全 文 ：Discussion on the Problem of Salt Gland of Glycine soja
ZHOU San , ZHAO Ke_Fu＊
(Institute of Plant Stress , Shandong Normal University , Jinan 250014 , China)
Abstract:　Glycine soja Sieb.et Zucc.plants living in saline soil in three provinces of China were treated
with different salinity concentrations under different laboratory culture conditions(including solution , sand and
field cultivation).The attachment shape and distribution on the surface of stalk and leaf of G.soja plants
were observed with scanning electron microscopy (SEM), and the ultrastructure of glandular hair with trans-
mission electron microscopy (TEM).Na+ and Cl- contents in the secretion of the leaf surface and inside the
leaf of G.soja subjected to different treatments were measured.The Na+ relative contents in glandular cells ,
epidermal cells and mesophyllous cells of leaves under different salinities were determined by X_ray microanal-
ysis.Results show that only glandular and epidermal hair exist on the surface attachments of leaves and stalks
of G.soja plants.These glandular hair were similar in shape to some salt glands of Gramineae halophytes ,
and they attached to the vein on the leaf surface.The cell structure of the glandular hair showed the character-
istics of common salt glands , such as big vacuoles , dense cytoplasm , a great deal of mitochondria , chloro-
plast , plasmodesmata and thicker cell walls , etc.The results of Na+ and Cl- contents in the leaf secretion
and inside the leaf showed that the glandular hair executed the function of salt_secretion , and when treated with
the salt gland inhibitor the salt_secretion process was inhibited.As a result , Na+ and Cl-were mainly accu-
mulated inside G.soja leaves.The results of Na+X_ray microanalysis under different salinities proved that
the three cells of the glandular hair , especially the top cell , possessed strong competence for Na+ accumula-
tion.Above all , the glandular hair were the salt gland , and no other kind of salt glands were found on G.soja
plants.The secreting mechanism of the salt gland was also discussed.
Key words:　Glycine soja;salt gland;ultrastructure;ion compartmentalization
　　Glycine soja Sieb.et Zucc.is awild relative species
of G.max.It possesses many useful characteristics and
has become an ideal material in G.max breeding(Zhuang , 1999).There are rich G.soja resources in
China.Recently , G.soja has been drawing more and
more attention , and some important achievements have
been attained(Zhuang , 1999), but very few studies have
been made on the salt_tolerance of G.soja till now.A
research report “Salt glands in G.soja L.China” , pub-
lished in Chinese Science Bulletin (Lu et al , 1998), es-
pecially attracted our interest.However , we noticed that
the discovery of the salt gland of G.soja plant was only
based on observing the shape of this gland , but its salt_
secreting function was not yet proven.In addition , we no-
ticed that the salt gland described and pictured in the
above report was actually a kind of intro_secreting salt
gland—salt bladder (Fahn , 1979;Zhao and Li , 1999).
However , it was different from salt bladders usually re-
ported since it possessed a secretory pore on the ball cell
of the salt bladder , so it is necessary to clearly re-study
the salt gland of G.soja.In this study , materials of G.
soja were collected from the salina in the estuary of the
Yellow River in Kenli County , Shandong Province of
China , as stated in Lu et al s report(1998), and other
materials of G.soja growing in saline soil were collected
from Jiangsu Province and Liaoning Province of China.In
the study of the shape of the epidermal attachments of
leaves and stalks of G.soja , the same specimen_making
and observing method was as Lu et al s report(1998).
1　Materials and Methods
1.1　Plant materials and treatments
Glycine soja Sieb.et Zucc.seedlings(identified by
Professor LI Fa_Zeng , Institute of Plant Stress , Shandong
Normal University)and seeds were collected respectively
in spring and autumn from the salina in the estuary of the
Yellow River in Kenli County , Shandong Province ,
China.Seeds from Jiangsu Province and Liaoning
Province were provided by the Agronomy Department of
Nanjing Agricultural University and the Institute of
Shenyang Applied Ecology , respectively.Seeds were ster-
ilized with 1%HgCl2 , washed with running water , plant-
ed in a seedbed after their coats were scarified for easy
absorption of water.Seedlings uniform in growth after ger-
mination were selected and cultured in Hoagland solution
with different salinities(0 , 50 and 100 mmol/L NaCl re-
spectively)under sand culture condition , at 22-30 ℃
and 80% humidity.At different time intervals of plant
growth and development , the shape , structure and physi-
ological function of the epidermal attachments of G.soja
plants were investigated.
