You are on page 1of 9

Plant Soil (2013) 370:615–623

DOI 10.1007/s11104-013-1661-9

REGULAR ARTICLE

Occluded C in rice phytoliths: implications to biogeochemical


carbon sequestration
Zimin Li & Zhaoliang Song & Jeffrey F. Parr &
Hailong Wang

Received: 17 October 2012 / Accepted: 25 February 2013 / Published online: 8 March 2013
# Springer Science+Business Media Dordrecht 2013

Abstract (leaf, stem, sheath and grains) on a dry weight basis in 5


Aims Carbon (C) bio-sequestration within the phytoliths rice cultivars range from 0.4 mgg−1 to 2.8 mgg−1, and the
of plants, a mechanism of long-term biogeochemical C C content of phytoliths from grains is much lower than
sequestration, may play a major role in the global C cycle that of leaf, stem and sheath. The data also show that the
and climate change. In this study, we explored the poten- PhytOC content of rice depends on both the content of
tial of C bio-sequestration within phytoliths produced in phytoliths and the efficiency of C occlusion within
cultivated rice (Oryza sativa), a well known silicon phytoliths during rice growth. The biogeochemical C
accumulator. sequestration flux of phytoliths in 5 rice cultivars is ap-
Methods The rice phytolith extraction was undertaken proximately 0.03–0.13 Mg of carbon dioxide (CO2)
with microwave digestion procedures and the determi- equivalents (Mg-e-CO2) ha−1 year−1. From 1950 to 2010,
nation of occluded C in phytoliths was based on about 2.37×108 Mg of CO2 equivalents might have been
dissolution methods of phytolith-Si. sequestrated within the rice phytoliths in China. Assuming
Results Chemical analysis indicates that the phytolith- a maximum phytoliths C bio-sequestration flux of
occluded C (PhytOC) contents of the different organs 0.13 Mg-e-CO2 ha−1 year−1, the global annual potential

Responsible Editor: Hans Lambers.

Z. Li : Z. Song (*) : H. Wang Z. Song


School of Environmental and Resource Sciences, College of Urban and Environmental
Zhejiang A & F University, No. 88 North Huancheng Road, Sciences, Peking University,
Lin’an, Zhejiang 311300, China Peking 100871, China
e-mail: songzhaoliang78@163.com

Z. Song : H. Wang
Zhejiang Provincial Key Laboratory of Carbon
Cycling in Forest Ecosystems and Carbon
Sequestration, Zhejiang A & F University,
Lin’an, Zhejiang 311300, China
J. F. Parr
Z. Song Southern Cross GeoScience,
Laboratory for Earth Surface Processes, Southern Cross University, Lismore,
Ministry of Education, Peking 100871, China NSW 2480, Australia
616 Plant Soil (2013) 370:615–623

