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Dynamic changes in radial oxygen loss and iron plaque

Dynamic changes in radial oxygen loss and iron plaque
Dynamic changes in radial oxygen loss and iron plaque

ORIGINAL PAPER

Dynamic changes in radial oxygen loss and iron plaque formation and their effects on Cd and As accumulation

in rice (Oryza sativa L.)

Xun Wang ?Haixin Yao ?Ming Hung Wong

?

Zhihong Ye

Received:18December 2012/Accepted:26March 2013/Published online:14June 2013óSpringer Science+Business Media Dordrecht 2013

Abstract Temporal variations and correlations

between radial oxygen loss (ROL),iron (Fe)plaque formation,cadmium (Cd)and arsenic (As)accumula-tion were investigated in two rice cultivars at four different growth stages based upon soil pot and deoxygenated solution experiments.The results showed that there were signi?cant differences in ROL (1.1–16l mol O 2plant -1h -1),Fe plaque formation (4,097–36,056mg kg -1),Cd and As in root tissues (Cd 77–162mg kg -1;As 49–199mg kg -1)and Fe plaque (Cd 0.4–24mg kg -1;As 185–1,396mg kg -1)between these growth stages.ROL and Fe plaque increased dramatically from tillering to ear emergence stages and

then were much reduced at the grain-?lling stage.

Furthermore,signi?cantly positive correlations were detected between ROL and concentrations of Fe,Cd

and As in Fe plaque.Our study indicates that increased Fe plaque forms on rice roots at the ear emergence stage due to the increased ROL.This stage could therefore be an important period to limit the transfer and distribution of Cd and As in rice plants when growing in soils contaminated with these toxic elements.

Keywords Iron plaque áDynamic changes áRice áCadmium áArsenic

Introduction

Cadmium (Cd)is an element that is of great environ-mental and toxicological concern due to its acute and chronic toxic effects on biota and human through contamination of the food chain (Liu et al.2007b ).It can be released into the environment by natural processes and through human activities,such as disposal of industrial ef?uents and mining wastes,and agricultural application of sewage sludge or

phosphate fertilizer (Ye et al.2000;Williams et al.

2009).Soil pollution by Cd has been of public concern since the occurrence of Itai–Itai disease in Japan in the

1950s and 1960s,due to high levels of Cd contained in rice (Oryza sativa L.)(Obata and Umebayashi 1997;McLaughlin et al.1999).In addition to Cd,arsenic (As)is a toxic metalloid pollutant,the risk of which for human health has also attracted the world’s attention in recent years (Williams et al.2006,2007;Zhu et al.

X.Wang áH.Yao áZ.Ye (&)

Key Laboratory of Biodiversity Dynamics and

Conservation of Guangdong Higher Education Institutes,School of Life Sciences,Sun Yat-sen University,Guangzhou 510006,People’s Republic of China e-mail:lsshzhh@https://www.doczj.com/doc/1e13165533.html,;lssyzhh@https://www.doczj.com/doc/1e13165533.html,

M.H.Wong

Croucher Institute for Environmental Sciences,Hong Kong Baptist University,Hong Kong SAR,People’s Republic of China

M.H.Wong

Department of Biology,Hong Kong Baptist University,Hong Kong SAR,People’s Republic of China

Environ Geochem Health (2013)35:779–788DOI 10.1007/s10653-013-9534-y

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统计,[数] 相关

去氧的n. [植] 分蘖;发棵

n. 污水污泥;下水污泥

痛痛病

2008b ;Hanh et al.2010).Grains of rice harvested

from As-contaminated paddy ?elds provide an impor-tant risk source of As for consumers,as a consequence

of the high ef?ciency of rice in accumulating As

(Williams et al.2009;Zhao et al.2010).Rice is the

staple food in Asia;therefore,minimizing grain Cd

and As in rice production,especially in Cd/As-contaminated areas,is of great importance.

In order to adapt to anaerobic conditions,rice develops aerenchyma to transfer O 2from the aerial

parts to its roots,resulting in O 2diffusing toward the

root apex and its rhizosphere (Justin and Armstrong

1987)—a process which is termed radial oxygen loss

(ROL)(Colmer 2003).ROL can oxidize rhizospheric

soil substances and cause precipitation of toxic metals

onto the rhizosphere soil and root surfaces (Otte et al.

