全 文 ：黑果腺肋花楸正常苗与玻璃化
苗茎叶显微结构的比较
高晔华 1，利爽 2，吕天舒 2，胡婷婷 3，吴荣哲 2*
（1.延边林业科学研究院，吉林延边 133001；
2.延边大学农学院，吉林延边 133000；3.延吉
市农业局，吉林延边 133001）
摘 要 以黑果花楸组培苗为试材，采用石蜡
切片和叶表皮撕取法，比较正常苗与玻璃化苗
茎叶横切面显微结构以及叶表面气孔特征 。
结果表明， 黑果腺肋花楸栅栏组织由 2~3 层
细胞构成，下表皮气孔下陷，小而密集，茎内分
布大量导管、筛管。玻璃化苗主脉不明显，细胞
排列凌乱不规则，栅栏组织与海绵组织的区别
不明显，气孔开口大近圆形，茎肿胀粗大髓腔
内薄壁细胞较多、已破裂。
关键词 黑果腺肋花楸；形态显微结构；正常
苗；玻璃化苗
作者简介 高晔华（1987-），女，内蒙古乌察布
人，助理农艺师，研究方向：园艺植物生物技术。
* 通讯作者，副教授，从事园艺植物生理及生物
技术方面的研究，E-mail：wurzh@ybu.edu.cn。
收稿日期 2014-08-18
修回日期 2014-09-30
Comparison of Morphological Microstructure of
Vitrified and Normal Shoots of Aronia
melanocarpa Elloit
Yehua GAO1, Shuang LI2, Tianshu LU2, Tingting HU3, Rongzhe WU2*
1. Yanbian Forestry Research Institute, Yanbian 133001, China;
2. Agricultural College of Yanbian University, Yanbian 133000, China;
3. Yanji Bureau of Agriculture, Yanbian 133001, China
*Corresponding author. E-mail: wurzh@ybu.deu.cn
Received: August 18, 2014 Accepted: September 30, 2014A
Agricultural Science & Technology, 2014, 15(10): 1667-1670, 1683
Copyright訫 2014, Information Institute of HAAS. All rights reserved Molecular Biology and Tissue Culture
A ronia melanocarpa Elloit(Rosaceae: Sorbus) is a de-ciduous shrub. It is an edible
and medicinal plant with ornamental
and other research value. During the
tissue culture of Aronia melanocarpa
Elloit, some vitrified plants have em-
erged. Vitrification occurs in the cul-
ture process of plant tissue, and it is
a physiological disorder or physiologi-
cal disease that occurs in plant in vitro.
Vitrification is an adaptive response
to stress[1]. It could occur in the tissue
culture of all kinds of plants. Vitrifica-
tion will not only affect the proliferation
and differentiation, but also affect the
growth of plants. The vitrified plants
will be difficult to survive in the trans-
planting. However, most of the vitrified
shoots could be reversed into normal
robust shoots under certain conditions,
such as changing the type of tissue
culture medium, the concentration of
exogenous hormones, the sealing
material of containers, etc [2]. In terms
of morphology, organization anatomy,
physiology and biochemistry, osmatic
balance, the changes of endogenous
hormones, etc, the further analysis in-
dicates that the abnormal internal
structure is the main cause for abnor-
mal external morphology of vitrified
plants, but the mechanism of vitrifica-
tion is not yet fully understood [3-5]. At
the morphology and anatomy, the cor-
tical parenchyma of vitrified plants
is overgrown and irregular; the gap
among adjacent cells is large and the
vessels are developed badly [6]. More-
over, the palisade tissue of leaf blades
has no or less layers of cells; there is
only spongy tissue; the stomas on the
surface of leaf blades open largely;
the guard cells cave in[7-10]. So it is nec-
essary to investigate the difference in
morphological characteristics and mi-
crostructure of leaf blades and stems
between normal and vitrified shoots.
The researches on vitrification in Aro-
nia melanocarpa Elloit have not been
reported, and the researched on
anatomical structure of vitrified Aronia
melanocarpa Elloit have not been re-
ported yet. In this study, the tissue
cultured Aronia melanocarpa Elloit
shoots were used as test material, and
Abstract The Aronia melanocarpa Elloit was used as test material. The microstruc-
ture of normal and vitrified shoots and the characteristics of their stomas on leaf
surface were compared by paraffin section and leaf epidermis-tearing method. The
results showed the palisade tissue of Aronia melanocarpa Elloit consists of 2-3 lay-
ers of cells. The stomas on lower epidermis cave in, and are small and dense.
There are abundant vessels and sieve tubes in stems. In contrast, the main veins
of vitrified shoots are unobvious, messy and irregular. The boundary between pal-
isade tissue and spongy tissue is not obvious. The stomas open circularly and
bigly. The stems are swelling and thick, but the pith parenchyma cells are broken.