1.2　Observation of the shape and distribution of the
Received:2002-06-28　Accepted:2003-02-17
Supported by the State Key Basic Research and Development Plan of China(G1999011700).
＊Author for correspondence.Tel:+86(0)531 6187879;Fax:+86(0)531 6180397;E_mail:.
植　物　学　报　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　
Acta Botanica Sinica　2003 , 45(5):574-580 http://www.chineseplantscience.com
glandular hair of G.soja plants
G.soja plants , growing naturally and cultivated in
the laboratory , were used.Fresh materials of stalk , leaf
and leaf stalk of different positions on the plants were ob-
served and photographed under Hitachi S_570 scanning
electron microscope (SEM).The number and the distri-
bution density of epidermis hair and glandular hair were
determined , and their sizes were measured.
1.3　Observation of the ultrastructure of the glandu-
lar hair of G.soja plants
The fresh leaf of G.soja was cut into 0.2 cm×0.2
cm pieces , fixed 2 h with 2.5% glutaraldehyde at room
temperature , air_freed and fixed for 1 d with 2.5% glu-
taraldehyde at room temperature.The material was then
washed with 0.1mol/L phosphate buffer solution , fixed 2
h with 1%OsO4 and washed again with 0.1mol/L phos-
phate buffer solution , before being dehydrated in an alco-
hol series concentration (30%, 50%, 70%, 80%,
95% alcohol for 20 min and in 100% alcohol for 3 h).
After an infiltration for 24 h with Epon 812 resin , samples
were embedded in the same resin , which was polymerized
for 24 h at 30 ℃, 45 ℃, 60 ℃, respectively.The ma-
terial was sliced with a glass knife on ULTRACUT_E
ultramicrotome to a thickness of 70 nm.The slice was
stained with uranium acetate and lead citrate , and was
observed and pictured with Hitachi H_800 transmission
electron microscope (TEM).
1.4　Determination of Na+and Cl- contents in the
secretion of the leaf surface and inside the leaf of G.
soja under different treatments
Seedlings , similar in size , were selected.Ten com-
pound leaves (30 leaflets)on the same position of each
plant were harvested.The surface of each leaflet was
thoroughly washed with distilled water.Seedlings were di-
vided into three groups and treated as follows:the 1st
group was continually cultured in Hoagland solution , the
2nd group was cultured in Hoagland solution supplement-
ed with 100 mmol/L NaCl , and the 3rd group was treated
as in the record group but the surfaces of chosen leaves
were sprayed with a salt gland secretion inhibitor—100
mmol/L choline benzenesulphonate (Larkum and Hill ,
1970).There were at least three replicates in each treat-
ment.The chosen leaves were washed again with a fixed
volume of distilled water after 72 h , and the washing so-
lution was preserved for further analysis.Afterwards , the
10 compound leaves that had been washed were chopped
up , dried , weighed , ashed , dissolved with concentrated
HNO3 , and the solution volume was adjusted with dis-
tilled water for Na+ and Cl- analyses.
The Na+ and Cl- contents in the washed solution
and dissolved solution were determined using an atom ab-
sorption spectrophotometer (Hitachi Z_8000)and the ni-
trate titration method , respectively.The ion secretion
quantity of the glandular hair on the leaf surface and the
ion content inside the leaf after 72 h treatment were calcu-
lated in each unit of leaf weight.