1989).Recent studies indicated that rice cultivars with

higher rates of ROL possess higher capacities for

limiting the transfer of Cd (Wang et al.2011)and As

(Mei et al.2009)to above-ground tissues.

Like other wetland plant species,rice can form Fe

plaque on its roots by oxidizing Fe 2?to Fe 3?,resulting

from ROL from plants (Taylor et al.1984)and

biological oxidation by microorganisms (Weiss et al.

2003).Due to the high capacity of functional groups

on Fe hydroxides,Fe plaque is able to sequester

metal(loid)s by adsorption and/or co-precipitation,

thus affecting the bioavailability of these elements in

the rhizosphere,which may lead to changes in the

uptake and accumulation of elements by the plants (Mei et al.2009,2012;Wang et al.2011).The

presence of Fe plaque has been reported to in?uence

metal tolerance and uptake in aquatic plants (Ye et al.

1997;Batty et al.2000).However,the exact effects of

Fe plaque on Cd and As uptake in rice plants are still

unclear.It has also been reported that the Fe plaque on

rice roots may have positive,negative or negligible

impacts on Cd uptake (Liu et al.2001,2007a ,b ,2008).

Liu et al.(2001)found that Cd in the plaque was at a

higher concentration than that in the root tissues of 14

rice cultivars,indicating that the plaque had a strong

absorbing capacity and blocked Cd absorption.In

contradiction,Liu et al.(2008)found that enhance-ment of Fe uptake by rice can diminish the negative

effects of Cd,but that Fe plaque on root surfaces is of

little signi?cance in affecting uptake and accumula-tion of Cd by rice plants.Ye et al.(1998)also

suggested that root tissue rather than Fe plaque is the

main barrier for Cd transport.Fe plaque may act as an effective Fe reservoir to increase Fe ion concentrations in active cells and then ameliorate metal toxicity.For As,it has been suggested that the formation of Fe plaque on the roots signi?cantly limits the uptake of As by rice,under both glasshouse (Chen et al.2005)and paddy ?eld conditions (Garnier et al.2010).Chen et al.(2005)reported that the Fe plaque had a signi?cant effect on the absorption kinetics of As by rice roots,decreasing arsenate uptake but increasing arsenite uptake.However,it should be emphasized that the rhizosphere effect on metal(loid)s (e.g.,Cd,As)uptake by rice plants is complex,and Fe plaque may serve as a sink or a source of metal(loid)s at different growth stages of plants (Zhao et al.2010).It is also worth noting that rice root Fe plaque may change in amount and composition during rice growth,and this variation may cause changes in the amount and speciation of Fe oxides in the rice rhizosphere (Zhang et al.2012).Nanzyo et al.(2010)reported that the quantity of root Fe plaque reaches its peak at the tillering stage,after which it gradually decreased.Besides that,root secretions may lead to an increase in dissolved organic matter (DOM),which could provide protons and electrons for reductive dissolution of Fe plaque (Zhang et al.2012).The level of DOM is the highest in the earing and ?owering stage and decreases gradually from the grain-?lling stage to the ripening stage (Wang et al.2004).However,there is a lack of information on how ROL and Fe plaque formation change over the entire growing season of rice,and what effects these changes may have on the accumu-lation and translocation of Cd or As in rice.Thus,the aims of the present study were to determine the effects of Fe plaque on Cd and As uptake by rice,investigate the dynamic changes of ROL and Fe plaque formation and temporal variations in Cd and As accumulation and translocation at four (tillering,bolting,ear emer-gence and grain ?lling)de?nable growth stages.Materials and methods Preculture of rice seedlings Two rice cultivars,a hybrid Tianyou 116(TY)and a conventional Huaxinzhan (HX),were selected for this investigation as they are both grown widely in China.Seeds were surface sterilized with 30%v/v H 2O 2for 30min and then washed thoroughly with deionized