rate of CO2 sequestrated in rice phytoliths would approx- and Parry 1981; Pearsall 1989; Hart and Humphreys
imately be 1.94×107 Mg. 1997; Parr 2006; Bowdery 2007). For example, it has
Conclusions Therefore rice crops may play a signifi- been reported that phytoliths in volcanic soils and
cant role in long-term C sequestration through the peatland sediments range in age from 0 to 8,000 year
formation of PhytOC. BP (Parr and Sullivan 2005). Wilding (1967) reported a
radiocarbon date of 13300±450 year BP for phytoliths.
Keywords Carbon sequestration . PhytOC . During the formation of phytoliths, some organic C
Phytolith . Rice can be occluded (Jones and Milne 1963). Compared
with other organic C fractions, phytolith occluded C
(PhytOC) is relatively stable and can persist in the soil
Introduction for millennia due to its strong resistance to decompo-
sition (Wilding et al. 1967; Wilding and Drees 1974;
Climate change is one of the major global environ- Mulholland and Prior 1993; Strömberg 2004; Parr and
mental issues (Schlesinger 1997). Increased green- Sullivan 2005; Prasad et al. 2005). PhytOC plays a
house gas (GHG) emissions are now widely accepted major role in soil C cycle (Parr and Sullivan 2005;
as the main cause of climate change since the Oldenburg et al. 2008) and the importance of the soil
Industrial Revolution (Schimel 1995; Falkowski et C cycle in relation to climate change has been empha-
al. 2000; IPCC 2007; Kosten et al. 2010). Recent sized by many researchers (Gifford 1994; Falkowski
research indicates that the global CO2 emission rate et al. 2000; Kosten et al. 2010). As an important part
had increased to 3.11× 1011 Mg per year by 2010 of terrestrial C, it can represent up to 82 % of total C in
(DOE 2008). With this in mind all mechanisms of some soil and sediments after 2,000 years of litter leaf
carbon (C) sequestration should be explored in order decomposition (Parr and Sullivan 2005). Recent re-
to identify relevant practical measures that may be search also indicates that plant phytoliths have great
taken to reduce the concentration of CO2 in atmo- potential to sequester CO2 from the atmosphere (Parr
sphere (Parr et al. 2010). Existing forestry, grasslands, et al. 2009; Parr et al. 2010; Parr and Sullivan 2011;
shrub lands and wetland plants play a key role in the Zuo and Lü 2011). For example, the phytolith C bio-
sequestration of atmospheric CO2. A large part of the sequestration fluxes from millet, wheat and sugarcane
C sequestrated by plants is returned into the soil as range up to 0.04, 0.25 and 0.36 Mg-e-CO2 ha−1 year−1,
plant residues (Harrison et al. 1993; Parr and Sullivan respectively (Parr et al. 2009; Parr and Sullivan 2011;
2005; Fang et al. 2007; Field 2001). It is estimated that Zuo and Lü 2011). Moreover, the flux of C occluded
soils contain about 1550 Pg inorganic C and 950 Pg within the phytoliths of bamboo species examined to
organic C (Zuo and Lü 2011). A minor variation in date ranges up to 0.71 Mg-e-CO2 ha−1 year−1, and,
such big C pool in terrestrial ecosystems may have a with current global bamboo forests covering an area
significant effect on the C flux (Harrison et al. 1993). of around 22 million ha could potentially be securely
However, due to environmental complexities, it is sequestering~1.56×107 Mg of atmospheric CO2 per
difficult to quantify soil C changes (McKenzie et al. year. Parr et al. (2010), have suggested that if all
2000; Skjemstad et al. 2000; Clark 2002). potentially arable land (4.1 billion ha) is exploited to
Phytoliths, the opaline amorphous silica formed in grow bamboo or other similar grass crops, the global
plant tissues (e.g., cell walls, cell lumina, and intercellular potential of phytolith C bio-sequestration is approxi-
spaces typically near evaporating surfaces), are present in mately 1.5×109 Mg CO2 per year. This would result in
most plants (Perry et al. 1987; Piperno 1988; Song et al. effectively reducing global CO2 emissions by a rate
2012a, b). When plants die and undergo decomposition, equivalent to 11 % of the current increased CO2 in
large amounts of these phytoliths are released directly atmosphere (Parr et al. 2010).
into the soil. When phytoliths encounter some Cultivated rice (Oryza sativa) is a main food source for
unpredictable situations, such as earthquake, dust storms, more that 50 % of the global population and is currently
flood (Baker et al. 1961), forest/grassland fire, erosion, cultivated on around 1.55×108 ha (IRRI 2011). China is
etc., they are often preserved in the environment (Baker the largest country involved in rice production, accounting
1961; Jones and Milne 1963; Wilding et al. 1967; Jones for up to 35 % of all world production, followed by India
and Handreck 1967; Wilding and Drees 1974; Sangster (22 %), Indonesia (9 %) and Bangladesh (7 %) (IRRI
Plant Soil (2013) 370:615–623 617