1.5　X_ray microanalysis
Sample preparation followed Fritz(1989)and Li and
Fritz(1990).Pieces(5 mm ×5 mm)of leaves of G.
soja plants were cut from the middle of the leaves near the
midrib.The tissue samples were put into small cages of
aluminum gauze and plunged into liquid N2_cooled
isopentane and propane (V/V=l/2)for rapid freezing
and freeze_drying.Modified T_shape valves were used for
infiltration with diethyl ether under vacuum at 27 ℃ for
24 h after freeze_drying.Samples were then reduced into
small pieces suitable for cutting with an ultramicrotome
and infiltrated with styrene_methacrylate , transferred into
gelatine capsules and polymerized for 7 d at 60 ℃.Em-
bedded materials were sectioned with ultramicrotome in
the dry state.Slices , 1 μm in thickness , were coated
with carbon.
Sections were examined under an HITACHI_H800
transmission electron microscope equipped with an EDAX_
9100 energy_disperse X_ray analyzer.The accelerating
voltage was 120 kV , the take_off angle 25°.The counting
time for all analyses was 60 live seconds.The data were
expressed as counts per second(CPS)at an element peak
after subtraction of the background.For each tissue com-
partment , seven measurements per section were carried
out.
2　Results
2.1　Shape and distribution of glandular and epider-
mal hair on the surfaces of leaves and stalks of G.
soja plants
As shown in both Fig.1 and Table 1 , the glandular
hair were protrusive on the surface of stalk and leaf ,
which appeared as a short round stick shape , 35-45μm
high , with a diameter of 17.0-18.5μm in the broadest
site and 10-12μm in the narrowest site.The glandular
hair were distributed on the surface of leaves , leaf stalks
and stalks , and their shapes were similar , but their dis-
tribution densities were different.On leaves , more glan-
dular hair were on the undersurface and fewer on the up-
persurface , but all the glandular hair of both sides were
attached to the veins:more on thicker veins and fewer on
thinner veins.Secretions could be observed on some se-
creting glandular hair.The distribution densities of glan-
dular hair on new leaves and aged leaves were different;
there were more glandular hair on new leaves but fewer on
the aged ones.In addition , there were many epidermal
hair on the surfaces of leaf and stalk , which were 95-
550μm long and were unevenly distributed on the stalk ,
leaf stalk , uppersurface and undersurface of leaves.
2.2 　Ultrastructure of glandular hair of G.soja
plants
By TEM , it was found that the glandular hair were
composed of two to three cells , among which the top and
basic cells were bigger , the middle cell was smaller , and
part of the basic cell was embedded in the epidermal cell
layer.There was a large vacuole in the top and basic
cells , and a small cytoplasm clung to their cell walls.
ZHOU San et al:Discussion on the problem of salt gland of Glycine soja 575　
Fig.1.　Shape and distribution of salt glands of Glycine soja plants.
A.The salt glands on tender leaf , ×1 000.B.The secreting salt gland on tender leaf , ×1 200.C.The salt glands on old leaf , ×500.D.
The salt glands on the undersurface of leaf , ×100.E.The salt glands on the uppersurface of leaf , ×100.F.The salt glands on stalk ,
×100.(The open arrows point to the glandular cell , and the solid arrows point to the epidermal hair).
576　 植物学报　Acta Botanica Sinica　Vol.45　No.5　2003
Table 1　The distribution density of glandular and epidermal hair on the surface of leaves , leaf stalks and stalks of Glycine soja plants(number
of glandular and epidermal hair/8.65×10-3 cm2)
Posi tion Leaf uppersurface Leaf undersurface Leaf stalk Stalk
Glandular hair 16.3±3.3 43.6±5.1 31.5±4.9 34.6±6.7
Epidermal hai r 51.6±4.9 103.8±11.2 165.6±22.0 180.7±19.1
Each value is mean±SE.
The cytoplasm in the middle cell was denser and there was
one more vacuole.A thicker cuticular plate could be seen
on the whole outer layer of the three cells , and there were
some protuberances on the cuticular plate of the top cell.
