Ecotoxicology and Environmental Safety 74 (2011) 13041309

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Ecotoxicology and Environmental Safety

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Iron plaque formation on roots of different rice cultivars and the relation
with lead uptake
Jianguo Liu n, Xuemei Leng, Mingxin Wang, Zhongquan Zhu, Qinghua Dai
School of Environmental and Safety Engineering, Changzhou University, Changzhou, Jiangsu 213164, China

a r t i c l e i n f o a b s t r a c t

Article history: The relationships between lead (Pb) uptake and iron/manganese plaque formation on rice roots were
Received 1 December 2010 investigated with three cultivars. The results showed that the rice cultivars with indica consanguinity
Received in revised form were more sensitive to soil Pb stress than the cultivar with japonica consanguinity. Pb concentrations
24 January 2011
and distribution ratios in root tissues were in the order: Shanyou 634 Yangdao 64 Wuyunjing 7, but
Accepted 27 January 2011
Available online 18 February 2011
Pb and Fe concentrations and distribution ratios in the plaques showed a reverse order. Mn
concentrations and distribution ratios in the plaques of Wuyunjing 7 were signicantly higher
Keywords: (P o 0.01 or 0.05) than those of Shanyou 63 and Yangdao 6. The results indicate that iron/manganese
Rice (Oryza sativa L.) plaque on rice root can provide a barrier to soil Pb stress. The plaque will increase sequestration of Pb
Lead
on rice root surface and in the rhizosphere, providing a means of external exclusion of soil Pb to some
Genotype
extent.
Iron plaque
& 2011 Elsevier Inc. All rights reserved.

1. Introduction exclusion of metals (Greipsson and Crowder, 1992; Hansel et al.,

Under anaerobic conditions, aquatic plants such as rice have a
common feature that is the formation of iron plaque on the root
surfaces. That is due to the release of oxygen and oxidants in the
rhizosphere, and the subsequent oxidation of ferrous to ferric iron
with the precipitation of iron oxide or hydroxide on the root
surface. In addition to Fe, the plaque contains a wide range of
other metal elements. In nature, the plaque element next in
importance to Fe is generally Mn (Taylor et al., 1984; Crowder
and Coltman, 1993). The main forms of iron plaque are the ferric
hydroxides, goethite, and lepidocrocite (Chen et al., 1980). Due to
the high adsorption capacity of functional groups on iron hydro-
xides, iron plaque may sequester a number of metal(loid)s by
adsorption or co-precipitation, and thus interferes with the
availability of these elements in the rhizosphere and their uptake
by and translocation in plants. So the presence of iron coating on
root surface may inuence metal uptake (Otte et al., 1989; Liu
et al., 2007).
Although some studies have been conducted on iron plaque
and metal uptake in wetland plants, the results were not
consistent with each other. Some of them indicated that the
formation of Fe and Mn precipitate (plaque) on root surface of
aquatic plants may provide a means of attenuation and external
n
Corresponding author.
E-mail addresses: liujianguo@cczu.edu.cn,
ljg@jpu.edu.cn, liu-jg703@sohu.com (J. Liu).
2002). Whereas, Ye et al. (1997, 1998) reported that there is no
difference between the presence and absence of iron plaque in
plant Cu and Zn uptake of Typha latifolia. Iron plaque on rice root
also affected patterns of metal uptake and accumulation, and
lower concentrations of Cu and Ni were found in the leaves of rice
plants with plaque than in the plants without plaque (Greipsson
and Crowder, 1992; Greipsson, 1994). However, Liu et al. (2008)
reported that iron plaque on rice root surface is of little signi-
cance in affecting uptake and accumulation of Cd by rice plant.
Furthermore, those works were mainly conducted under hydro-
ponic conditions and/or different levels of Fe additions, few
experiments were conducted under soil conditions with natural
Fe concentrations.
Lead is a worldwide contaminant in the environments. Owing
to industrial development and population expansion, the concen-
trations of heavy metals in agricultural soils have also become
increasingly serious in China (Li et al., 2000; Ji et al., 2000). Pb is
toxic to many organ systems of human body, especially to central
nervous systems of children (Wasserman et al., 1997; Silbergeld
et al., 2000).
Paddy rice is one of the most important crops in the world,
especially in Asia. It was reported that there were differences
between rice genotypes in Fe content in iron plaque (Geng et al.,
2005). Our previous studies showed that rice cultivars varied
signicantly in Pb accumulation and tolerance (Liu et al., 2003).
However, limited information is available on the relationship
between plant Pb uptake and Fe/Mn plaque formation on root
for different rice cultivars. The present research was conducted to