2011). There is a long history of rice production in (Parr et al. 2010). Each phytolith extraction solution
China (Cao et al. 2007) and its current area of produc- was used 0.8 mol/l potassium dichromate to examine
tion is around 2.96×107 ha (NBSC 2011). Rice is able to the extraneous organic materials outside of the phytolith
be cultivated twice per year under the high temperature cells. If the color of solution would not change within
and abundant rainfall environment of south-eastern 5 min, it showed that the extraneous organic materials
China (Cao and Zhang 2004; Lin et al. 2004). outside of the phytoliths were thoroughly removed. The
Recently the C sequestration potential of phytoliths phytoliths extracted were oven-dried at 75 °C for 24 h in
within crops of millet (Zuo and Lü 2011) and wheat a centrifuge tube of known weight. The samples were
(Parr and Sullivan 2011) has been reported. However, allowed to cool and then weighed to obtain the phytolith
the variability of PhytOC in rice has not been investi- quantities. Each sample was checked with an optical
gated yet, even though cultivated rice is a well known microscope (Olympus CX31, Japan) to further ensure
silica accumulator (Epstein 1994, 1999). Therefore, the that all extraneous organic materials outside of the
objective of this study is to explore the potential of C phytoliths were thoroughly removed (Murphy 2002).
occluded within phytoliths in rice cultivars. Based on methods of Kroger et al. (2002), the dried
phytoliths samples were treated with 1 mol/L HF at 45 °
C for 60 min to dissolve phytolith-Si. The organic C
Materials and methods released from phytoliths after HF treatment was dried at
45 °C and determined for C content using the classical
Collection of rice plant materials potassium dichromate method (Lu 2000). The organic C
data was monitored with standard soil samples of
In this study, the rice plants used in experiment were GBW07405. The precision is better than 7 %.
grown under the same conditions to eliminate factors
that might influence silica uptake and deposition. The
stem, leaf, sheath and grains from 5 commonly planted Results
rice cultivars (Table 1) were used to determine the
content of phytoliths and PhytOC in different organs, Microscopy check show that all external organic mate-
and to analyze the relationship of two aspects in dif- rials were completely removed (Fig. 1). The contents of
ferent rice cultivars and organs of the same rice plant. phytoliths in different rice organs (leaf, stem, sheath,
The rice plant samples (three replicates) of different grains) range from 15 mgg−1 to 144 mgg−1 and show a
cultivars were collected during the harvest season in similar trend (sheath>leaf>stem>grains) in 5 rice cul-
October, 2010, from the regional trials of new varieties tivars (Table 1). There are substantial variations in the C
of crops grown at the demonstration base of Zhejiang content (14 mgg−1–34 mgg−1) of phytoliths from organs
Soil and Fertilizer Station (30°56′06.3″N and 120°51′ in rice cultivars (Table 1). Generally, the C content of
52.9″E) in Jiaxing, Zhejiang Province, southeast China. phytoliths from the stem is higher than that of sheath,
The Base is located in Hangjiahu Plain which experi- grains and leaf (Table 1). The PhytOC contents of
ences a typical subtropical humid monsoon climate, organs on a dry weight basis in the 5 rice cultivars vary
with an average annual precipitation of 1,200 mm. The from 0.4 mgg−1 to 2.8 mgg−1 and have a consistent
annual mean temperature is 16 °C and the number of trend (sheath>leaf>steam>grains) (Table 1).
frost free days is 230. Soil is classified as the Earth- A weak negative correlation exists between the con-
cumuli-Orthic Anthrosols (CSSD 2012). tent of phytoliths and the C content of phytoliths from
different organs in 5 rice cultivars (R2 =0.2236, p>0.05)
Phytoliths occluded C analysis (Fig. 2). However, there are strong positive correlations
between the content of phytoliths and the PhytOC
In the laboratory, the phytolith extraction of rice or- content of different organs (R2 = 0.8651, p < 0.01)
gans (around 50 mg each test) was undertaken with (Fig. 3). The correlation (R2 =0.2733, p>0.05) between
microwave digestion procedures (Parr et al. 2001). the C content of phytoliths and the PhytOC content of
This process was followed by a Walkley–Black type grains is weaker than that of leaf (R2 =0.8241, p<0.01),
digest (Walkley and Black 1934) to thoroughly re- stem (R2 =0.9214, p<0.01) and sheath (R2 =0.5168,
move extraneous organic materials in the samples p<0.05) (Fig. 4).
618 Plant Soil (2013) 370:615–623

Table 1 Rice cultivars and organs, the phytolith content as a equivalents (Mg-e-CO2) for rice (according to grains yields of
percentage of organs biomass, the PhytOC content of the rice single and double rice crops between 9.3 and 18.6 Mg-ha−1)
phytoliths, the content of PhytOC in organs on a dry weight (data offered by Zhejiang Soil and Fertilizer Station)
basis and the estimated fluxes of PhytOC per ha in Mg of CO2

Rice Rice Phytolith contents PhytOC of phytoliths PhytOC of organs Estimated PhytOC fluxes
cultivars organs Mean (s. d) mgg−1 Mean (s. d) mgg−1 Mean (s. d) mgg−1 (Mg-e-CO2 ha−1 year−1)

Xiushui 09 grains 20.56 (0.26) 20.16 (1.05) 0.41 (0.03) 0.04–0.10


stem 30.87 (3.36) 21.06 (1.16) 0.65 (0.11)
sheath 132.95 (5.67) 21.27 (1.46) 2.83 (0.30)
leaf 55.47 (8.16) 18.66 (3.05) 1.04 (0.32)
Ning 81 grains 26.57 (3.86) 18.47 (1.57) 0.49 (0.13) 0.03–0.12
stem 37.36 (11.56) 28.85 (8.36) 1.08 (0.65)
sheath 108.87 (3.76) 19.56 (1.14) 2.13 (0.20)
leaf 64.57 (9.18) 24.46 (1.36) 1.58 (0.32)
Xianghu 301 grains 27.23 (5.76) 23.47 (0.14) 0.63 (0.14) 0.05–0.13
stem 39.12 (2.44) 33.56 (0.36) 1.31 (0.09)
sheath 118.57 (5.76) 20.47 (0.13) 2.43 (0.14)
leaf 77.87 (2.03) 22.13 (0.37) 1.72 (0.07)
Zhejing 37 grains 15.47 (1.48) 23.16 (0.27) 0.36 (0.04) 0.04–0.08
stem 40.23 (2.27) 24.75 (0.46) 1.00 (0.08)
sheath 103.06 (2.37) 22.14 (1.14) 2.28 (0.16)
leaf 79.27 (5.24) 25.67 (1.47) 2.03 (0.25)
Jiahua 11 grains 21.57 (2.14) 26.16 (2.04) 0.56 (0.09) 0.04–0.11
stem 39.95 (5.04) 27.27 (1.47) 1.09 (0.20)
sheath 143.95 (6.96) 14.18 (1.68) 2.04 (0.34)
leaf 74.57 (1.14) 23.01 (0.37) 1.72 (0.03)