780

Environ Geochem Health (2013)35:779–788杂交

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去离子水

adj. 受污染的;弄脏的

adj. [微] 厌氧的,[微] 厌气的;没有气而能生活的

n. [组织] 组织,薄的纱织品;面巾纸

n. [植] 根围(围绕植物根系的区域)

n. [化学] 离子

在分蘖期铁膜的形成达到顶峰分泌物

分蘖、拔节、抽穗、灌浆期

water.The seeds were then germinated in acid-washed quartz sand for 10days.Afterward,each seedling was transferred to a PVC pot and grown in 1/2-strength Hoagland’s nutrient solution for 3days and then for a further 26days in full strength solution with the following nutrient composition (l mol L -1):NH 4NO 3500,K 2SO 4200,CaCl 2400,MgSO 4á7H 2O 1,500,KH 2PO 41.3,Fe-EDTA 50,H 3BO 310,ZnSO 4á7H 2O 1.0,CuSO 4á5H 2O 1.0,MnSO 4á5H 2O 5.0,Na 2MoO 4á2H 2O 0.5,CoSO 4á7H 2O 0.25.The nutrient solution was adjusted to pH 5.8with NaOH or HCl and changed every 3days.

Pot experiment under waterlogged conditions Topsoil was collected from a paddy ?eld (0–20cm depth)at the campus of South China Agricultural University.The soil contained 9.8%organic mater,1.35g kg -1total N, 1.04g kg -1total P and 0.62g kg -1total K.The soil pH was 6.48and the total Cd and As concentration in the soil was below the detection limit (0.01mg kg -1).After being air-dried and passed through a 2-mm mesh,1kg of soil was weighed into each pot (13cm diameter and 12cm high).The soil was then used to produce the following three treatments:no Cd or As (CK),amended with 50mg Cd kg -1as CdCl 2áH 2O only (Cd treatment)and amended with 50mg As kg -1as NaHAsO 4only (As treatment).Soils were mixed thoroughly and sub-merged for 2weeks with 2cm of water above the soil surface.

Seedlings were washed carefully to remove adhering quartz sand and transplanted to pots (two plants each pot).There were 4replicates per treatment.Soil in each pot was kept submerged using deionized water.The growth temperature was controlled at 25/20°C day/night and relative humidity 60/80%day/night,with ambient light intensity (light period 16h per day with natural sunlight supplemented with sodium vapor lamps to maintain a photon ?ux density of [350l mol m -2S -1)in the glasshouse.Plants were harvested at their tillering stage (the stage when side shoots are produced,day 25),bolting stage (elongation stage,the stage when rice plants grow at greatest rate,and stems are elongating,day 46),ear emergence stage (the stage when developing ears are visible,day 72)and grain-?lling stage (the stage when grains are being ?lled,day 88),after transplant,respec-tively.The soil adhering onto roots was collected and considered as ‘rhizosphere soil.’Plant roots were then

washed in tap water to remove soil particles,followed by a ?nal washing using deionized water.Fe plaque was extracted as below,the roots and shoots were oven-dried at 60°C to constant weight and stored at 4°C,awaiting chemical analysis.

Pot experiment under deoxygenated nutrient condition

The same seeds used in the soil experiment were used in a deoxygenated nutrient experiment.Seedlings were selected and transplanted to PVC pots (c.0.8l,one plant per pot)?lled with deoxygenated 1/2-strength Hoagland’s nutrient solution containing 0.1%(w/v)agar.The pots were placed in the glasshouse,and solutions were renewed once every 6days.Measure-ments of ROL amount were taken at the same stages (days 18,40,69and 81after transplanting)as in the waterlogging experiment.

Sample analysis

DCB extraction of Fe plaque

The harvested roots were washed twice with deionized water and then incubated for 60min in 40ml cold solution of dithionite–citrate–bicarbonate (0.03M sodium citrate and 0.125M sodium bicarbonate with the addition of 0.6g sodium dithionite,DCB)(Otte et al.1989).After incubation,roots were rinsed three

times with deionized water and then added to the DCB extract.The resulting solution was made up to 100ml with deionized water for the analysis.