0147-6513/$ - see front matter & 2011 Elsevier Inc. All rights reserved.
doi:10.1016/j.ecoenv.2011.01.017
 J. Liu et al. / Ecotoxicology and Environmental Safety 74 (2011) 13041309
investigate the relationship and to understand some mechanisms
causing the variations between rice cultivars with regard to Pb
uptake and accumulation.
2. Materials and methods
2.1. Soil preparation and analysis
The soils for the pot experiments were collected from uncontaminated elds
(020 cm). After air-drying and passing through a 2-mm sieve, the following soil
properties were tested: (1) particle size with hydrometer method; (2) pH with pH
meter (soil : distilled water 1: 2); (3) organic matter with sequential extraction
method; (4) cation exchange capacity with ammonium acetate method; and
(5) total Fe, Mn and Pb with AAS following H2O2HFHNO3HClO4 digestion
(Sparks, 1996). The testing results are shown in Table 1.
Four kilogram of soil was placed in each pot (18 cm in diameter and 20 cm in
height). Pb in the form of PbCl2 was added to the soil to obtain Pb levels of 500 and
1000 mg kg  1 (dry weight).
PbCl2 was dissolved in deionized water and poured into the soil slowly while
mixing the soil at the same time. The thoroughly mixed soils were stored in pots
and submerged in water (23 cm above the soil surface) for a month before rice
seedlings were transplanted. The soils without adding Pb served as control.
2.2. Rice plant materials
Based on our previous studies (Liu et al., 2003), three rice cultivars of different
genotypes with diverse Pb uptake abilities were used in this experiment, and they
are Shanyou 63 (hybrid indica, a high Pb accumulator), Yangdao 6 (indica, a
moderate Pb accumulator), and Wuyunjing 7 (japonica, a low Pb accumulator).
Rice seeds were submerged in a water bath for about 48 hr at room temperature
(2025 1C) and germinated under moist condition (seeds were covered with two
layers of moist gauze cloth) at 32 1C for another 30 hr and the germinated seeds
were grown in uncontaminated soils. After 30 days, the uniform seedlings were
selected and transplanted into the pots (3 plants per pot). The pot soil was
maintained under ooded conditions (with 23 cm of water above soil surface)
during the rice growth period.
2.3. Experimental design
The experiments were carried out under open-air conditions. The pots were
arranged in a randomized complete block design with ve replicates. 0.5 g of urea
(containing 46% N) and 0.5 g of K2HPO4  3 H2O were applied to each pot at two
times, one on the 3rd day before seedling transplant, and the other on day 20 after
the transplant.
2.4. Cold DCB extraction of iron and manganese plaque
One entire plant (including shoot and root system) was sampled from each pot
at day 40 after seedling transplant. The plant samples were washed thoroughly
with tap water and deionized water, and then separated into roots and shoots.
Iron and manganese plaque on the roots were extracted using a modied cold
dithionitecitratebicarbonate (DCB) method (Taylor and Crowder, 1983; Otte
et al., 1989; Liu et al., 2008). The entire root systems were rstly incubated for 2 h
at 20 1C in 80 ml 0.03 mol L  1 sodium citrate (Na3C6H5O7  2H2O) and
0.125 mol L  1 sodium bicarbonate (NaHCO3) with the addition of 1.6 g sodium
dithionite (Na2S2O4), and the asks were shaken during the incubation. The
extracts were then transferred into 100-ml glass asks. Roots were rinsed three