Discussion millet has no direct relationship with the actual content


of silica (phytoliths) taken up by the plant, and mainly
The mechanism of C occlusion within rice phytoliths depends on the efficiency of the C occluded within
and its application phytoliths during plant growth. However, our study
results on C occlusion within rice phytoliths are quite
A number of researchers (e.g., Parr et al. 2009, 2010; different from the above studies. The strong positive
Parr and Sullivan 2011; Zuo and Lü 2011) found that correlations between the phytolith content and the
the PhytOC content in bamboo, wheat, sugarcane and PhytOC content of organs in 5 rice caltivars tested

Fig. 1 Rice phytoliths


extracted by the microwave
method followed by a
Walkley–Black digest using
standard light microscopy
(Walkley and Black 1934;
Parr et al. 2001)
Plant Soil (2013) 370:615–623 619

36 3.0
leaf
y = 0.0494x + 1.304 stem
y = -0.051x + 26.09 2 sheath
2 R = 0.5168, P < 0.05 grains
R = 0.2241, P > 0.05
27

PhytOC content in organs


C content of phytolith

2.0
y = 0.0474x - 0.2274

( mg g )
-1
2
y = 0.1247x - 1.242 R = 0.9214, P < 0.01
(mg g )

18
-1

2
R = 0.8241, P< 0.01

1.0
9
y = 0.0174x + 0.1127
2
R = 0.2733, P > 0.05

0 0.0
0 40 80 120 160 0 5 10 15 20 25 30 35 40
-1 C content of phytoliths (mg g-1)
Phytolith content (mg g )

Fig. 2 Correlation of the phytolith content of organs and the C Fig. 4 Correlation of PhytOC content of organs and the C
content of phytoliths in 5 rice cultivars content of phytoliths in 5 rice cultivars. (PhytOC content in
organs as a function of the C content of phytoliths)

(R2 =8651, P<0.01) (Fig. 3), and between the C con-


2001; Ding et al. 2005; Parr and Sullivan 2011; Zuo
tent of the phytoliths in each organ and the PhytOC
and Lü 2011). The different shape of phytoliths be-
content of organs in 5 rice caltivars (R2 =0.2733–
tween different organs (e.g., between rice stem and
0.9214) (Fig. 4) indicate that the PhytOC content in
rice sheath) may also cause differences in the C con-
rice plants might depend on both the content of
tent of phytoliths because of differences in specific
phytoliths and the nature of silica occluding C within
surface area (Table 1; Bartoli and Wilding 1980;
cells of the phytoliths during plant growth. Thus, all
Bartoli 1985). However, these indirect factors remain
factors influencing the content of phytoliths and the
to be examined in further studies.
content of silica occluding C within phytoliths could
It is possible to improve the sequestration rate of
result in significant variations of the PhytOC content
PhytOC simply by selecting cultivars of high-
in plants. For example, factors such as varieties, loca-
phytolith content and high-C content of phytoliths
tion, disease resistance, and fertilizer requirements can
for cropping (Parr et al. 2010; Parr and Sullivan
play a major role in accumulating phytoliths during
2011). However, as pointed out by Parr et al. (2009)
plant growth (Ma et al. 2002; Korndorfer and Lepsch
cultivars cannot be selected solely on the basis of their
PhytOC content but rather should be considered in
combination with other desirable traits such as bio-
3.0
mass and yield. In addition, other limiting factors,
y = 0.0166x + 0.315
2 such as differences in location and climatic conditions
R = 0.8651, P < 0.01
(Parr et al. 2009; Parr and Sullivan 2011), and taste
PhytOC content in organs

2.0 preferences also need to be considered. As we have


shown in this study for rice there is a strong relation-
(mg g )
-1

ship between silica (phytolith) production and PhytOC


content (Figs. 3 and 4) thus even if rice crops with a
1.0 relatively weak ability for occluding C in their
phytoliths are planted, it may still be possible to
enhence the PhytOC content for the plants by regulat-
ing silicon supply during growing period. For exam-
0.0
0 40 80 120 160 ple, previous studies have demonstrated that the
Phytolith content (mg g-1) content of the silica (phytoliths) in crops may be
enhanced through adding manganese zinc silicon fer-
Fig. 3 Correlation of the phytolith content of organs and
PhytOC content of organs in 5 rice cultivars. (PhytOC content tilizer, straw (Bao et al. 1996), organic fertilizer,
in organs as a function of phytolith content) calcium-magnesium phosphate fertilizer (Chen et al.
620 Plant Soil (2013) 370:615–623

Table 2 Comparison of PhytOC contents in plant tissues, estimated PhytOC fluxes per hat-CO2 equivalents and global total PhytOC
sequestration rate in different plants

Plants species PhytOC contents of dry PhytOC sequestration fluxes Global PhytOC sequestration References
material (mgg−1) (Mg-e-CO2 ha−1-year−1) rates (Mg -e-CO2 -year−1)

