ACTA ECOLOGICA SINICA

Volume 28, Issue 7, July 2008

Online English edition of the Chinese language journal

Cite this article as: Acta Ecologica Sinica, 2008, 28(7), 3228í3235. RESEARCH PAPER

Arsenic tolerance, uptake and translocation by seedlings of

three rice cultivars
Liu Zhiyan1, Chen Guizhu1,*, Tian Yaowu2
1 School of Environmental Science and Engineering, Sun Yat-sen University, Guangzhou 510275, China
2 College of Environment & Planning, Liaocheng University, Liaocheng 252059, China
3 School of Life Science, Sun Yat-sen University, Guangzhou 510275, China

Abstract: By simulating the anaerobic conditions with agar nutrient solutions, effect of arsenic (As) on the growth and As uptake
by hybrid, conventional and glutinous rice cultivars were studied. It showed insignificant effect of As on the root dry weights of
three rice cultivars when treated by As of different concentrations. The shoot dry weights of hybrid and glutinous decreased with As
concentrations increasing, while low concentrations of As (0.5 mg L–1) could enhance the growth of conventional rice. Generally, As
concentrations in roots and shoots increased as As concentrations of treatment solutions increasing. The root system had strong abil-
ity to uptake and accumulate As. The root As concentrations ranged from 156 to 504 mg kg–1, representing 63.40%–81.90% of the
total As concentrations in rice, which were much higher than shoot As concentrations. The fact that the glutinous rice had higher
biomass, higher tolerance, and lower As concentrations in its roots and shoots than the other two rice cultivars proved that the gluti-
nous rice was more applicable to As-polluted soils.

Key Words: cultivar; arsenic (As); hybrid rice; conventional rice; glutinous rice

Rice (Oryza sativa L.) is the most stable food in many re-
gions of the world, especially South-East Asia[1]. The food
safety issue of rice has been much concerned by many scien-
tists. A large number of Arsenic (As) has been put into envi-
ronment through anthropogenic pathways[2,3]. The groundwa-
ter has also faced with As contamination in many countries of
the world[4]. The soils have been contaminated seriously by As
and some other heavy metals in some mines of South Guang-
dong Province, China. As concentrations in field soils around
mines reached 344.11 mg kg–1 according to our soil investiga-
tion recently, which greatly overstepped As concentrations (1–
40 mg kg–1) in uncontaminated soils[2]. Once grown in As-
contaminated soils or irrigated by As-contaminated water, rice
can accumulate high levels of As in roots, straws and grains[5–7].
As accumulates in people’s body through eating As- contami-
nated rice grains, and also As can risk people’s health through
the food chain of straw-livestock-human beings[7].
A lot of studies on rice cultivars have focused on their ag-
ronomic and physiological characteristics[8–10]. Different rice
cultivars have different resistances to environmental adversi-
ties, such as floods, acid rains, plant diseases and insect pests[9,11].
Some reports have shown that different rice varieties (genes)
have different endurances and accumulation abilities to
As[12–14]. However, there are not so many studies about rice
cultivars on As uptake, translocation and accumulation. In this
paper, 3 normal rice cultivars including hybrid, conventional
and glutinous rice in Guangdong Province, South China were
studied to deal with the serious problem of As contamination
in mines, South of Guangdong Province. By simulating an-
aerobic wetland conditions with agar nutrient solutions which
were similar to the practical conditions of rice growing[15–17],
As uptake, translocation and accumulation by different rice
cultivars were studied to provide some scientific supports for
the choice of low-accumulation and high-endurance rice cul-
tivars (varieties).
1 Materials and methods
1.1 Experimental materials