times with deionized water and the eluates were added to the DCB extracts.
Finally the solutions were ltered into plastic containers for subsequent analysis.
Table 1
Selected properties of the soil used in this experiment (n 3).
Soil type Soil texture Particle size (g kg  1) pH
Sand Silt Clay
Paddy soil Sandy loam 587.5 233.2 179.3 6.8
a
Organic matter.
b
Cation exchange capacity.
1305
2.5. Determination of Pb concentrations in rice plants
After DCB extraction, the roots and shoots were oven-dried at 70 1C to a constant
weight. The oven-dried samples were ground with a stainless steel grinder to pass
through a 100-mesh sieve. Pb concentrations of the samples were determined with an
Atomic Absorption Spectrophotometer (Solaar S4+Graphite Furnace System 97,
Thermo Elemental, USA) following HNO3HClO4 (HNO3:HClO4 4:1, V:V) digestion
procedures (Allen, 1989).
Data were analyzed with the statistical package SPSS11.5 and EXCEL2003 for
Windows. Two signicant levels of P r 0.05 and 0.01 were used in presenting the
results.
3. Results
Table 1 shows that the soil used was a sandy loam with a high
proportion of sand and neutral pH. The soil contained a moderate
level of organic matter, CEC, Fe and Mn, and a low level of Pb.
3.1. Effects of soil Pb treatments on plant biomasses of different rice
cultivars
Table 2 shows that the effects of soil Pb treatments on rice
growth differed with rice cultivars, soil Pb levels and plant organs.
In general, Yangdao 6 and Shanyou 63 were more sensitive to soil
Pb stress than Wuyunjing 7.
Under soil Pb treatment of 500 mg kg  1, the biomass of
Shanyou 63 increased by about 10%, as compared to the control,
and root and total weights of rice plant increased signicantly
(P o0.05). However, the biomass of Yangdao 6 decreased by about
8%, and the root weight decreased signicantly (Po0.05). The
biomass of Wuyunjing 7 changed little.
Under soil Pb treatments of 1000 mg kg  1, root and shoot
weights decreased for all the rice cultivars. But the magnitudes of
decreases varied largely with cultivars and plant organs. With
regard to root weights, the reduction rates were more than 20%
for Yangdao 6 and Shanyou 63, but less than 10% for Wuyunjing 7.
With respect to shoot weights, the decreases were signicant at
Po0.01 level for Yangdao 6, and at Po0.05 level for Shanyou 63.
However, the decrease rate was only 5% for Wuyunjing 7, an
insignicant level (P40.05).
Under soil Pb stress, the change rates in dry weights (decrease
or increase) were generally higher for roots than for shoots.
3.2. Pb uptake and distributions in the plants of different rice
cultivars
The differences among the rice cultivars in Pb concentrations
and distributions in roots are shown in Table 3.
In general, Pb concentrations in root tissues and shoots were in
the order: Shanyou 634Yangdao 64Wuyunjing 7. The differences
between the cultivars were signicant (Po0.05), except for the
difference between Shanyou 63 and Yangdao 6 in shoot Pb con-
centrations under soil Pb level of 500 mg kg  1. But Pb concentra-
tions in plaques showed a reverse order. They were the highest for
Wuyunjing 7, and the lowest for Shanyou 63, except for the
difference between Shanyou 63 and Yangdao 6 of the control.
OMa CECb Total Fe Total Mn Total Pb
(g kg  1) (cmol kg  1) (mg kg  1) (mg kg  1) (mg kg  1)
27.6 13.6 8981 683.9 36.5
1306 J. Liu et al. / Ecotoxicology and Environmental Safety 74 (2011) 13041309