Rice 0.4–2.8 0.03–0.13 1.94×107 This study


Bamboo 2.4–5.2 0.01–0.71 1.56×107 Parr et al. 2010
Sugarcane 3.1–15.4 0.12–0.36 0.72×107 Parr et al. 2009
Wheat 0.6–6.0 0.01–0.25 5.3×107 Parr and Sullivan 2011
Millet 0.4–2.7 0.01–0.04 0.27×107 Zuo and Lü 2011

2008), slag mucks (Zhang et al. 2008), and silicon grains of the single and double rice cropping systems
fertilizers (Matichenkov et al. 1999; Alvarez and (Anthoni et al. 2004; Zhang et al. 2010), we estimate that
Datnoff 2001; Ma and Takahashi 2002; Liang et al. the C flux of rice phytoliths is 0.03–0.13 Mg-e-CO2 ha−1-
2006; Mecfel et al. 2007) during the plant growth. In year−1. Compared with the other studies (Table 2), the flux
fact, silica (phytolith) accumulation also provides plants of the phytolith C sequestration in this study is noticeably
with some competitive advantages during the crops lower than that of bamboo, wheat and sugarcane (Parr et
growth, such as enhancement of yield and growth, re- al. 2009; Parr et al. 2010; Parr and Sullivan 2011). The
sistance to disease and increased shoot rigidity, etc. main causes for this significant difference in PhytOC
(Epstein 2001) through the regulation of silicon nutrient content may be as follows: a) the C content of phytoliths
supply for crops. It is likely to enhance PhytOC content in the 5 rice cultivars tested is simply not as high as that for
in crops through regulating silicon nutrient supply for bamboo, wheat and sugarcane; b) the rice produces less
crop plants in agricultural production. biomass than other phytolith species, e.g., bamboo leaf
(Parr et al. 2010), perhaps rice cultivars with stronger
PhytOC traits are yet to be discovered or c) the method
Carbon sequestration potential through C occlusion of C quantification used in this study (Lu 2000) which
within the phytoliths in rice plants varied considerably from previous studies (Parr et al.
2009; Parr et al. 2010; Parr and Sullivan 2011) provides
Using the PhytOC content of rice plant material on a dry a significantly different result; d) Genetic reasons and
weight basis (leaf, stem, sheath, grains) (Wang et al. 2008) different physiology/relationship to water may affect the
and the mean annual production (9.3–18.6 Mg-ha−1) of

4.0
40
C occlusion rates (10 Mg-e-CO2)

35
3.0
Rice-planted area (10 ha)

6
6

30

2.0

25

1.0
20
1950 1960 1970 1980 1990 2000 2010
1950 1960 1970 1980 1990 2000 2010
Fig. 6 Estimated the lowest and highest potential rate of CO2
Fig. 5 The variation trend of rice planted-areas from 1950 to sequestration through PhytOC accumulation in rice plants between
2010 in China (NBSC 2011 and CSDN 2011) 1950 and 2010 in China
Plant Soil (2013) 370:615–623 621