Chemical analysis

All plant samples harvested were weighed separately for biomass,ground to a ?ne powder (IKA A11basic;IKA-WerkeGmbH,Germany)and digested by HNO 3in a microwave oven (MARS-X;CEM,USA).The soil samples were air-dried,ground to a ?ne powder and digested with aqua regia (HCl:HNO 3=3:1,w/v).Concentrations of Cd and As in digests and DCB-extracted solutions were determined by atomic absorption spectroscopy (AAS)(for Cd)and induc-tively coupled plasma-optical emission spectrometry

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adv. 彻底地,完全地

DCB-Fe 的测定方法

(ICP-OES,Optima2000DV,Perkin Elmer,USA) (for As),respectively.Blanks,tea standard material (GBW-08303)and soil standard material(GBW-07401)(China Standard Materials Research Center, Beijing,P.R.China)were used for quality control.The Cd and As recovery rates were both90±10%.

ROL measurement

Radial oxygen loss amounts were measured using a titanium(III)citrate buffer method(Kludze et al. 1994).The actual ROL amount was calculated using the following formula:

ROL amount?c yàz

eT

where ROL is measured as l mol O2plant-1h-1,c is the initial volume of Ti3?added to each tube(in l),y is the concentration of Ti3?in the solution of the control (without plants)(in l mol Ti3?l-1),and z is the concentration of Ti3?in solution after6h treatment with plants(in l mol Ti3?in solution plant-1l-1). Statistical analysis

Data were analyzed and evaluated using the SPSS17.0 software package.Results are presented as means with standard errors attached(n=4),and means were compared by least signi?cant difference(LSD)at the 5%level.

Results and discussion

ROL and Fe plaque formation

Radial oxygen loss amounts were signi?cantly differ-ent among the growing stages,ranging on average from1.1to16l mol O2plant-1h-1(Table1).ROL increased as rice plants grew,reaching its peak at the ear emergence stage,and?nally decreased at the grain-?lling stage(Fig.1a).Our results indicated that rice roots were most active and released largest amounts of O2into rhizosphere at the ear emergence stage.Furthermore,Tianyou116had signi?cantly higher ROL than Huaxinzhan during the entire growth

cycle.This result supports those reported by Wang (2010)who found that the amounts of ROL ranged from0.32to1.11mmol O2plant-1h-1in the25rice cultivars studied.

Fe concentrations in Fe plaque increased by 2.6–5.1-fold from tillering to ear emergence stages and then decreased by56.2–73.4%at the?lling stage(Table2).Furthermore,there was a signi?cant Table1ROL of rice plants exposed to0.1%agar with 1/2-strength Hoagland’s solution at different growth stages (l mol O2plant-1h-1,mean±SE,n=4)

Growth stage Tianyou116Huaxinzhan Tillering stage 1.1±0.2a 1.3±0.1a Bolting stage8.2±1.9b 3.4±1.3b Ear emergence stage16±1.1c10±2.1c Grain-?lling stage7.6±2.2b 2.7±1.1b Different letters within the same column indicate signi?cant difference between the stages at the level of P\

0.05

Fig.1a Dynamic changes of ROL in two rice cultivars (Tianyou116and Huaxinzhan)at four growth stages(tillering stage at day18,bolting stage at day40,ear emergence stage at day69and grain-?lling stage at day81).Data are mean±SE (n=4);b relationship between ROL and Fe concentrations in Fe plaque at different growth stages

我的研究表明水稻根系在抽穗期活动和分泌氧气的能力是最强的

positive correlation detected between ROL amounts and Fe concentrations in Fe plaque (R 2=0.87,P \0.001;Fig.1b).Previous studies showed that ROL is one of the important physiological factors controlling Fe plaque formation (Taylor et al.1984;Wu et al.2012).Wetland plants with high rates of ROL tend to form more Fe plaque on root surfaces and in the rhizosphere (Li et al.2011).The present results demonstrate that rice plants with higher ROL at the ear emergence stage release more O 2from roots than the other three stages,resulting in a higher degree of oxidation of Fe(II)ferric oxide or hydroxide-forming Fe plaque on root surfaces.