Received date: November 6, 2007; Accepted date: April 18, 2008

*Corresponding author. E-mail: chenguizhu@yeah.net
Copyright © 2008, Ecological Society of China. Published by Elsevier BV. All rights reserved.
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235
The experimental rice varieties are popular in South China,
including 2 hybrid rice varieties: Youyou 998 (YY) and Wu-
fengyou 2168 (WFY), 2 conventional rice varieties: Fengfuz-
han (FF) and Huangsizhan (HS), and 2 glutinous rice varieties:
NuoGW (NGW) and Huangkenuo (HKN).
1.2 Experimental design
(1) Seeds of 6 varieties were surface-sterilized with 30%
H2O2 for 15 min., washed thoroughly with deionized water and
then germinated in moist perlite for 3 weeks.
(2) 1/10 strength of stagnant agar nutrient solution (Hogland
solution) was prepared[18]. The full strength nutrient solution is
composed of macronutrients and micronutrients. The macro-
nutrients (mmol L–1) were: NH4NO3, 5.0; K2SO4, 2.0; CaCl2,
4.0; MgSO4·7H2O, 1.5; KH2PO4, 1.3. The micronutrients (μmol
L–1) were: Fe(II)-EDTA, 50; H3BO4, 10; ZnSO4·7H2O, 1.0;
CuSO4·5H2O, 1.0; MnSO4·5H2O, 5.0; Na2MoO4·2H2O, 0.5;
CoSO4·7H2O, 0.25. To prepare the stagnant nutrient solution,
highly pure agar was mixed in a flask at a rate of 0.1% (w/v)
with the nutrient solution (minus MgSO4·7H2O and KH2PO4)
and heated at 100qC for 1 h. The solution was kept stirring by
a magnetic stirrer during the heating period and when it re-
turned to the ambient temperature, MgSO4·7H2O, KH2PO4 and
micronutrients were added. The pH of agar nutrient solution
was adjusted to 5.5 with 0.1 M KOH or HCl to avoid deposi-
tion[3,16]. Then, 0.5, 1.0, 2.0 and 4.0 mg L–1 As were prepared
by adding Na2HAsO4·7H2O and expressed by As0.5, As1.0,
As2.0 and As4.0, respectively. The control was made without
As and expressed by As0. One PVC pot (10 cm diameter and
13 cm height) contained 500 ml agar nutrient solution with
and without As.
(3) The strong and uniform rice seedlings were selected and
transplanted to the PVC pot containing disposed agar nutrient
solutions (be careful in transplanting rice seedlings to prevent
hurting the root system). There were 6 seedlings in one pot
and 4 replicates in each treatment. The agar nutrient solutions
were replaced once per week. The plants were harvested after
5 weeks’ metal treatment. The growth indexes of rice seed-
lings were determined. The plant samples were thoroughly
washed with tap water, then rinsed with ultra-pure water, and
then separated into shoots and roots. The sub-samples were
oven-dried at 65qC to constant weight for determination and
analysis.
1.3 Indexes and measurements
1.3.1 Measurements of growth indexes
The longest root length of the 6 seedlings from one pot was
measured at the first day after adding metals and the end of
the experiment to determine the longest elongation.
The dry weights of seedlings were determined.
1.3.2 Measurements of As concentrations
Dried plant materials were ground, weighed and put into
dry digestion tubes. Concentrated HNO3 (5 ml) was added and
allowed to stay overnight. On the following day, the tubes
with plant samples were digested at 120–130qC for 24 h. After
digestion the solutions were cooled, diluted with ultra-pure
water, and the As concentrations in acid digests were meas-
ured by an atomic fluorescence spectrometry (AFS-820, Bei-
jing Jitian Analytical Instrument Co., Beijing, China). A blank
and a standard reference of plant material (GBW0763) from
the Department of Earth and Main of China were used to ver-
ify the accuracy of As determination.
1.4 Statistical analysis
(1) Tolerance index (TI)
The tolerance of different rice cultivars (varieties) to As was
expressed by tolerance index (TI), which was quantified by
comparing rates of the longest root elongation in agar nutrient
solutions with and without As additions[19]:
growth in solution+As