C content of phytoliths in different plants. These are areas, content of phytoliths in grains is much lower than that of
which, while beyond the scope of the current study, other organs, such as leaf, stem and sheath. The data also
necessarily require further investigation. show that the PhytOC content in rice mainly depends on
According to the records for the area of rice planted both the content of silica (phytoliths) and the efficiency
from 1950 to 2010 in China (NBSC 2011 and CSDN of the C occluded within phytoliths during rice growth.
2011) (Fig. 5), we have estimated the trend of the lowest Based on our results, it is estimated that the flux of C
and highest potential rates of CO2 occluded within rice occluded within rice phytoliths is between 0.03 and
phytoliths from 1950 to 2010 (Fig. 6). Our results estimate 0.13 t-e-ha−1-year−1. Rice may sequester between
that between 0.81×106 and 3.88×106 Mg-e-CO2 per year 0.81×106 and 3.88×106 Mg-e-CO2 annually and up to
is occluded within the rice phytoliths in China. Applying 2.37×108 Mg-e-CO2 within phytoliths of cultivated rice
the largest flux (0.13 Mg-e-CO2 ha−1 year−1) of the largest during a 60-year period in China. According to the
PhytOC sequestration flux in this study, our results indi- global rice planting area (1.55×108 ha), we estimate that
cate that rice phytoliths have potentially occluded around 1.94×107 Mg-e-CO2 from the atmosphere could have
2.37×108 Mg-e- CO2 during the past 60 years. In 2010, been sequestered in rice phytoliths annually, which
the rice planting-area of the world was around 1.55×108 could play an important role in mitigation of global
ha (IRRI 2011). Taking the largest flux (0.13 Mg-e-CO2 greenhouse gas emission. However, more studies on
ha−1 year−1) of rice phytoliths, about 1.94×107 Mg-e-CO2 the capacity of the PhytOC accumulation in arable crops
per year would have been sequestrated in rice phytoliths are needed to quantify potential of phytoliths in global C
globally. Although the annual CO2 occlusion within the sequestration and to reduce greenhouse gas concentra-
rice phytoliths of unit area is likely to be lower than that of tion in the atmosphere.
other plants such as bamboo, wheat and sugarcane
(Table 2), based on the total area of rice production the Acknowledgments We are grateful for support from National
global CO2 sequestration (1.94×107 Mg-e-CO2 year−1) in Natural Science Foundation of China (Grant No. 41103042), Zhe-
rice phytoliths is greater than those of reported, for exam- jiang Province Key Science and Technology Innovation Team
(NO.2010R50030), Zhejiang Provincial Natural Science Founda-
ple, bamboo leaf litter (1.56×107 Mg-e-CO2 year−1) (Parr tion Program (Grant No. Y5080110 and Z5080203), Opening Pro-
et al. 2010) and sugarcane leaf (0.72×106 Mg-e-CO2 ject of State Key Laboratory of Environmental Geochemistry
year−1) (Parr et al. 2009). When rice straw is returned to (SKLEG9011), Opening Project of Ministry of Education Labora-
the paddy fields, phytoliths are released into the soil after tory for Earth Surface Processes, Peking University. We thank Pr.
Dr. Zhihong Cao and Miss Fang Huang for their help in sampling.
straw decomposition. The PhytOC from rice is very stable
and can be preserved in soils for more than 6,000 years
(Cao et al. 2006, 2007; Zheng et al. 2003). For example,
fan type (bulliform cell) rice phytoliths could be found References
intact in ancient paddy soil (6,280 a BP) at the Chuodun
site (Cao et al. 2007). Thus, the PhytOC can be considered
Alvarez J, Datnoff LE (2001) The economic potential of silicon
as an important part of soil stable organic C and plays an for integrated management and sustainable rice production.
important role in long-term C sequestration (Parr and Crop Prot 20:43–48
Sullivan 2005) and mitigation of global climate change Anthoni PM, Freibauer A, Kolle O, Schulze E (2004) Winter
(Parr et al. 2010; Parr and Sullivan 2011; Zuo and Lü wheat carbon exchange in Thuringia, Germany. Agr For
Meteorol 121:55–67
2011). It is especially important to further quantify and Baker G (1961) Opal phytoliths and adventitious mineral parti-
enhance the potential of PhytOC of cultivated plants such cles in Wheat dust. CSIRO, Melbourne
as rice and other arable crops under different soil and Baker G, Jones LHP, Wardro ID (1961) Opal phytoliths and
climate conditions. mineral particles in the rumen of sheep. Aust J Agric Res
12:462–471
Bao SD, Yang XR, Li XQ, Zhang MJ (1996) The effect of wheat
yields on silicon nutrition and the zinc silicon fertilizer in
Conclusions calcareous soils. Soil 6:311–315 (In Chinese)
Bartoli F (1985) Crystallochemistry and surface properties of
biogenic opal. J Soil Sci 36:335–350
Our study reveals that the PhytOC content of the differ- Bartoli F, Wilding LP (1980) Dissolution of biogenic opal as a
ent organs (leaf, stem, sheath and grains) in 5 rice function of its physical and chemical properties. Soil Sci
cultivars ranged from 0.4 mgg−1 to 2.8 mgg−1. The C Soc Am J 44:873–878
622 Plant Soil (2013) 370:615–623