Table 2
Effects of soil Pb treatments on plant biomasses of different rice cultivars (n 3).

Cultivars Soil Pb concentration Root Shoot Total

(mg kg  1)
Dry weight RC a( 7%) Dry weight RC (7 %) Dry weight RC (7%)
(g pot  1) (g pot  1) (g pot  1)

Shanyou 63 Control 5.31 11.42 16.73

500 5.99n 12.81 12.48 9.28 18.47n 10.40
1000 4.18nn  21.28 9.87n  13.57 14.05nn  16.02
Yangdao 6 Control 4.34 9.88 14.22
500 3.91n  9.91 9.15  7.39 13.06  8.16
1000 3.27nn  24.65 7.92nn  19.84 11.19nn  21.31
Wuyunjing 7 Control 2.14 4.54 6.68
500 2.18 1.87 4.69 3.30 6.87 2.84
1000 1.93n  9.81 4.31  5.07 6.24  6.59

a
Relative changes under soil Pb treatment compared to the control. RC( 7%) ((weight of Pb treatment  weight of control)/weight of control)  100.
nn
Signicant difference between Pb treatment and the control at P r0.01.
n
Signicant difference between Pb treatment and the control at P r0.05.

Table 3
Pb concentrations in rice plants and distribution in the roots (n 3).

Soil Pb concentration Cultivars Root tissue Plaquea Total Pb concentration Pb concentration in

(mg kg  1) of root (mg g  1) shoot (mg g  1)
Pb concentration % Pb concentration %
(mg g  1) (mg g  1)

Control Shanyou 63 8.34 a 26.10 23.62 a 73.90 31.96 a 7.43 a

Yangdao 6 6.72 b 22.63 22.98 a 77.37 29.70 a 5.48 b
Wuyunjing 7 4.12 c 14.02 25.27 b 85.98 29.39 a 3.82 c
500 Shanyou 63 2299 a 91.28 219.56 a 8.72 2519 a 114.36 a
Yangdao 6 1747 b 87.98 238.73 b 12.02 1986 b 115.49 a
Wuyunjing 7 998 c 78.57 272.33 c 21.43 1271 c 90.97 b
1000 Shanyou 63 6201 a 93.72 415.68 a 6.28 6617 a 273.24 a
Yangdao 6 5199 b 91.51 482.67 b 8.49 5682 b 245.91 b
Wuyunjing 7 4239 c 88.58 546.32 c 11.42 4785 c 228.75 c

Different letters beside the data in the same treatment indicate signicant difference between the cultivars at P r 0.05.
a
Pb concentrations in plaques are also based on root dry weights.