Table 2Concentrations of Fe in iron plaque at different growth stages (mg kg -1D.W.,mean ±SE,n =4)

Fe concentration Tianyou 116

Huaxinzhan

CK

Tillering stage 10,886±586a 4,097±144a Bolting stage 23,698±780b 8,088±360c Ear emergence stage 35,197±1513c 20,709±512d

Grain-?lling stage 11,793±422a 6,519±172ab ?Cd

Tillering stage 9,774±329a 4,395±430a Bolting stage 20,375±608b 6,756±342ab Ear emergence stage 36,056±1983c 19,682±2460c Grain-?lling stage 9,597±500a 5,599±329ab ?As

Tillering stage 9,904±835a 4,329±216a Bolting stage 16,076±1901b 11,163±533b Ear emergence stage 26,035±2765c 15,773±1846c Grain-?lling stage

11,410±341ab

4,918±713a

Different letters within the same column and same treatment indicate signi?cant difference between the stages at the level of P \0.05

Table 3Biomass of rice plants at grain-?lling stage under different treatments (g,mean ±SE,n =4)

Tianyou 116Huaxinzhan

Shoot Root Shoot Root CK 4.2±0.30c 1.7±0.25c 3.5±0.88b 1.4±0.54b ?Cd 2.5±0.60a 1.4±0.40a 2.3±0.69a 0.9±0.13a ?As

3.2±0.62b

1.6±0.15b

2.2±0.21a

1.1±0.21a

Different letters within the same column indicate signi?cant difference between the treatments at the level of P \0.05

T a b l e 4C o n c e n t r a t i o n s o f C d a n d A s i n s h o o t ,r o o t a n d F e p l a q u e a t d i f f e r e n t g r o w t h s t a g e s (m g k g -1D .W .,m e a n ±S E ,n =4)

C d c o n c e n t r a t i o n

A s c o n c e n t r a t i o n

S h o o t

R o o t

F e p l a q u e

S h o o t

R o o t

F e p l a q u e

C K

?C d

C K

?C d

C K

?C d

C K

?A s

C K

?A s C K

?A s

T i a n y o u 116

T i l l e r i n g s t a g e

0.03±0.01a

8.2±2.5a

1.8±0.9a

86±3.0b

0.3±0.03b

2.5±1.0a

0.13±0.05a

6.9±0.8b

2.8±0.9a

199±14c

44±25a

298±49a

B o l t i n g s t a g e

0.02±0.00a

7.5±0.8a

1.2±0.4a

77±5.0a

0.1±0.04a

13±5.6b

0.22±0.08a

4.7±1.8a

3.2±0.5a

162±3.1b

52±20a

986±161c

E a r e m e r g e n c e s t a g e

0.04±0.02a

7.9±0.6a

1.7±0.3a

132±4.0c

0.2±0.2a b

24±15c

0.14±0.02a

5.3±0.3a

2.7±0.3a

112±10a

101±5b

1,396±204d

G r a i n -?l l i n g s t a g e

0.03±0.00a

8.0±1.9a

1.3±0.3a

162±0.3d

N D

1.8±0.5a

0.23±0.07a

9.2±0.8c

2.3±0.8a

105±24a

71±29a b

542±66b

H u a x i n z h a n

T i l l e r i n g s t a g e

0.02±0.01a

13±3.6a

2.5±1.0b

136±22a

0.04±0.02a

0.4±0.1a

0.23±0.05a

9.0±1.1b

2.5±1.0a b

181±8.8c

34±5.7a

267±26a b B o l t i n g s t a g e

0.05±0.01b

11±4.1a

1.3±0.6a

125±7.9a

0.05±0.04a

3.0±0.9b

0.23±0.04a

8.3±2.6b

3.3±0.6b 109±10b

93±41b

520±18b

E a r e m e r g e n c e s t a g e

0.04±0.01b

11±2.1a

2.4±1.5b

124±16a

0.1±0.08b

14±1.8c

0.14±0.08a

5.0±0.7a 2.4±0.5a 49±2.5a

104±19b 641±281b G r a i n -?l l i n g s t a g e

0.07±0.03b

10±2.0a 1.8±0.5a 138±1.8a

N D

0.9±0.3a b

0.17±0.07a

13±2.3c 1.8±1.5a

68±12a b 33±1.9a

185±36a

D i f f e r e n t l e t t e r s w i t h i n t h e s a m e c o l u m n a n d t h e s a m e c u l t i v a r i n d i c a t e s i g n i ?c a n t d i f f e r e n c e b e t w e e n t h e s t a g e s a t t h e l e v e l o f P \0.05

最近的研究表明水稻在抽穗期根系释放更多的径向氧气

Cd and As concentrations in rice plants and Fe

plaque

Rice growth was inhibited by Cd and As at most of its

growing period (Table 3).The toxicity of Cd and As

became much more important as rice grew.Rice

growth is generally suppressed when grown in soils with high levels of Cd (Zhuang et al.2009)or As

(Akter et al.2005;Williams et al.2005).Liu et al.