Bowdery D (2007) Phytolith analysis: sheep, diet and fecal International Rice Research Institute (IRRI) (2011) http://
material at Ambathala Pastoral Station, Queensland, Aus- beta.irri.org/
tralia. In: Madella M, Débora Z (eds) Plant, people and Jones L, Handreck K (1967) Silica in soils, plants and animals.
places–recent studies in phytolith analysis. Oxbow, Oxford Adv Agron 19:107–149
Cao ZH, Zhang HC (2004) Phosphorus losses to water from Jones LHP, Milne AA (1963) Studies of silica in the oat plant.
lowland rice fields under rice-wheat double cropping sys- Plant Soil XVIII:207–220
tem in the Tai Lake region. Environ Geochem Health Korndorfer GH, Lepsch I (2001) Effect of silicon on plant growth
26:229–236 and crop yield. In: Datnoff LE, Snyder GH, Korndorfer GH
Cao ZH, Ding JL, Hu ZY, Knicker H, Kögel-Knabner I, Yang (eds) Silicon in agriculture. Elsevier Science B V, Amsterdam
LZ, Yin R, Lin XG, Dong YH (2006) Ancient paddy soils Kosten S, Roland F, Da Motta Marques DML, Van Nes EH,
from the Neolithic age in China’s Yangtze River Delta. Mazzeo N, Stemberg LDSL, Scheffer M, Cole JJ (2010)
Naturwissenschaften 93:232–236 Climate-dependent CO2 emissions from lakes. Global
Cao ZH, Yang LZ, Lin XG, Hu ZY, Dong YH, Zhang GY et al Biogeochem Cy 24. doi:10.1029/2009GB003618
(2007) Morphological characteristics of paddy fields, pad- Kroger N, Lorenz S, Brunner E, Sumper M (2002) Self-
dy soil profile, phytoliths and fossil rice grain of the Neo- assembly of highly phosphorylated silaffins and their func-
lithic age in Yangtze River Delta. Acta Pedologica Sin tion in biosilica morphogenesis. Science 298:584–586
44:839–847 (In Chinese) Liang Y, Hua H, Zhu YG, Cheng C, Romheld V (2006) Impor-
Chen JG, Zhang YZ, Zeng XB, Zhou WJ, Zhou J (2008) Effect tance of plant species and external silicon concentration to
of long-term various fertilization on exchangeable Ca and active silicon uptake and transport. New Phytol 172:63–72
Mg, and available S and Si contents in paddy soils. Ecol Lin XG, Yin R, Zhang HY, Huang JF, Chen RR, Cao ZH (2004)
Environ 17:2064–2067 Changes of soil microbiological properties caused by land
China Sannong Data NetWork (CSDN) (2011) http:// use changing from rice-wheat rotation to vegetable culti-
www.sannong.gov.cn vation. Environ Geochem Health 26:119–128
China Soil Scientific Database (CSSD) (2012) http://mirror. Lu RK (2000) Methods for soil and agrochemical analysis.
soil.csdb.cn/page/showItem. vpage?id=cn. csdb. soil. tax- China Agriculture Press, Beijing
onomy. cst Yagang/ Ma JF, Takahashi E (2002) Amsterdam: Elsevier Science; Soil,
Clark DA (2002) Are tropical forests an important carbon sink? fertilizer, and plant silicon research in Japan, 1st edn.
Reanalysis of the long-term plot data. Ecol Appl 12:3–7 Elsevier Science, Amsterdam
Ding TP, Ma GR, Shui MX, Wan DF, Li RH (2005) Silicon Ma JF, Tamai K, Ichii M, Wu K (2002) A rice mutant defective
isotope study on rice plants from Zhejiang Province, Chi- in active Si uptake. Plant Physiol 130:2111–2117
na. Chem Geol 218:41–50 Matichenkov V, Calvert D, Snyder G (1999) Silicon fertilizers
DOE (2008) International Energy Outlook 2008 Energy Infor- for citrus in Florida. Proc Fla State Hort Soc 112:5–8
mation Administration Office of Integrated Analysis and McKenzie N, Ryan P, Fogarty P, Wood J (2000) Sampling,
Forecasting. US. Department of Energy, Washington, DC measurement and analytical protocols for carbon estima-
Epstein E (1994) The anomaly of silicon in plant biology. Proc tion in soil, litter and coarse woody debris, Australian
Natl Acad Sci USA 91:11–17 Greenhouse Office, National Carbon Accounting System,
Epstein E (1999) Silicon. Annu Rev Plant Physiol Plant Molec Technical Report no 14:1–42
Biol 50:641–644 Mecfel J, Hinke S, Goedel WA, Marx G, Fehlhaber R, Bǎucker
Epstein E (2001) Silicon in plants: facts vs. concepts. In: E, Wienhaus O (2007) Effect of silicon fertilizers on
Datnoff LE, Snyder GH, Korndorger GH (eds) Silicon in silicon accumulation in wheat. J Plant Nutr Soil Sci
agriculture). Elsevier Science B.V, Amsterdam, pp 1–15 170:769–772
Falkowski P, Scholes RJ, Boyle E, Canadell J, Canfield D, Elser J, Mulholland SC, Prior CA (1993) AMS radiocarbon dating of
Gruber N, Hibbard K, Högberg P, Linder S, Mackenzie FT, phytoliths. In: Pearsall DM, Piperno DR (eds) MASCA
Moore B III, Pedersen T, Rosenthal Y, Seitzinger S, Smetacek research papers in science and archaeology. University of
V, Steffen W (2000) The global carbon cycle: a test of our Pennsylvania, Philadelphia, pp 21–23
knowledge of earth as a system. Science 290:291–296 Murphy D (2002) Fundamentals of light microscopy and elec-
Fang JY, Guo ZD, Piao SL, Chen AP (2007) Terrestrial vegeta- tronic imaging. A John Wiley & Sons, Inc., 121 pp
tion carbon sinks in China, 1981–2000. Sci China Ser D- National Bureau of Statistics of China (NBSC) (2011) http://
Earth Sci 50:1341–1350 www.stats.gov.cn/
Field CB (2001) Plant physiology of the “Missing” carbon sink. Oldenburg CM, Torn MS, DeAngelis KM, Ajo-Franklin JB,
Plant Physiol 125:25–28 Amundson RG, Bernacchi CJ, et al. (2008) Biologically
Gifford RM (1994) The global carbon cycle: a viewpoint on the enhanced carbon sequestration: Research needs opportuni-
missing sink. Aust J Plant Biol 21:1–15 ties. Report on the Energy Biosciences Institute Workshop
Harrison K, Broecker W, Bonani G (1993) A strategy for esti- on Biologically Enhanced Carbon Sequestration, October
mating the impact of CO2 fertilization on soil carbon 29 2007, Berkeley, CA, LBNL–713E
storage. Global Biogeochem Cy 7:69–80 Parr JF (2006) Effect of fire on phytolith coloration. Geo-
Hart DM, Humphreys GS (1997) Plant opal phytoliths: an archaeology 21:171–185
Australian perspective. Quatern Aust 15:17–25 Parr JF, Sullivan LA (2005) Soil carbon sequestration in
Intergovernmental Panel on Climate Change (IPCC) (2007) phytoliths. Soil Biol Biochem 37:117–124
Climate change 2007: the scientific basis. Cambridge Uni- Parr JF, Sullivan LA (2011) Phytolith occluded carbon and silica
versity Press, UK variability in wheat cultivars. Plant Soil 342:165–171
Plant Soil (2013) 370:615–623 623