With regard to Pb distributions in roots (between root tissues
and the plaques), Pb distribution ratios in root tissues were much
higher for soil Pb treatments than for the control, but the ratios in
the plaques were much lower for soil Pb treatments than for the
control. Pb distribution ratios increased with the increase of soil
Pb levels in root tissues, but decreased with the increase of soil Pb
levels in plaques. On the differences among the rice cultivars, Pb
distribution ratios in root tissues were in the order: Shanyou
634Yangdao 64Wuyunjing 7. But Pb distribution ratios in
plaques showed a reverse order, which were the highest for
Wuyunjing 7 and the lowest for Shanyou 63.
With respect to Pb accumulations in rice plants (Table 4),
Shanyou 63 was much higher than Yangdao 6, and Yangdao 6 was
much higher than Wuyunjing 7, irrespective of in root tissues,
plaques or shoots. The differences between two cultivars of indica
consanguinity (Shanyou 63 and Yangdao 6) and the cultivar of
japonica consanguinity (Wuyunjing 7) were very enormous.
On the distributions of Pb in different parts of rice plants, it
varied with rice cultivars, soil Pb levels and the parts of rice plants.
For the control, Pb distribution ratios in different parts of rice plants
were in the order: plaque4shoot4root tissue, about half or more
of the Pb absorbed by rice plant was stored in plaques, about 22
33% in shoots, and about 1117% in root tissues. But for soil Pb
treatments, most of the Pb was accumulated in root tissues (from
about 68% to about 85%), about 619% in plaques, and about 913%
in shoots. Pb distribution ratios in the plaques dropped sharply
with the increase of soil Pb levels. Under soil Pb treatment of
1000 mg kg  1, only 10% or much less of Pb absorbed by rice plants
was restrained in the plaque. Concerning the differences among the
rice cultivars in Pb distributions, the distribution ratios in root
tissues were in the order: Shanyou 634Yangdao 64Wuyunjing
7, but the ratios in the plaques showed a reverse order, with the
highest for Wuyunjing 7 and the lowest for Shanyou 63. The
differences among the rice cultivars in Pb distribution ratios in
shoots varied with soil Pb levels.
3.3. Fe and Mn concentrations in rice plants and distribution in the
roots
Fe concentrations in different parts of rice plants and distribu-
tion in roots are displayed in Table 5. On Fe concentrations in root
tissues and in plaques, they were signicantly higher (Po0.01)
for Wuyunjing 7 than for Yangdao 6, and signicantly higher
(Po 0.01 or 0.05) for Yangdao 6 than for Shanyou 63, irrespective
of soil Pb levels. The magnitudes of the differences among the rice
cultivars were higher under soil Pb treatments (500 mg kg  1 and
1000 mg kg  1) than for the control. The differences among the
rice cultivars in shoot Fe concentrations varied with soil Pb levels.
On Fe distribution in rice roots, absolute majority of Fe
accumulated in the plaques (more than 98.5%), and only less than
1.5% of Fe was transferred into root tissues. Fe distribution ratios in
the plaques of three rice cultivars were in the order: Wuyunjing
74Yangdao 64Shanyou 63, but the ratios in root tissues showed
 J. Liu et al. / Ecotoxicology and Environmental Safety 74 (2011) 13041309 1307

Table 4
Pb accumulations and distributions in different parts of rice plants (n 3).

Soil Pb Cultivars Root tissue Plaque Shoot Total Pb

concentration accumulation
(mg kg  1) in rice plant
(mg pot  1)
Pb accumulation % Pb accumulation % Pb accumulation %
(mg pot  1) (mg pot  1) (mg pot  1)

Control Shanyou 63 44.29 a 17.40 125.42 a 49.27 84.85 a 33.33 254.56 a

Yangdao 6 29.16 b 15.93 99.73 b 54.49 54.14 b 29.58 183.04 b
Wuyunjing 7 8.82 c 10.99 54.08 c 67.40 17.34 c 21.61 80.24 c
500 Shanyou 63 13774 a 83.40 1315 a 7.96 1427 a 8.64 16516 a
Yangdao 6 6830 b 77.44 933 b 10.58 1057 b 11.98 8820 b
Wuyunjing 7 2176 c 68.08 594 c 18.57 427 c 13.35 3196 c
1000 Shanyou 63 25921 a 85.39 1738 a 5.72 2697 a 8.88 30356 a
Yangdao 6 17002 b 82.82 1578 b 7.69 1948 b 9.49 20528 b
Wuyunjing 7 8181 c 80.04 1054 c 10.32 986 c 9.65 10221 c

Different letters beside the data in the same treatment indicate signicant difference between the cultivars at Pr 0.05.

Table 5
Fe concentrations in different parts of rice plant and distribution in the root (n 3).