(2007b )reported that rice tillering,plant height,leaf

area,dry matter accumulation and grain yield were

dramatically reduced when exposed to 100mg kg -1

of Cd.Williams et al.(2005)and Mei et al.(2009)

have both shown that the grain and straw biomass of

rice were signi?cantly reduced when grown in soil

with the addition of 100mg kg -1

As.

Concentrations of Cd and As in shoots,roots and Fe

plaque of our experimental rice plants are presented

in Table 4.Cd concentrations did not show any

signi?cant differences in shoots between the growth stages,whereas in roots,the two cultivars differed.As concentrations in shoots of rice plants decreased by 23.2–44.4%from tillering to ear emergence stages,

but increased to 1.7–2.6-fold higher at the grain-?lling stage.However,As concentrations in roots continued to decrease during the four stages,ranging from 49to 199mg kg -1.Zheng et al.(2011)also reported that As concentrations in roots of rice decreased by 83.5%during growth stages,while the concentrations in leaves and internodes increased 2–3-fold after ?ow-ering to reach a peak level.There were no signi?cant differences between the four growing stages for Cd and As concentrations in rice plants in the CK treatment,and concentrations were maintained at a relatively lower level (Table 4),indicating that the control plants were not contaminated by Cd or As.Concentrations of Cd and As in Fe plaque followed similar trends in Fe concentrations,ranging from 0.4–24mg kg -1to 267–1,396mg kg -1,respectively.Furthermore,signi?cant positive correlations were detected between ROL and Cd (R 2=0.85,P \0.01;Fig.2a)and As (R 2=0.83,P \0.01;Fig.2b)con-centrations in Fe plaque.

A number of studies have reported that the presence

of Fe plaque can inhibit the uptake of phytotoxic

metal(loid)s into plant tissues due to the high adsorp-tion capacity of iron (hydr)oxides (Ye et al.2001;Batty et al.2002;Chen et al.2005).Liu et al.(2008)found that the enhancement of Fe plaque can increase the sequestration of Cd in the rhizosphere and onto root surfaces of rice,and therefore,Fe plaque may act as a buffer to reduce Cd uptake into rice roots.It has also been reported that Fe plaque plays an important role in the behavior of As on rice root surfaces,such as inhibiting translocation of As from roots to shoots (Liu et al.2004).The present results indicate that rice roots with higher ROL at the ear emergence stage tend to form more Fe plaque,which can enhance the sequestration of Cd and As in the rhizosphere or onto root surfaces,leading to less Cd and As being translocated to shoots.Cd in Fe plaque accounted for a low proportion of the total Cd amount in rice plants from tillering to grain-?lling stages,ranging from 0.2to 13.9%.Most Cd was accumulated in root tissues (Fig.3a).The ?nding was in line with previous studies (Liu et al.2007a ,2008)which report that root tissues rather than Fe plaque on the root surfaces are a major barrier

to Fig.2Relationships between ROL and concentrations of Cd (a )and As (b )in Fe plaque at different growth stages 784

Environ Geochem Health (2013)35:779–788分蘖

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节间存在

组织

氢氧化物

隔离

根际

v. 抑制;控制(inhibit 的过去分词)

v. 镇压,禁止(suppress 的过去时和过去分词)adj. 抑制的,发育不全的adj. 植物性毒素的

铁膜可以充当一个阻挡Cd 进入水稻根系的缓冲区域

n. 障碍物,屏障;界线

Cd uptake and translocation within rice plants.How-ever,Liu et al.(2007a )suggested that the uptake and

translocation of Cd appears to be related to Fe nutritional levels in the plants.Therefore,Fe plaque may act not only as a Cd ‘barrier’which can to some extent sequester Cd on root surfaces,but also as a Fe ‘reservoir’which can provide added Fe nutrition to plants to diminish the effects of the Cd.Unlike Cd,As was primarily absorbed in Fe plaque on root surfaces,ranging from 55.7to 92.0%of the total As amount in the rice plants (Fig.3b).A similar phenomenon was also observed by Liu et al.(2004)who found that As in DCB-extracted solutions accounted for 71.7–89.0%of total As in rice plants.It has been reported that most (70–80%)of the As in the Fe plaque on the roots of rice is As(V),with the remaining (20–30%)As being As(III)(Liu et al.2006).The presence of Fe plaque enhances As(III)and decreases As(V)uptake by rice