TI
growth in solution As
(2) Translocation factor (TF)
The translocation factor (TF) for As within a plant was ex-
pressed by the ration of Asshoot and Asroot to show As transloca-
tion properties from roots to shoots[20].
(3) Specific arsenic uptake (SAU)
The specific arsenic uptake showed the As uptake abilities
of rice roots:
Croot-As u Root biomass  Cshoot-As u Shoot biomass
SAU
Root biomass
where C expressed the As concentrations in rice seedlings.
The data analysis was performed using SAS 8.0 and Excel.
2 Results and analysis
2.1 Influence of different concentrations of As on growth
of rice seedlings
2.1.1 Root and shoot biomass of experimental rice
Based on variance analysis, there was no significant differ-
ence in the root dry weights among different cultivars treated
by the same dose of As (P > 0.05). For most of the As treat-
ments, the root dry weights were between 0.04 and 0.05 g
except for As1.0, where the root dry weights of hybrid rice
were 0.03 g. The dry weights of the same rice cultivar showed
insignificant differences among different As treatments (P >
0.05), which showed that root biomass of the cultivars was not
obviously restrained by As in the environment (Fig. 1), but
which could also be induced by the smaller root biomass or the
relatively shorter period of treatment. In treatments As0–4.0, the
shoot dry weights of hybrid rice were the lightest among the 3
cultivars. The shoot dry weights of glutinous rice were the
heaviest among the cultivars in As0, 2.0, 4.0 treatments. Those
of hybrid rice decreased with As concentrations increasing in
solutions (P > 0.05). Those of conventional rice increased from
As0 to As0.5, and then decreased with As concentrations in-
creasing (P < 0.05). Those of glutinous rice significantly de-
creased with As concentrations increasing (P < 0.05, Fig. 2).
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235
Fig. 1 Root biomass of rice cultivars growing in solutions
with different As concentrations
Error bars mean standard errors. Different letters in the same group
indicate significant difference at P < 0.05 according to Duncan’s multi-
ple range tests. The same for figures as below
2.1.2 Tolerance degree of rice cultivars to As
The tolerance degrees to As (expressed by tolerance index
TI) of conventional and glutinous rice both significantly de-
creased with As dose increasing (P < 0.01). The TI values of
hybrid rice decreased with As concentrations increasing from
As0.5 to As2.0, while increased from As2.0 to As4.0 (P <
0.05). The TI values of glutinous rice were obviously higher
than those of the other two cultivars in As0.5 to 2.0 treatments
with the ranking of glutinous > conventional > hybrid, and
Fig. 2 Shoot biomass of rice cultivars growing in solutions with
different As concentrations
Fig. 4 As concentrations in root of rice cultivars growing in solu-
tions with different As concentrations
which followed the order of hybrid > glutinous > conventional
in As 4.0 treatment (Fig. 3).
2.2 As accumulation in the seedlings of different rice
cultivars
2.2.1 As accumulation in the roots and shoots of
experimental rice
As concentrations in roots of hybrid rice were significantly
higher than those in the other two cultivars in As0.5 to 2.0
treatments, respectively. As concentrations in roots of gluti-
nous rice were the highest and those in hybrid rice were the
lowest when growing in As4.0 treatment. For As concentra-
tions in roots of rice cultivars, hybrid and conventional rice
both became higher and higher along with As concentrations
increasing in solutions (P < 0.01). Glutinous rice had no ob-
viously change in As0.5 to 2.0 treatments and was up to the
highest in As4.0 treatment (P <0.01, Fig. 4). In As0.5 and 1.0
treatments, As concentrations in shoots of rice cultivars fol-
lowed the order of hybrid > conventional > glutinous. In As2.0
and 4.0 treatments, they followed the orders of conventional >
hybrid > glutinous and hybrid > glutinous > conventional,
respectively. As concentrations in shoots of the three rice cul-
tivars significantly increased with As concentrations increas-
ing in agar nutrient solutions (P < 0.01, Fig. 5). As a whole,
As concentrations in roots and shoots of hybrid rice were rela-
Fig. 3 Tolerance indexes of rice cultivars growing in solutions
with different As concentrations
Fig. 5 As concentrations in shoot of rice cultivars growing in
solutions with different As concentrations
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235