Soil Pb concentration Cultivars Root tissue Plaquea Total Fe concentration Fe concentration in

(mg kg  1) of root (mg g  1) shoot (mg g  1)
Fe concentration % Fe concentration %
(mg g  1) (mg g  1)

Control Shanyou 63 684.36 c 1.31 51485 c 98.69 52169 c 233.42 a

Yangdao 6 749.68 b 1.27 58268 b 98.73 59018 b 208.79 b
Wuyunjing 7 862.19 a 1.18 72468 a 98.82 73330 a 195.85 b
500 Shanyou 63 715.43 c 1.35 52394 c 98.65 53110 c 239.37 ab
Yangdao 6 798.42 b 1.29 60984 b 98.71 61782 b 258.74 a
Wuyunjing 7 995.46 a 1.21 81596 a 98.79 82591 a 236.53 b
1000 Shanyou 63 732.09 c 1.48 48716 c 98.52 49447 c 265.56 a
Yangdao 6 853.77 b 1.39 60637 b 98.61 61491 b 271.36 a
Wuyunjing 7 1101.49 a 1.29 84478 a 98.71 85580 a 238.43 b

Different letters beside the data in the same treatment indicate signicant difference between the cultivars at Pr 0.05.
a
Fe concentrations in plaques are also based on root dry weights.

a reverse order, irrespective of soil Pb levels. Nevertheless, the
magnitudes of the differences among the rice cultivars were small.
Mn concentrations in different parts of rice plants and dis-
tribution in roots are displayed in Table 6. Mn concentrations in
plaques increased with the increase of soil Pb levels for three rice
cultivars, but Mn concentrations in root tissues decreased with
the increase of soil Pb levels. In general, Mn concentrations in root
tissues of Wuyunjing 7 were signicantly lower (Po0.01) than
those of Shanyou 63 and Yangdao 6, but Mn concentrations in
plaques of Wuyunjing 7 were signicantly higher (P o0.01 or
0.05) than those of Shanyou 63 and Yangdao 6 (except for the
differences between Wuyunjing 7 and Shanyou 63 of the control).
With regard to Mn distributions in the roots, most of Mn was
absorbed in plaques (about 6388%), only about 1237% was
accumulated into root tissues. Mn distribution ratios in root
tissues of Wuyunjing 7 were signicantly lower (Po0.01) than
those of Shanyou 63 and Yangdao 6, but the ratios in plaques of
Wuyunjing 7 were signicantly higher (Po0.01) than those of
Shanyou 63 and Yangdao 6.
4. Discussion
4.1. Variations among rice cultivars in Pb tolerance and the relation
with iron and manganese plaque formation on roots
The levels of Pb in soils that are toxic to plant are not easy to
evaluate, and the toxic symptoms of Pb in plant are not very specic.
Many researches showed that Pb phytoxicity resulted in oxidative
stress and retardation of plant growth. The inhibitory effects may be
due to interference with enzymes essential for normal metabolic
and development, photosynthetic processes, water and mineral
nutrients absorption, and changes in cell ultrastructure (Stefanov
et al., 1995; Paivoke, 2002; Zeng et al., 2006).
It has been reported that plant species differed greatly in their
tolerance to soil Pb stress (Rooney et al., 1999; Paivoke, 2002),
and the genotypes within a same plant species also differed
greatly in their tolerance to soil Pb stress (Liu et al., 2003). But
the mechanisms have not been fully understood.
It was reported that deposition of iron oxide plaque or coating
on rice root ameliorated the toxic effects of Cu, Ni, and Cu +Ni on
plant growth (Greipsson and Crowder, 1992; Greipsson, 1994).
However, other reports showed no differences in root and shoot
dry weights and leaf elongation between presence and absence of
the plaque, when Cu or Ni were added to solutions (Ye et al.,
1997).
In our present research, effects of soil Pb stress on rice
biomasses varied largely with cultivars. Under relatively lower
soil Pb treatment (500 mg kg  1), the biomasses changed little
(P 40.05) for Wuyunjing 7, but those increased largely for
Shanyou 63 (signicant at Po0.05 for root weight), decreased
obviously for Yangdao 6 (signicant at Po0.05 for root weight).
Pb is not an essential element for rice growth and metabolism, the
stimulating effects of relatively lower level of Pb on rice growth
may be another response of rice plant to Pb stress, but the
underlying mechanisms need further research.
1308 J. Liu et al. / Ecotoxicology and Environmental Safety 74 (2011) 13041309

Table 6
Mn concentrations in different parts of rice plant and distribution in the root (n 3).