Tillering stage

Bolting stage

Ear emergence stage

Grain-filling stage

Tillering stage

Bolting stage

Ear emergence

stage

Grain-filling stage

Root Cd

Root As

Environ Geochem Health (2013)35:779–788785障碍

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容器

使减少

(Chen et al.2005).Furthermore,Zhao et al.(2009)

reported that Fe plaque might be responsible for the

oxidation of arsenite to arsenate and thus reducing the

toxicity of As contamination in soil–plant systems.

These ?ndings all suggest that Fe plaque may serve as

a barrier which could both weaken the toxic effects of

As through oxidation of As(III)to As(V)and reduce

the uptake of As [especially As(V)]into root tissues.

Moreover,the highest proportion of Cd and As in Fe

plaque in our experiment was observed at ear emer-gence stage,suggesting that this would be an ef?cient

way to reduce the accumulation of Cd and As in rice

plants at this stage.It is noteworthy that more of both Cd and As were accumulated in shoots of Huaxinzhan than in Tianyou

116(P \0.05)during the entire growth period (Table 4).Previous studies indicated that Cd (Liu et al.2005;Wang et al.2011)and As (Zhu et al.2008a ;Mei et al.2009)levels in both straw and grains varied greatly between different cultivars.Considering that

Tianyou 116had signi?cantly higher ROL than

Huaxinzhan,the data suggest that a rice cultivar with

higher ROL is inclined to accumulate less Cd and As in its shoots.The results from our study are also supported by Wang et al.(2011)and Mei et al.(2009,

2012)who found that rice cultivars with higher rates of

ROL had higher capacities for limiting the transfer of Cd and As to straw and grain.Thus,it can be concluded that,by using cultivars with high ROL and

strong plaque-forming abilities,and also taking some

measures at the ear emergence stage to stimulate more Fe plaque formed (such as adding Fe),it may be possible to reduce the transfer of Cd and As to above-ground parts of rice plants growing in contaminated areas.Conclusions

Our study has shown that signi?cant differences exist

in ROL,Fe plaque formation,Cd and As in root tissues and in Fe plaque between the four growth stages.There were signi?cant positive correlations between ROL and concentrations of Fe,Cd and As in Fe plaque on rice root surfaces.Higher ROL at the ear emergence stage induced more Fe plaque formation on roots,which in turn increase the amount of Cd and As absorbed on Fe plaque,leading to a lesser proportion of Cd and As accumulated within plant

tissues of rice.A rice cultivar with higher ROL tends to enhance Fe plaque formation,reducing the accu-mulation of Cd and As in shoots.The present study contributes to our understanding of the dynamic changes of ROL,Fe plaque formation and their relationships with Cd and As in the Fe plaque and shows that the ear emergence stage could be an important period to reduce the accumulation of Cd and As in rice.Acknowledgments We sincerely thank Prof.HQ Zhou (Guangdong Rice Research Institute)for providing the rice

seeds.We are also grateful to the National Natural Science

Foundation of China (30770417),the National ‘863’project of China (2013AA062609),the Natural Science Foundation of

Guangdong (07003650)and Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education

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我喜欢微笑。不管是我对别人,还是别人对我,微笑同样真切。我也喜欢音乐。从古典优美的声音中,我觉得一切都那么充一切都那么完美。我喜欢朋友。他们在我倾诉烦恼时认真地听,开导我,每当这时,我就觉得什么东西流入心底,十分温暖。 我喜欢生活,感激一切让我喜欢的事物! 我喜欢作文(二): 我喜欢 我喜欢清澈的小溪,喜欢小溪叮咚的声音;但更喜欢将我的脚,当作小木桨,在水面上轻轻划过;让那清爽的感觉,流进我的血液。 我喜欢郁郁葱葱的森林,喜欢森林清鲜的空气;但更喜欢将我的手,当作一阵风,在树身上慢慢拂过:让那自然的感觉,传播我的身体。 我喜欢明朗的天空,喜欢天空偶尔的几声鸟叫;但更喜欢将我的耳朵,当作收音机,在天空中悄悄收着;让那清脆的声响,传送我的四肢。 我喜欢广阔的大地,喜欢大地上的一花一草;但更喜欢将我的眼睛,当作望眼镜,在大地上静静地看着:让那美丽的风景,镶在我的大脑。

