
weight compared to Xiangzaoxian 24. The pericarp
plays an important role in caryopsis development (Gu
et al, 2002). As long as the pericarp develops well and
enlarges fully and freely, the endosperm is able to
continuously develop. In this study, a larger number
of starch granules were observed in the cells of ovary
wall during the early stage of caryopsis development;
then, the starch granules disappeared with the endosperm
expansion, indicating that the pericarp supplied the
endosperm growth with nutrients and energy. Moreover,
the dorsal vascular bundle is an important pathway for
photosynthate transferring into endosperm, and thus
its structure and transport ability also affect grain
filling (Gu et al, 2002). In the present report, the vascular
bundle of Yangdao 6 had larger cross-sectional areas,
intact structure and stronger transport activity, which
can allow the caryopsis to absorb more assimilates,
and finally result in increased grain weight.
Relationship between structural development and
grain quality
Rice quality is affected not only by genetic factors but
also by environment and cultivation system (Wu et al,

147

1985; Qu et al, 1991; Lin et al, 2003; Wang et al,

2003). Rice quality is a combination trait, consisting
of milling, appearance, cooking, tasting and nutrient
qualities, which is closely related to structural
development of the caryopsis. If the amyloplasts in
grains were well developed, and of angular polyhedral
appearance, then the rice grains were transparent and
non-chalky, and thus not only had good appearance
but also was not easily broken when milled, resulting
in good milling quality (Guo et al, 1983; Tamaki et al,
2009). In contrast, if the amyloplasts were poorly
developed and of spherical shape with large spaces
between amyloplasts, this resulted in chalky grains of
poor appearance, milling and taste, which gave rise to
decreased market price. In terms of protein, more
accumulated protein bodies lead to a higher nutrient
value but poor taste (Wang et al, 2003).
In this study, SEM results showed that amyloplasts
of caryopses in Yangdao 6 were well developed,
showing crystalloid of polyhedral appearance, and
were compactly arranged with small spaces between
starch grains. We also found that Yangdao 6 had a
longer duration of grain filling (30 d), intact dorsal
vascular bundle and well-developed aleurone compared
to Xiangzaoxian 24. These features showed that
Yangdao 6 had a greater capacity for unloading and
transporting nutrients for endosperm. Slim and long
grains aid transport of nutrients into endosperm,
resulting in decreased chalkiness, increased transparency
and improved appearance, which is similar to the
reports of Jun (1985) and Lin et al (2003). Therefore,
the structure of caryopsis indicates that Yangdao 6 is a
cultivar of high quality and low chalkiness.
Development of endosperm
The endosperm develops from the fertilized polar nucleus
and is composed of aleurone and starchy endosperm.
The aleurone is formed from surface endosperm cells,
while inner endosperm cells differentiate into starchy
endosperm. Most starch in amyloplast and protein
accumulate in the starchy endosperm. The developmental
status of amyloplasts and protein bodies determines
the quality and quantity of rice grains (Yamagata and
Tanaka, 1986; Lopes and Larkins, 1993; Otegui et al,
1999). In present study the development of endosperm
included the development of aleurone and starchy
endosperm.
Causes of chalkiness formation
White-belly or white-core of mature rice caryopses is

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148

an interesting abnormality. High or low temperature

affects the sequence and pattern of enzyme changes
during grain filling, resulting in increased rice
chalkiness (Shen et al, 1997). Over-fertilization or the
competitions for nutrients between grains are also
related to white-belly or white-core (Guo et al, 1983;
Tamaki et al, 2009; Yang and Zhang, 2010). In the
present study, the ventral area of endosperm showed a
loose arrangement of cell content (Fig. 9). We
speculated that white-belly was closely related to the
pathway of nutrients into endosperm. The dorsal
vascular bundle was the main pathway of nutrients
into endosperm. The vascular bundle is not directly
connected with endosperm (Wang et al, 1995). At the
early stage, the dorsal vascular bundle does not
develop fully, and at that time the nutrients for
endosperm cell division and formation of cell walls
come mainly from nucellus cells and the antipodal cell
which would degenerate (Wang et al, 1995). During
grain filling, nutrients from the dorsal vascular bundle
are firstly unloaded in the nucellus projection, and
then there are two ways for nutrients to enter the inner
endosperm (Fig. 9-A) (Wang et al, 1995). One is via
the nucellus epidermis, and nutrients are transported
into the surrounding endosperm, and finally enter the
inner endosperm via aleurone. The other is direct
entry into the endosperm via aleurone near the
degenerated nucellus. Because the belly endosperm is
relatively far from the dorsal vascular bundle, it is
easy to interrupt the final grain-filling and lead to
loose packing of starch granules, thereafter resulting
in endosperm opacity or white-belly (Fig. 9-A, -D and
-E). Therefore, the cause of white-belly or white-core
is earlier development of the ventral area and the
longer distance of nutrient transport.

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