Parr JF, Dolic V, Lancaster G, Boyd WE (2001) A microwave silicon and carbon: Implications for biogeochemical carbon
digestion method for the extraction of phytoliths from sequestration. Earth Sci Rev 319–331
herbarium specimens. Rev Palaeobot Palynol 116:203–212 Strömberg CAE (2004) Use phytolith assemblages to reconstruct
Parr JF, Sullivan LA, Quirk R (2009) Sugarcane phytoliths: the origin and spread of grass-dominated habitats in the great
encapsulation and sequestration of a long-lived carbon plains of North America during the late Eocene to early
fraction. Sugar Tech 11:17–21 Miocene. Paleogeogr Paleoclimatol Paleoecol 207:239–275
Parr JF, Sullivan LA, Chen B, Ye G, Zheng W (2010) Carbon Walkley A, Black IA (1934) An examination of the Degtjareff
bio-sequestration within the phytoliths of economic bam- method for determining soil organic matter, and a proposed
boo species. Global Change Biol 16:2661–2667 modification of the chromic acid titration method. Soil Sci
Pearsall DM (1989) Paleoethnobotany: a handbook of proce- 37:29–38
dures. Academic, London Wang SM, Zhang CH, Hu FX, Zeng K, Zhang WH, Wang WJ
Perry CC, Williams RJP, Fry SC (1987) Cell wall biosynthesis (2008) The quantitative analysis of rice aboveground bio-
during silicification of grass hairs. J Plant Physiol 126:437– mass and net primary productivity. Chinese Agr Sci Bull
448 24:201–205 (In Chinese with English abstract)
Piperno DR (1988) Phytolith analysis: an archaeological and Wilding LP (1967) Radiocarbon dating of biogenetic opal.
geological perspective. Academic, London Science 156:66–67
Prasad V, Strömberg CAE, Alimohammadian H, Sahni A (2005) Wilding LP, Drees LR (1974) Contributions of forest opal and
Dinosaur coprolites and the early evolution of grasses and associated crystalline phases to fine silt and clay fractions
grazers. Science 310:1177–1180 of soils. Clay Clay Miner 22:295–306
Sangster AG, Parry DW (1981) Ultrastructure of silica deposits Wilding LP, Brown RE, Holowaychuk N (1967) Accessibility
in higher plants. In: Simpson TL, Volcani BE (eds) Silicon and properties of occluded carbon in biogenetic opal. Soil
and siliceous structures in biological systems. Springer, Sci 103:56–61
New York, pp 383–407 Zhang YL, Yu L, Liu MD, Yu N (2008) Silicon liberation
Schimel D (1995) Terrestrial ecosystems and the carbon cycle. characteristics of soil and its effect factors after applying
Global Change Biol 1:77–91 slag mucks I Relationships between Calcium, Magnesium,
Schlesinger WH (1997) Biogeochemistry: an analysis of global Iron and Aluminium and Silicon liberation. Chinese J Soil
change. Academic, New York Sci 39:722–725 (In Chinese)
Skjemstad JO, Spouncer LR, Beech A (2000) Carbon conver- Zhang WJ, Wang XJ, Xu MG, Huang SM, Peng C (2010) Soil
sion factors for historical soil carbon data. 15, CSIRO Land organic carbon dynamics under long-term fertilizations in
and Water, Adelaide arable land of northern China. Biogeosciences 7:409–425
Song ZL, Liu HY, Si Y, Yin Y (2012a) The production of Zheng Y, Matsui A, Fujiwara H (2003) Phytoliths of rice
phytoliths in China’s grasslands: implications to the bio- detected in the Neolithic sites in the valley of the Taihu
geochemical sequestration of atmospheric CO2. Global Lake in China. Env Archaeol 8:177–184
Change Biol 18:3647–3653 Zuo XX, Lü HY (2011) Carbon sequestration within millet
Song ZL, Wang HL, Strong PJ, Li ZM, Jiang PK (2012b) Plant phytoliths from dry-farming of crops in China. Chinese
impact on the coupled terrestrial biogeochemical cycles of Sci Bull 56:3451–3456

You might also like