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有许多小朋友来的时候都哭了,但我没哭,我比同龄人都坚强,我理解妈妈,她是为了我, 为了我能过上更好的生活妈妈每逢过年过节,都会回来看我,我觉得这样就够了,三年 了,我见到妈妈的次数用手指头都能查过来这样我上了初中,每天都在别人家吃住。看 着别人的脸色,这些年我受到的委屈,她都不知道,我也不想告诉她,少女的心事,不想被 人知道,我在梦中经常喊到的就是她的名字,醒来时,枕头已被泪水浸湿,我无数次因为太 想她,而掉眼泪,终于我自甘堕落,上课开始溜号,注意力开始不集中,老师注意到我的变 化,找我长谈,我什么都没有和老师说,老师把电话打到妈妈那里,第二天下午,她回来了, 她不分青红皂白,劈头盖脸一顿说,说我不上进,不省心,我也忍不住了,积压在心里多年 的情绪终于爆发。我大喊这么多年你管过我吗?你只给我物质上的满足,可你知道我要的不 是这些,你记得我的生日吗?你给我过过生日吗?你给我参加过家长会吗?说完我转身就走, 不给她解释的机会晚上,她回来了,看见我在洗衣服,她说你长大了,都会洗衣服了我

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专题05 感悟类作文-中考语文高分作文之道破解版

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心愿作文700字_心愿作文

心愿作文700字_心愿作文 心愿作文700字 “我亲爱的人类伙伴们,你们好!”就在那不远处的小山丘上,传来了低吟声。在朦胧的月光帷幕中,有一个矮小的身影闪现着。那是一棵秃头树。它的枝叶也枯黄了,凋零了,枝干上光秃秃的,就像一个秃头。树腰上也有了一个个蛀洞,手臂也折了,留下来一道丑陋的刀疤。 “我是一棵年轻的树。”或许令人大吃一惊,但它的话听起来有几分伤感,“以前我也曾羽叶丰满、翠绿,身材高大、威猛,是这片森林的绿色卫士。以前这片森林绿树成荫,百草丰茂。我和我的兄弟姐妹们守望相助,过着无忧无虑的生活,当地的百姓也很善待我们。我们也常常帮助他们净化空气,改善环境。” “可随着经济科学的发达,人类发明出了一种叫锯子的工具,天天拿着它上山给我们做人工手术,把我们的兄弟姐妹整得不是手残,就是臂断。我亲眼见证着一条条无辜的生命在这片乐土倒下…”说到

这里,秃头树掉下了一串晶莹的眼泪,“树的同胞们都知道了‘锯子’是比‘蛀虫’还厉害的东西。于是,从此以后,我们一直过着提心吊胆的生活,我们的家族日渐人丁稀少。我肝肠寸断,也渐渐消瘦了下来。每天我遭受着百般折磨,打出了腰间盘突出,变得年纪轻轻就像一个老头子,整天弓着背,下雨天会隐隐作痛。手也被锯断了。”,说着,秃头树皱起了眉头,敲起了那弯着的背,长长地吁了一口气,“你们人类都在说‘笑一笑,十年少;愁一愁,白了头’,所以,我的叶子渐渐枯黄掉落。” 秃头树说着轻笑了一下:“城市既然拒绝了森林。同时,他们也会遭到了大自然的惩罚。近年来,人类遭遇水土流失,泥石流袭击,沙尘暴席卷,这不是自然灾害,而是你们自作自受啊!看着一棵棵参天大树长老的倒下,闻着从窗户中冒出的滚滚浓烟的臭味,令我身心憔悴。没有了我们,天空也灰暗,河流也混沌,高山也光秃。一串串灾难接踵而至,你们还不悔悟吗?人类既然来自于我们森林,也离不开我们,我们是你们赖以生存的自然屏障。”说着,秃头树望了望四周的

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