Many mitochondria and chloroplasts were observed in the
three gland cells , and many plasmodesmata in the cell
walls among the top , middle and basic cells(Fig.2).
2.3　Na+ and Cl- contents in the secretion of the
leaf surface and inside the leaf of G.soja
Table 2 shows that the Na+ and Cl- contents in the
leaf secretion and inside the leaf of G.soja plant in con-
trol are much lower than those in the NaCl treated ones.
Under 100 mmol/L NaCl treatment condition , the Na+
and Cl- contents in the secretion of the leaf surface in-
creased distinctly.It also increased to a certain degree in-
side the leaf of G.soja plant.But , when the salt gland
inhibitor was used , the Na+ and Cl-secretion of the leaf
surface was inhibited:Na+ and Cl- contents in the leaf
secretion was much fewer , while that inside the leaf in-
creased significantly.
2.4　Change of Na+ relative content in glandular ,
epidermal and mesophyllous cells of G.soja leaves
under different salinity conditions
As illustrated in Table 3 , the Na+ relative content
in cytoplasm and vacuoles of glandular cells , epidermal
cells and mesophyllous cells of leaves of G.soja plants
increased with salt concentration treatment.Under 50
mmol/L NaCl , the Na+ relative content in cytoplasm and
vacuoles increased step by step from the basic to the top
gland cells:it was higher in the top gland cells , whose
Na
+
relative contents in cytoplasm and vacuoles had re-
spectively increased by 29.6 and 51.4 times than that in
the control.Meanwhile , the Na+ relative contents in cy-
toplasm and vacuoles of epidermal cells and mesophyllous
cells augmented as well , but in a small margin.Under
NaCl 100 mmol/L treatment , the Na+ relative contents in
the cytoplasm and vacuoles of top glandular cells in-
creased further up to 37.0 and 66.8 times than that in the
control.It also rose up in cytoplasm and vacuoles of epi-
dermal cells but within a smaller margin.No apparent in-
crease was noticed in mesophyllous cells.Results also
showed that the Na+ content in vacuoles was higher than
that in cytoplasm , indicating that Na+ was mainly com-
partmented into vacuoles.
3　Discussion
By observing the surfaces of leaves and stalks of G.
soja specimens from three provinces of China (including
the same site as Lu et al s report , 1998)and in different
developmental stages of G.soja plants , the salt gland
described in Lu et al s report(1998)could not be found
in this study , but glandular and epidermal hair were
Table 2　The Na+ and Cl-contents(μmol/g FW)in the secretion of leaf surface and inside the leaf of Glycine soja plants
Treatments Control 100mmol/ L NaCl Inhibitor+100 mmol/L NaCl
Leaf Na+ 0.29±0.09 8.08±0.12 15.65±0.36
Leaf secretion Na+ 0.10±0.01 5.02±0.19 0.13±0.01
Leaf Cl- 0.32±0.05 9.18±0.23 16.90±0.28
Leaf secretion Cl - 0.16±0.02 4.46±0.17 0.17±0.01
Each value is mean±SE.
Table 3　Na+ relative content in glandular , epidermal and mesophyllous cells of Glycine soja leaves under different salinity conditions by
X_ray microanalysis
Treatments Control 50mmol/ L NaCl 100 mmol/ L NaCl
Gland top cell Cytoplasm 2.44±0.36 72.28±9.44 90.37±4.97
Vacuole 2.05±0.20 107.45±9.55 138.99±5.65
Gland middle cell Cytoplasm 2.67±0.54 31.19±2.87 45.49±4.21
Vacuole 2.39±0.43 46.92±1.71 58.45±4.72
Gland basic cell Cytoplasm 2.64±0.38 29.74±1.46 38.09±2.15
Vacuole 2.95±0.15 38.56±1.96 44.90±1.18
Epidermal cell Cytoplasm 1.76±0.05 13.92±2.26 19.04±1.12
Vacuole 1.78±0.09 23.16±3.65 30.05±3.32
Mesophyllous cell Cytoplasm 1.60±0.04 10.64±1.94 10.14±1.33
Vacuole 1.90±0.10 21.63±1.81 23.03±3.71
The datum unit i s cps(count per second).Each datum is mean±SE(n=7).