Key words Aronia melanocarpa Elloit; Morphological microstructure; Normal shoot;
Vitrified shoot
Agricultural Science & Technology 2014
Table 1 Comparison of leaf tissue microstructure of normal and vitrified shoots
Microstructure Leafthickness∥μm
Vein
thickness∥μm
Palisade tissue
thickness∥μm
Spongy tissue
thickness∥μm Leaf tightness∥%
1st-level normal shoots 116.2 bz 265.2 a 52.3 a 23.6 a 45.0 a
2nd-level normal shoots 124.9 b 268.6 a 54.8 a 21.6 b 43.3 a
3rd-level reversed shoots 121.2 b 239.6 b 43.9 b 19.6 c 36.1 b
4th-level vitrified shoots 150.6 a 203.3 c 38.7 c 18.3 c 26.8 c
5th-level vitrified shoots 161.1 a 181.4 d 33.9 d 15.4 d 22.3 c
Z indicates the result of multiple comparison tests at the 0.05 significance level, the same below.
the difference in morphological char-
acteristics and microstructure be-
tween normal and vitrified shoots
was investigated, providing a refer-
ence for the definition of formation
mechanism for vitrification and pre-
vention of vitrification.
Material and Methods
Material
In order to investigate the mor-
phological microstructure of tissue
cultured Aronia melanocarpa Elloit
shoots, the 40 d aged Aronia
melanocarpa Elloit shoots with same
subcultured cycles were divided into 5
levels according to the specific mor-
phology. 1st-level normal shoots, the
leaves are large, verdant and stretch-
ing with transverse diameter ranging
among 1.1 -1.3 cm. 2nd-level normal
shoots, the leaves are large, verdant
and stretching with transverse diame-
ter ranging among 0.8 -1.0 cm. 3rd-
level vitrified but reversed shoots, the
leaves are moderate in size; the leaf
color is shallower than that of the 1st-
level and 2nd-level shoots; the trans-
verse diameter of leaf blades ranges
among 0.5 -0.7 cm; 3/4 of the whole
plants is normal, but 1/4 of the leaf
blades and lower stems (near the root
end) are soaked-like. 4th-level vitrified
shoots, the leaves are relatively small;
the transverse diameter ranges among
0.3-0.5 cm; the leaves are curling lon-
gitudinally; the leaf blades and the
lower 3/4 of the stems (near the root
end) are soaked-like. 5th-level vitrified
shoots, the leaf blades are small; the
transverse diameter of leaf blades is
about 0.2 cm; the leaves are curling
longitudinally; the leaf blades and
stems are almost soaked-like. All the
above 5 levels of plants were collected
at 10:00 am, and the leaf blades with
the same morphology and size on the
3rd internodes were collected. The leaf
fragments in size of 0.5 cm × 0.5 cm
along both sides of the main vein were
cut, and the stem fragments in length
of 0.5 cm on the 3rd internodes of tis-
sue cultured shoots were cut. All the
sampled fragments were fixed imme-
diately with FAA (V (70% ethanol)∶V
(formaldehyde)∶V (acetic acid)=90∶5∶5)
as the test material.
Methods
The central and edging parts (2/3
of the leaf blades) of the 5-level leaf
blades of 40 d aged Aronia
melanocarpa Elloit shoots with same
subcultured cycles were cut with size
of 0.5 cm × 0.5 cm. The stem frag-
ments in length of 0.5 cm on the 3rd
internodes of tissue cultured shoots
were cut. There were 10 repeats for
each treatment. The fixation was last-
ed for as long as 24 h for making the
test material sink to the bottom.
Data analysis
All the tests adopted the random-
ized complete design. The test data
was processed with Excel 2003. The
significant difference analysis was
performed by using the Duncan’s mul-
tiple range test of SPSS 11.5. The sig-
nificant level was 0.05.
Results and Analysis
The morphological structure of
leaves, among all the plant organs,
could clearly reflect the ecological
conditions of plants. The leaves of
normal Aronia melanocarpa Elloit
shoots were verdant and stretching,
the stems were erecting and the plants
were robust. The leaf color of reversed
shoots was shallower than that of nor-
mal shoots, but the leaves were
stretching and verdant. The leaves of
vitrified shoots were light green,
soaked-like, swelling and curling lon-
gitudinally, and the plants were dwarf
and small. Under a microscope, the
leaf blades of Aronia melanocarpa El-
loit consisted of epidermis, mesophyll
and veins. The upper epidermis was
made up with a layer of tightly connect-
ed flat cells, while the lower epidermis
was made up with a layer of square
cells and was distributed with stomas.
The palisade tissue consisted with 2 -
3 layers of cells. The spongy tissue
was not developed well . The gap
among adjacent cells was small. The
stems of tissue cultured Aronia
melanocarpa Elloit shoots consisted of
epidermis, cortex and vascular cylin-
der from outer to inner. And the vascu-
lar cylinder was made up with vascular
bundles, pith and pith ray. There were
vessels and sieve tubes distributed in
the stems.