tively higher, while those in glutinous rice were relatively
lower.
2.2.2 Ability of As uptake by rice root system
As shown in Fig. 6, for the specific arsenic uptake (SAU) in
As0.5 and 1.0 treatments, hybrid rice was significantly higher
than conventional and glutinous rice, and there were insig-
nificant differences between conventional and glutinous rice.
The SAU values of conventional rice were the highest among
the three cultivars and insignificantly different from those of
hybrid rice, while those of the hybrid and conventional rice
cultivars were both significantly higher than those of glutinous
rice cultivar. The SAU values of glutinous rice were signifi-
cantly higher than those of hybrid and conventional rice
growing in As4.0 treatment. The SAU values of experimental
rice cultivars significantly increased along with As concentra-
tions increasing in agar nutrient solutions (P < 0.01).
2.3 As translocation from roots to shoots of rice plants
There were no obvious changing rules in the As transloca-
tion capability from roots to shoots of rice plants (expressed
by translocation factor, TF) among the three cultivars with
different As treatments. The values of TF showed different
changing trends along with As doses increasing. There was no
obvious change in the TF values of hybrid rice among three
doses of As treatments, As0.5, As1.0 and As2.0, and the high-
est values of TF (in As2.0 treatment) were just 0.022 units
higher than the lowest ones (in As1.0 treatment). The TF val-
ues significantly increased in As4.0 treatment (P < 0.01). The
TF values of conventional rice did not show significant dif-
ferences among different As treatments. The TF values of
glutinous rice significantly decreased with As concentration
increasing in solutions (P < 0.01, Fig. 7).
2.4 Correlations between biomass, root-As, shoot-As, TI,
TF and SAU
As shown in Table 1, root dry weights had significantly
positive relationship with shoot dry weights, which indicated
that the accumulation of root biomass had positive effect on
the accumulation of shoot biomass. Shoot dry weights had
significantly negative relationship with As concentrations in
roots and shoots, which expressed that As accumulation in
roots and shoots was toxic to rice plants and decreased the
growth of rice plants. The relationships between As concen-
trations in roots and shoots and the TF values were signifi-
cantly positive, which showed that As translocation from roots
to shoots was definite. The As accumulation in roots was the
key factor to influence the TF values, and both of them also
determined the As accumulation in shoots. The fact that TF
and SAU values both had negative relationships with shoot
biomass indicated that the toxicity of As to rice plants reduced
the accumulation of dry matters.
3 Discussion
3.1 Influence of As on the growth of rice
As did not have significant influence on accumulation of
root biomass of experimental rice cultivars. The shoot dry
weights were restrained to a certain degree by relatively

Fig. 6 Specific arsenic uptake of rice cultivars growing in solu- Fig. 7 Translocation factor of rice cultivars growing in solutions
tions with different As concentrations with different As concentrations