ZHOU San et al:Discussion on the problem of salt gland of Glycine soja 577　
Fig.2.　Ultrastructure of salt glands of Glycine soja plants.
A.The whole salt gland of G.soja plants , showing the top , middle and basic cell , ×1 500.B.The cuticularized cell wall of the top cell ,
showing the protuberantia and mitochondria , ×9 000.C.The mitochondria of the top , middle and basic cell , ×9 000.D.The cell wall be-
tween the middle and basic cell , showing the plasmodesma , ×12 000.E.The cell wall between the middle and top cell , showing the plas-
modesma, ×20 000.F.The chloroplast of the top cell , ×9 000.
found in different G.soja specimens(Fig.1).The salt
gland described and pictured in the above mentioned re-
port was a kind of typical salt bladder , which was com-
posed of a large number of ball cells on the upper side
and a small handle cells at the base.It is well known that
the salt bladder is an easy_broken structure and its salt_
secreting manner is exactly through breaking themselves(Zhao and Li , 1999).In Lu et al s specimen_making ,
salt bladders could not be observed because they should
have been broken.Furthermore , it was incredible that
there was a secretory pore on the salt bladder (shown in
Fig.3 of Lu et al s report , 1998).Regardless of whether
the salt gland as Lu et al discovered exists or not , it was
worthwhile to study the interesting question of the salt
gland of G.soja.
Glandular and epidermal hair on the surface of
leaves and stalks of G.soja plants were found in this
study.According to the common laws of plant morpholo-
gy , epidermal hair could not possess a secretory function.
By studying the shape , ultrastructure and function of the
glandular hair , which were discovered in this study , it
was proven that the glandular hair was the salt gland of
G.soja.This assumption can be justified by the follow-
ing reasons:(1)The shape of the gland resembled hair_
like salt glands of some Gramineae species , which was
composed of two to three hair_like cells (Luttage , 1971;
Waisel ,1972;Liphschitz and Waisel , 1974;1982;Zhao
and Li , 1999);the ultrastructure of glandular cells
showed the characteristics of common salt glands(Waisel , 1972;Liphschitz and Waisel , 1974;1982;
578　 植物学报　Acta Botanica Sinica　Vol.45　No.5　2003
Flowers et al ,1977;Drennan et al ,1987;Vassilyer and
Stepanova , 1990;Zhao and Li , 1999), such as large
vacuoles , dense cytoplasm , a great deal of mitochondria ,
chloroplast , plasmodesmata and thicker cell walls , etc.;
on the leaves the gland hair were exclusively attached to
the leaf vein , which was probably for easy collection of
salt from the vascular bundle(Figs.1 ,2;Table 1).(2)
The results of Na+ and Cl- contents in the leaf secretion
and inside the leaf showed that this gland functioned for
salt_secretion , and , when treated with salt gland in-
hibitor , its salt_secretion was inhibited while Na+ and
Cl
-
accumulated inside the leaf (Table 2).(3)Results
of Na
+
X_ray microanalyses under different salinity treat-
ments proved that the Na+ relative contents in the three
gland cells , especially in the top cell , were much higher
than those in other cells , suggesting that the glandular
cells possessed a strong competence for Na+ accumulation(Table 3).
Results also showed that the content and distribution
of salt ions in the leaf and stalk tissue of G.soja plants
under salinity condition could be regulated by
salt_secretion of the salt gland.The lower salt ion content
in the functional tissue (such as mesophyll)of G.soja
plants was maintained(Tables 2 and 3), and the physio-
logical function(such as photosynthesis of mesophyll)of
G.soja plants could continue normally because the salt
injury was avoided.
According to the ultrastructure andNa+relative con-
tent of the salt gland , it was deduced that the basic cell
was a collecting cell and the top cell was a secreting cell.
Na+was collected by the basic cell , then transported into
the middle cell , and finally into the top secreting cell.