Leaf microstructure of normal and
vitrified Aronia melanocarpa shoots
As shown in Table 1, the leaf
thickness of tissue cultured Aronia
melanocarpa Elloit shoots was in-
creased with the increase of leaf level.
The leaf thickness of vitrified shoots
(4th-level, 150.6 μm； 5th-level, 161.1
μm) was significantly higher than that
of normal shoots (1st-level, 116.2 μm；
2nd-level, 124.9 μm). There was no sig-
nificant difference in leaf thickness be-
tween reversed shoots (121.2 μm) and
normal shoots. The vein thickness of
vitrified shoots was significantly lower
than that of normal shoots with the
vein thickness of 5th-level vitrified
shoots accounting for only 68.4% of
that of 1st-level normal shoots. The
Fig.1-a showed the main veins of vit-
rified shoots were not obvious; the pal-
isade tissue of normal shoots consist-
ed of 2 - 3 layers of cells; the long axis
of cells was vertical to the upper epi-
dermis; the cells were dense (Fig.1
A1, A2). While the palisade tissue of
vitrified shoots consisted of only 1 lay-
er of cells; the boundaries between
palisade tissue and spongy tissue
were not obvious; the gap among ad-
1668
Agricultural Science & Technology2014
Table 2 Microstructure of the upper and lower epidermises of normal and vitrified Aronia melanocarpa Elloit shoots
Microstructure Upper epidermisthickness//μm
Lower epidermis
thickness//μm
Stoma size (longitude diameter ×
horizontal diameter, μm × μm)
Stoma density
n/mm2
1st-level normal shoots 19.9 dz 15.4 c 25.3 × 9.2 c 416 c
2nd-level normal shoots 19.3 d 16.0 c 23.4 × 10.1 c 409 c
3rd-level reversed shoots 23.8 c 18.2 bc 23.5 × 10.3 c 456 b
4th-level vitrified shoots 32.9 b 24.2 b 23.8 × 12.6 b 491 a
5th-level vitrified shoots 39.6 a 35.6 a 25.0 × 14.5 a 502 a
jacent cells was large (Fig.1 A4, A5);
the thicknesses of palisade tissue and
spongy tissue were all lower than that
of normal shoots, and compared to the
thickness of palisade tissue of 1st-level
normal shoots, the thickness of pal-
isade tissue of 5th-level vitrified shoots
was decreased by 35.2%. With the in-
creased extent of vitrification, the mor-
phology of palisade tissue had also
changed. The cylindrical cells were
changed into short cylindrical or
spherical cells, and the tight junction
was changed into loose junction. The
morphological structure of palisade
tissue and spongy tissue of 3rd-level
reversed shoots was similar to that of
normal shoots (Fig.1 a4, a5). At the
same time, with the increased extent
of vitrification, the leaf tightness (CTR)
was decreased. The leaf tightness of
5th-level vitrified shoots was only
22.3%, less than half of that of 1st-level
normal shoots.
Microstructure of upper and lower
epidermises of leaf blades of nor-
mal and vitrified Aronia melano-
carpa shoots
As shown in Table 2, the thick-
ness of upper epidermis of vitrified
shoots (4th-level, 32.9 μm; 5th-level,
39.6 μm) was about 2 times that of
normal shoots (1st-level, 19.9 μm; 2nd-
level, 19.3 μm). The thickness of upper
epidermis of reversed shoots was
about 23.8 μm, lying in the middle of
that of normal and vitrified shoots. The
thickness of lower epidermis of vitrified
shoots (4th-level, 24.2 μm; 5th-level,
35.6 μm) was significantly higher than
that of normal (1st-level, 15.4 μm; 2nd-
level, 16.0 μm) and reversed shoots
(3rd-level, 18.2 μm). The upper epider-
mis of normal and vitrified Aronia
melanocarpa Elloit shoots was all con-
sisted of 1 layer of cells with few
stomas. The upper epidermic cells of
normal shoots were rectangular and
flat, and they connected with each oth-
er tightly with no gap; while the upper
epidermic cells of vitrified shoots were
square and swelling with obvious cel-
lular gap (Fig.1 A4, A5). There were
stomas on the lower epidermis of both
normal and vitrified Aronia
melanocarpa Elloit shoots. The
stomas of normal shoots consisted of
2 kidney-shaped guard cells, which
were similar in size, and the stomas
were elliptical and relatively small
(Fig.2 B5); while the stomas of vitrified
shoots were relatively large, and the 2
guard cells were different in size. With
the increased extent of vitrification, the
opening extent of stomas was in-
creased gradually. The 5th-level vitri-
fied shoots had the largest and nearly
spherical stomas with size of 25.0 μm
(longitudinal diameter) × 14.5 μm (hor-
izontal diameter). The stoma dense of
vitrified shoots (4th-level, 491/mm2; 5th-
level, 502/mm2) was significantly high-
er than that of reversed (3rd-level,
456/mm2) and normal shoots (1st-level,
416/mm2; 2nd-level, 409/mm2), reflect-
ing the stress-adaptability of vitrified
shoots.