Table 1 Person’s product-moment correlation matrix for biomass, As concentrations in roots and shoots, and TI, TF, SAU of rice varieties
(n = 30)
Shoot DW Root-As Shoot-As TI TF SAU
Root DW 0.415* –0.014 –0.068 0.232 –0.008 –0.145
Shoot DW — –0.639** –0.603** –0.032 –0.445* –0.688**
Root-As — 0.827** 0.238 0.532** 0.963**
Shoot-As — 0.036 0.774** 0.868**
TI — 0.318 0.194
TF — 0.641**
Note:* shows that correlation is significant at the 0.05 level; ** shows that correlation is significant at the 0.01 level; DW means dry weight.
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235
higher doses of As (As1.0 to 4.0 mg L–1) in solutions, but ef-
fects were not obvious. The relatively lower doses of As in
solutions were not always toxic to rice, which could enhance
growth of some rice cultivars (varieties). The results showed
that the shoot dry weights of conventional rice growing in
As0.5 treatment solutions were heavier than those of conven-
tional rice growing in As0 treatment solutions. In the experi-
ment done by Liu et al.[12], 6 different rice genetics (6-weeks
old) were exposed to nutrient solutions of 20 M As (V), 10 M
As (III) and 10 M As (V) for 2 weeks. The rice biomass in
their results was higher than that in our results probably be-
cause of the different experimental conditions, experimental
times and rice varieties. Their results suggested that As ad-
vanced the growth of the root system, but reduced the growth
of shoots, which was partially similar to our results. Also, the
results in our experiment were similar to some results under
edaphic conditions. For example, Abedin et al.[5] used 0.2 to
0.8 mg L–1 Na2HAsO4·7H2O to irrigate rice. Their results
showed that the plant height was restrained, but the rice bio-
mass did not obviously reduce with As doses increasing.
Tsutsumi[21] added As (V) into soils and observed that the rice
height did not significantly decrease when As was below 125
mg kg–1. However, the height reduced 63% in soil with 312.5
mg kg–1 As. So low doses of As may not always hurt rice
plants, which may be correlated with the physical and chemi-
cal characteristics of As. As and phosphate are in the same
family[22]. Therefore, on the one hand, the deoxidization ac-
tions of As could improve the activity of oxidases to make
unavailable supply of phosphate effective; on the other hand,
the toxicity of As could kill or restrain harmful pathogen to
enhance plant growth[23]. However, the high doses of As were
harmful to plants.
Conventional rice was more strongly advanced by low As
concentrations than hybrid and glutinous rice. Growth of the 3
cultivars was all reduced by relatively higher concentrations
of As. For the As tolerance of rice cultivars, glutinous rice was
the strongest, hybrid rice was the weakest and conventional
rice was in the middle, which conformed with the ranking of
biomass accumulation: glutinous > conventional > hybrid. So
it could be illuminated that there were close relationships be-
tween root growth strength and dry matter accumulation. Al-
though hybrid rice had the characteristics of fertility, steady-
productivity, extensive-adaptability, etc., hybrid rice had a
weaker As tolerance than conventional and glutinous rice, and
growth of its roots was more severely decreased by As than
that of the other 2 rice cultivars.
3.2 As accumulation and translocation characteristics of
rice
Generally, As concentrations in roots and shoots increased
along with As concentrations increasing in treatment solutions.
However, some opposite circumstances appeared in some As
treatments, probably because the TF values of As translocation
from roots to shoots were different among different As treat-
ments. That was, besides the close relationship of As concen-
trations in environment with As accumulation in roots and
shoots, As translocation capability of seedlings from roots to
shoots (TF) was not neglectable.
As shown in our results, there were significantly positive
relationships between As concentrations in roots and shoots.
The correlation coefficient reached 0.827 (Table 1). However,
As accumulation in roots (156.31–504.03 mg kg–1, represent-
ing 63.40%–81.90% of the total As concentrations in rice) was
far higher than that in shoots (10.52–73.14mg kg–1, represent-
ing 18.10%–36.60% of the total As concentrations in rice).
The TF values of rice cultivars were relatively lower (0.043 to
0.275), probably because As was deposited in the roots by
sulphydryl in proteins of roots to prevent As translocation
from roots to shoots[24]. Therefore, only a small part of As was
translocated to shoots, though roots had strong As uptake ca-
pability. The results of Liu et al.[12] showed that the root As
concentrations ranged from 447 mg kg–1 to 1010 mg kg–1, while