When the Na+ content reached a certain level , it was se-
creted out of the secreting cell by hydrodynamic pressure(Arisz et al , 1955).The mechanism of salt secretion was
based on the following hypothesis(Fig.3)set up by Poli-
jakoff_Mayer and Gale (1975):
　　　　　　　　　　Symplast　　　　　Symplast　　　Hydrodynamic pressure
Salt ion of leaf tissue······※Collective cell······※Secreting cell※Out of secreting cell　　　　　　　　　　　　　　Active process　　　　Active process　　　　Active and physical process
Fig.3.　Salt secretion mechanism of leaves(Polijakoff_Mayber A and Gale J , 1975).
　　Some researches have indicated that salt gland cells
could transport salt ions against concentration gradient ac-
tively from their neighbor cells and the salt ion content in
their vacuoles could reach a high level for secretion(Arisz
et al , 1955;Berry , 1970;Pollak and Waisel , 1970;
Polijakoff_Mayer and Gale , 1975).Results of Na+ rela-
tive content by X_ray microanalysis showed that the salt
gland cells of G.soja possess strong competence for Na+
compartment(Table 3), which could probably be decided
by the structural characteristics of the salt gland cells.
Many mitochondria of these cells could provide energy for
ion transport , and many plasmodesmata in the cell wall
between gland cells could be the channels of ion transport
(Fig.2).Other researches have indicated that Na+ com-
partment within cells of higher plants was through Na/H
antiporter , which functioned with the energy produced by
H+_ATPase and PPase on the tonoplast(Barkla and Pan-
toja , 1996;Hasegawa et al , 2000).Further studies on
the mechanism of ion transport on the tonoplast of the
G.soja salt glands are needed.
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(Managing editor:WANG Wei)
关于野大豆盐腺问题的探讨
周　三　赵可夫＊
(山东师范大学逆境植物研究所 , 济南 250014)
摘要:　以中国 3 个省的盐生野大豆(Glycine soja Sieb.et Zucc.)为材料 ,在沙基培养 、溶液培养和大田种植 3 种种植
条件下用不同浓度的盐处理 ,观测了茎叶表面附着物的形态分布和腺毛的超微结构 , 测定了叶片腺毛分泌物中和
叶片组织内部Na+和 Cl-的含量变化 ,并对腺毛的 3 个细胞以及表皮细胞和叶肉细胞内的 Na+相对含量变化进行
了X 射线微区分析。结果发现:盐生野大豆茎叶表皮上生长的附着物中只有表皮毛和腺毛 , 腺毛的形态类似于禾
本科植物中的一些盐腺 ,叶片上的腺毛均生长在叶脉上;腺毛细胞内部结构具有一般盐腺的特点 , 如有大液泡 , 稠
密的细胞质 ,大量的线粒体 、叶绿体 、胞间连丝以及较厚的细胞壁等。通过测定在无盐对照 、盐处理和盐处理加盐
腺泌盐抑制剂条件下盐生野大豆叶片腺毛分泌物中和叶片组织内部的 Na+和 Cl-含量 , 结果显示 ,盐生野大豆腺毛
具有泌盐功能 ,加入泌盐抑制剂后 , 其泌盐作用停止;腺毛的 3 个细胞以及表皮细胞和叶肉细胞内的 Na+在不同的
盐浓度下的微区定位分析结果表明 ,盐生野大豆叶片的腺毛细胞有较强的积累 Na+的能力。综合分析认为 ,盐生
野生大豆的腺毛就是具有泌盐功能的盐腺 ,没有发现其他类型的盐腺。
关键词:　野大豆;盐腺;超微结构;离子区域化
中图分类号:Q945　　　文献标识码:A　　　文章编号:0577-7496(2003)05-0574-07
收稿日期:2002-06-28　接收日期:2003-02-17
基金项目:国家重点基础研究发展规划项目(G1999011700)。
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580　 植物学报　Acta Botanica Sinica　Vol.45　No.5　2003