Soil Pb concentration Cultivars Root tissue Plaquea Total Mn concentration Mn concentration

(mg kg  1) of root (mg g  1) in shoot (mg g  1)
Mn concentration % Mn concentration %
(mg g  1) (mg g  1)

Control Shanyou 63 16.62 a 34.34 31.78 ab 65.66 48.40 a 432.75 a

Yangdao 6 17.59 a 36.66 30.39 b 63.34 47.98 a 426.35 a
Wuyunjing 7 12.37 b 26.90 33.62 a 73.10 45.99 a 375.48 b
500 Shanyou 63 15.58 a 28.94 38.25 b 71.06 53.83 b 384.92 a
Yangdao 6 14.28 a 28.82 35.27 b 71.18 49.55 b 336.76 b
Wuyunjing 7 10.49 b 15.60 56.76 a 84.40 67.25 a 336.32 b
1000 Shanyou 63 12.75 a 18.44 56.38 b 81.56 69.13 a 178.66 c
Yangdao 6 12.38 a 21.00 46.57 c 79.00 58.95 b 217.53 b
Wuyunjing 7 8.73 b 11.76 65.51 a 88.24 74.24 a 242.31 a

Different letters beside the data in the same treatment indicate signicant difference between the cultivars at P r 0.05.
a
Mn concentrations in plaques are also based on root dry weights.