the shoot As concentrations were between 21.3 and 42.0 mg
kg–1. The lowest root As concentrations were more than 10
times larger than the highest shoot As concentrations. There
were partially similar results in the edaphic experiment done
by Cui[24] that when paddy rice fields were irrigated with As
polluted waste water, As concentrations in roots (163.96 mg
kg–1) were greatly higher than that in shoots, and the former
presented 96.5% of the total As concentrations in all parts of
rice seedlings.
The normal order of As accumulations in rice is root >
shoot > crust > grain[14,25]. Some reports have suggested that
rice grains accumulate little As, though the rice plants grow in
polluted environments with higher concentrations of As. For
example, Abedin[5] reported that under 8 mg As L–1 hydro-
ponic conditions, root As concentrations were 107.5 mg kg–1,
but As concentrations in grains were only 0.15–0.42 mg kg–1,
which were lower than the food security standard of Austra-
lia[26]. Du et al.[13] in their edaphic experiment found that when
rice grew in 0 to 200 mg As kg–1 soils by adding Na2HAsO4· 7H2O,
As concentrations in brown rice increased with As concentra-
tions increasing in soils, but the increased extent was very
small, and the highest As concentrations in brown rice were
0.4 mg kg–1, which were not higher than the food security
standard of China (0.7 mg kg–1). Reports showed that only
10% of the grain yield was lost when the grain grew in As
polluted paddy fields with enough fertilizers of N, P, K, etc.
Also, the grain yield in As polluted soils with enough fertiliz-
ers was higher than that in unpolluted soils without enough
fertilizers[27].
Besides, the characteristics of As uptake by rice were espe-
cial compared with other crops. In the experiment done by
Yang and Ge[28], the pollution effects of As on rice, spring
wheat, maize and radish growing in 10 mg kg–1 to 500 mg kg–1
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235
As by adding Na2HAsO4·7H2O and Na3AsO3 into meadow
cinnamon soils were studied. Their results suggested that As
concentrations in radish had already overstepped the food
security standard of China. However, the grains of spring
wheat had not, and the shoots and roots had low levels of As
accumulation. The As storage abilities of rice plants were
much higher than those of other plants; As concentrations in
grains were very low, and the highest was 0.55 mg kg–1. The
detection analysis of Williams et al.[6] showed that in Bangla-
desh suffering serious As pollution, As concentrations in rice
grains ranged between 0.18 mg kg–1 and 0.31 mg kg–1, which
were not higher than the local food security standard for crop
of 1.0 mg kg–1. However, the levels of As in Alocasia odora,
eggplant, cucumber, chayote, coriander, potato, long bean,
radish leaf, dasheen and papaya were 1.93, 1.59, 1.17, 1.06,
0.98, 0.89, 0.87, 0.79, 0.69 mg kg–1 and 0.69 mg kg–1, respec-
tively, which were higher than the local food security stan-
dards for root and tuberous vegetables: 0.79 mg kg–1; fruit
vegetables: 0.56 mg kg–1; leafy vegetables: 0.39 mg kg–1, re-
spectively.
From all above, 2 points could be concluded about As ac-
cumulation and translocation by rice: (1) the root system had
strong abilities to accumulate As; (2) the As uptake by rice
was mostly fixed in roots, and only a small parts of As were
translocated to shoot parts. Rice was more applicable to As
polluted soils compared with other crops. The cultivars (varie-
ties) with low As accumulation abilities in roots and shoots,
high endurance (high TI values), and low As translocation
capability (low TF values) should be selected when planting
rice in As polluted environments. In our results, glutinous rice
had higher biomass, higher tolerance, lower As concentrations
in its roots and shoots, lower translocation factor (TF), lower
specific As uptake ability (SAU) than the other rice cultivars.
Therefore, glutinous rice was more applicable to As polluted
soils compared with the other two rice cultivars, and conven-
tional rice was the second applicable one. It should be forbid-
den to plant hybrid rice in As polluted soils because hybrid
rice had strong abilities to absorb and accumulate high levels
of As, though it had special abilities of antivirus, anti-lodging,
high-productivity, stable-productivity, etc.
Acknowledgements
The authors thank Prof. Zhang Renduo for polishing the ab-
stract and thank Dr. Yang Bing for offering great help and
support when writing the paper. This project was financially
supported by the National “985” Engineering Project of Sci-
ence and Technology for Environmental and Pollution Control
of China.
References
[ 1 ] Abedin M J, Feldmann J, Meharg A A. Uptake kinetics of ar-
senic species in rice plants. Plant Physiology, 2002, 128(3):