Stem microstructure of normal and
vitrified Aronia melanocarpa shoots
Table 3 showed there was signifi-
cant difference in stem thickness be-
tween normal and vitrified shoots. The
cross section diameter of stems of vit-
rified shoots was 891.1 μm, and the
stems were thick and swelling (Fig.3c).
The epidermis of both vitrified and nor-
mal shoots consisted of 2 layers of
cells. The epidermic cells of normal
shoots were full and arranged tightly;
while the epidermic cells of vitrified
shoots were swelling and square. The
epidermis thickness of the vitrified
shoots was 32 μm, which was signifi-
cantly higher than that of normal
shoots. The epidermic cells of normal
shoots were regular; but the cortical
parenchyma of vitrified plants was
overgrown and irregular, the gap a-
mong adjacent cells was large and the
thickness of epidermis was signifi-
cantly higher than that of other shoots.
The epidermis thickness of reversed
shoots was close to that of normal
shoots, indicating their epidermic cell
morphology had gradually returned to
A: Mesophyll, 10 × 40; a: Cross section of leaf blades, 10 × 10; 1, 1st-level normal shoots; 2, 2nd-level normal shoots; 3, 3rd-level reversed
shoots; 4, 4th-level vitrified shoots; 5, 5th-level vitrified shoots.
Fig.1 Morphological anatomy microstructure of leaf blades of tissue cultured Aronia melanocarpa Elloit shoots
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Agricultural Science & Technology 2014
Table 3 Microstructure of stems of normal and vitrified Aronia melanocarpa Elloit shoots
μm
Microstructure Stemthickness
Epidermis
thickness
Cortex
thickness
Pith
thickness
1st-level normal shoots 802.6 bz 22.8 b 101.6 b 263.0 b
2nd-level normal shoots 830.7 b 23.6 b 103.7 b 274.7 b
3rd-level reversed shoots 833.9 b 31.1 a 103.3 b 286.3 b
4th-level vitrified shoots 889.3 a 32.3 a 118.7 a 320.8 a
5th-level vitrified shoots 891.1 a 32.9 a 120.3 a 339.8 a
C: Cross section of stems, 10 × 40; c: Cross section of stems, 10 × 10; 1, 1st-level normal shoots; 2, 2nd-level normal shoots; 3, 3rd-level re-
versed shoots; 4, 4th-level vitrified shoots; 5, 5th-level vitrified shoots.
Fig.3 Morphological anatomy microstructure of stems of tissue cultured Aronia melanocarpa Elloit shoots
B: Stomas on the lower epidermis, 10 × 40; b: Stomas on the lower epidermis, 10 × 10; 1, 1st-level normal shoots; 2, 2nd-level normal shoots;
3, 3rd-level reversed shoots; 4, 4th-level vitrified shoots; 5, 5th-level vitrified shoots.
Fig.2 Morphological anatomy microstructure of stomas on the lower epidermises of tissue cultured Aronia melanocarpa Elloit shoots
normal. Fig.3C showed the vascular
bundles of normal shoots were sepa-
rated, but the spacing was small.
There were abundant vessels and
sieve tubes. The vascular bundles of
vitrified shoots were developed badly
with unobvious boundaries among pith
rays, and there was fault between two
adjacent vessels or sieve tubes, dis-
facilitating the transport of substances.
Discussion and Conclu-
sions
The organ morphology of plants
will be closely adaptive to their physio-
logical function and growing environ-
ment. Under the long-term effects of
external ecological factors, leaves
have the largest variability and plastici-
ty in morphological structure. This
means leaves will make most obvious
response to ecological conditions [11].
The appearance of vitrified shoots was
soaked-like. Their main veins were not
obvious. There were no obvious
boundaries between loose palisade
tissue and spongy tissue. The upper
epidermic cells of tissue cultured Aro-
nia melanocarpa Elloit shoots were ar-
ranged closely. The stomas on the
lower epidermis of vitrified shoots were
small and dense, and they opened
largely and nearly circularly. The
stoma density of vitrified shoots was
significantly higher than that of normal
shoots. The epidermic cells were
messy and irregular. There were de-
veloped vessels and sieve tubes dis-
tributed in the tissue cultured Aronia
melanocarpa Elloit shoots. However,
the stems of vitrified shoots were
swelling and thick; the vascular bun-
dles developed badly; the pith
parenchyma cells were abundant but
broken.
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