It can be seen from present results that Shanyou 63 and
Yangdao 6 (with indica consanguinity) were more sensitive to
soil Pb stress than Wuyunjing 7 (with japonica consanguinity).
Our results also showed that Fe and Mn concentrations in the
plaques of Wuyunjing 7 were signicantly higher (Po0.01,
oro0.05) than those of Shanyou 63 and Yangdao 6, and the
magnitude of the difference among the cultivars was higher for
soil Pb treatments than for the control.
So iron and manganese plaques on rice roots may provide a
barrier to soil Pb stress in Pb-contaminated soils.
4.2. Variations among rice cultivars in plant Pb uptake and the
relation with iron and manganese plaque formation on roots
Literature reports indicated that presence of iron plaque
inhibited uptake of heavy metals by plants, and Cu concentrations
were lower in the plants of Lobelia dortmanna and Phragmites
australis in the presence of plaque than when it was absent
(Christensen and Jensen, 1998; Batty et al., 2000). However, the
results of Ye et al. (1997) also presented that the plaque of Typha
latifolia did not alter Cu uptake and translocation but increased Ni
uptake and translocation. Ye et al. (1998) also showed that Zn was
absorbed fairly equally on root surfaces with and without plaque,
but the roots with plaque absorbed more Pb, but less Cd, than
those without. The presence of plaque did not alter Zn, Pb, and
Cd translocation in the seedlings of Typha latifolia in nutrient
solutions.
Our present research showed that there were great differences
among three rice cultivars in plant Pb uptake, specically
between the cultivars of indica consanguinity (Shanyou 63 and
Yangdao 6) and the cultivar of japonica consanguinity (Wuyunj-
ing 7). Pb concentrations in root tissues and shoots in the order:
Shanyou 63 4Yangdao 64 Wuyunjing 7. But Pb concentrations in
plaques showed a reverse order. With regard to Pb distributions
in roots, Pb distribution ratios increased with the increase of soil
Pb levels in root tissues, but decreased with the increase of soil Pb
levels in plaques. Pb distribution ratios in root tissues were in the
order: Shanyou 634Yangdao 64Wuyunjing 7. But Pb distribu-
tion ratios in plaques showed a reverse order, and they were the
highest for Wuyunjing 7 and the lowest for Shanyou 63.
Meanwhile, Fe concentrations in the plaques were signi-
cantly higher (Po0.01) for Wuyunjing 7 than for Yangdao 6,
and signicantly higher (Po0.01 or 0.05) for Yangdao 6 than for
Shanyou 63 (Table 5). The magnitudes of the differences among
the rice cultivars were higher under soil Pb treatments
(500 mg kg  1 and 1000 mg kg  1) than for the control. The dis-
tribution ratios of Fe in iron plaques of different rice cultivars
were in the order: Wuyunjing 74Yangdao 64Shanyou 63, but
the ratios in root tissues showed a reverse order. In general, Mn
concentrations and distribution ratios in plaques of Wuyunjing
7 were signicantly higher (Po0.01 or 0.05) than those of
Shanyou 63 and Yangdao 6, but Mn concentrations and distribu-
tion ratios in root tissues of Wuyunjing 7 were signicantly lower
(Po 0.01) than those of Shanyou 63 and Yangdao 6.
So it can be concluded that formation of iron and manganese
plaque on rice root can increase the sequestration of Pb on root
surface and in the rhizosphere, providing a means of external
exclusion of soil Pb. But the functions of iron and manganese
plaque were limited, only effective in relatively lower or moder-
ate Pb-contaminated soil (for example less than 1000 mg kg  1),
because Pb distribution ratios in the plaques dropped sharply
with the increase of soil Pb levels (Tables 3 and 4). Under soil Pb
level of 1000 mg kg  1, only 10% or much less of Pb absorbed by
rice plants was restrained in the plaque. Under higher Pb levels,
the capacity of iron plaque in sequestrating Pb may turn to
saturation.
5. Conclusions
In general, Shanyou 63 and Yangdao 6 (with indica consangui-
nity) were more sensitive to soil Pb stress than Wuyunjing 7 (with
japonica consanguinity). Pb concentrations and distribution ratios
in root tissues were in the order: Shanyou 634Yangdao
64Wuyunjing 7. But Pb concentrations and distribution ratios
in iron plaques showed a reverse order. On Fe concentrations in
plaques, they were signicantly higher (Po 0.01) for Wuyunjing
7 than for Yangdao 6 and Shanyou 63. Fe distribution ratios in the
plaques were in the order: Wuyunjing 7 4Yangdao 6 4Shanyou
63, but the ratios in root tissues showed a reverse order. Mn
concentrations and distribution ratios in plaques of Wuyunjing
7 were signicantly higher (Po0.01 or 0.05) than those of
Shanyou 63 and Yangdao 6, but Mn concentrations and distribu-
tion ratios in root tissues of Wuyunjing 7 were signicantly lower
(Po 0.01) than those of Shanyou 63 and Yangdao 6.
On the basis of our results, it can be concluded that iron and
manganese plaques on rice roots can provide a barrier to soil Pb
stress in Pb-contaminated soils. The plaque will increase the
sequestration of Pb on the root surface and in the rhizosphere,
providing a means of external exclusion of soil Pb. But the
functions of plaque are limited, only effective in relatively lower
 J. Liu et al. / Ecotoxicology and Environmental Safety 74 (2011) 13041309
or moderate Pb-contaminated soil (for example less than
1000 mg kg  1), because Pb distribution ratios in the plaques
dropped sharply with the increase of soil Pb levels. Under soil
Pb concentration of 1000 mg kg  1, only 10% or much less of Pb
absorbed by rice plants was restrained in the plaque.
Acknowledgments
This work was supported by the National Natural Science
Foundation of China (No. 31071350, 70901035), the Key Project
for Agricultural Science and Technology of Changzhou (No.
CE20102023) and the Natural Science Foundation of Jiangsu
Province (No. BK2008144).
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