1120–1128.
[ 2 ] Mandal B K, Suzuki K T. Arsenic round the world: a review.
Talanta, 2002, 58(1): 201–235.
[ 3 ] Liu W J, Zhu Y G, Smith F A, et al. Do iron plaque and geno-
types affect arsenate uptake and translocation by rice seedlings
(Oryza sativa L.) grown in solution culture? Journal of Ex-
perimental Botany, 2004, 55(403): 1707–1713.
[ 4 ] Silva-Gonzaga M I, Santos J A G, Ma L Q. Arsenic chemistry
in the rhizosphere of Pteris vittata L. and Nephrolepis exaltata
L. Environmental Pollution, 2006, 143(2): 254–260.
[ 5 ] Abedin M J, Cresser M S, Meharg A A, et al. Arsenic accumu-
lation and metabolism in rice (Oryza sativa L.). Environmental
Science and Technology, 2002, 36(5): 962–968.
[ 6 ] Williams P N, Islam M R, Adomako E E, et al. Increase in rice
grain arsenic for regions of Bangladesh irrigating paddies with
elevated arsenic in groundwaters. Environmental Science and
Technology, 2006, 40(16): 4903–4908.
[ 7 ] Xie Z M. Huang C Y. Control of arsenic toxicity in rice plants
grown on an arsenic-polluted paddy soil. Communications in
Soil Science and Plant Analysis, 1998, 29(15): 2471–2477.
[ 8 ] Wang J L, Xu Z J, Ma D R. Comparison on grain filling char-
acters between hybrid and conventional rice in northern China.
Chinese Journal of Rice Science, 2004, 18(5): 425–430.
[ 9 ] Chen Y H, Zhao S, Yan Q Q, et al. Effect of different submer-
gence stress on agronomic traits and biochemical characteris-
tics in hybrid and traditional rice. Chinese Journal of Rice
Science, 2006, 20(5): 512–516.
[10] Liu Y F, Xiao L T, Tong J H, et al. Comparative studies on
vascular bundles of flag leaf in super hybrid rice and conven-
tional rice. Chinese Agricultural Science Bulletin, 2007, 23(7):
151–155.
[11] Li H S, Nie C R, Hu Y G. Effects of simulated acid rain on
hybrid, common and wild rice varieties. Journal of Agro-En-
vironment Science, 2004, 23(2): 284–287.
[12] Liu W J, Hu Y, Bi S Q, et al. Study of genotypic differences on
arsenic uptake by and translocation in rice seedlings. Chinese
Agricultural Science Bulletin, 2006, 22(6): 356–360.
[13] Du D D, Li Y X, Dai B Q, et al. Effect of As to rice growth and
its residual in rice. Environmental Science and Technology,
1987, (3): 11–13.
[14] Liu W J, Zhu Y G, Hu Y, et al. Arsenic sequestration in iron
plaque, its accumulation and speciation in mature rice plants
(Oryza sativa L.). Environmental Science and Technology,
2006, 40(18): 5730–5736.
[15] Wiengweera A, Greenway H, Thomson C J. The use of agar
nutrient solution to simulate lack of convection in waterlogged
soils. Annals of Botany, 1997, 80(2): 115–123.
[16] Colmer T D, Gibberd M R, Wiengweera A, et al. The barrier to
radial oxygen loss from roots of rice (Oryza sativa L.) is in-
duced by growth in stagnant solution. Journal of Experimental
Botany, 1998, 49(325): 1431–1436.
 LIU Zhiyan et al. / Acta Ecologica Sinica, 2008, 28(7):3228–3235
[17] Armstrong W. Radial oxygen losses from intact rice roots as
affected by distance from the apex, respiration and waterlog-
ging. Physiologia Plantarum, 1971, 25: 192–197.
[18] Hewitt E J. Sand and water culture methods used in the study
of plant nutrition. 2nd ed. UK: The Eastern Press, 1966.
[19] Wilkins D A. The measurement of tolerance to edaphic factors
by means of root growth. New Phytologist, 1978, 80: 623–633.
[20] Stoltz E, Greger M. Accumulation properties of As, Cd, Cu, Pb
and Zn by four wetland plant species growing on submerged
mine tailing. Environmental and Experimental Botany, 2002,
47(3): 271–280.
[21] Tsutsumi M. Intensification of arsenic toxicity to paddy rice by
hydrogen sulphide and ferrous iron I. Induction of bronzing
and iron accumulation in rice by arsenic. Soil Science and
Plant Nutrition, 1980, 26: 561–569.
[22] Meharg A A, Macnair M R. Suppression of the high affinity
phosphate uptake system: a mechanism of arsenate tolerance in
Holcus Lanatus L. Journal of Experimental Botany, 1992,
43(249): 519–524.
[23] Zhang G X, Yang J R, Hua L. Arsenic in soil condition and the
ecological effect. Soil, 1996, 28(2): 64–68.
[24] Cui R P. Study of Arsenic distribution in different part of rice.
Guizhou Environmental Protection Science and Technology,
1995, 1(1): 31–32.
[25] Tang L J, Li X D, Liu J H, et al. Effect of arsenic on growth
development of rice and residual rules in the loam paddy soil.
Journal of Jilin Agricultural University, 1996, 18(3): 62–65.
[26] National Food Authority. Australian Food Standard Code: Aus-
tralian Government Publication Service. Canberra, 1993.
[27] Shao H C, Chen J Y, Shen D F. Effect of As contamination to
rice and transform measurements. Shanghai Agricultural Sci-
ence and Technology, 2002, (1): 81.
[28] Yang J Y. Ge J M. Studies on the effects of arsenic on farmland
ecosystem. Acta Ecologica Sinica, 1984, 4(1): 34–45.