Front. Earth Sci.

2016, 10(4): 683–690

DOI 10.1007/s11707-015-0549-9

RESEARCH ARTICLE

Evaluation of the occluded carbon within husk phytoliths of

35 rice cultivars

Xing SUN1, Qin LIU (✉)1, Jie GU1,2, Xiang CHEN1, Keya ZHU1,3
1 Institute of Soil Science, Chinese Academy of Sciences, Nanjing 210008, China
2 College of Forest Resources & Environment, Nanjing Forestry University, Nanjing 210037, China
3 College of Resources and Environmental Sciences, Nanjing Agricultural University, Nanjing 201195, China

© Higher Education Press and Springer-Verlag Berlin Heidelberg 2015

Abstract Rice is a well-known silicon accumulator. 1 Introduction

During its periods of growth, a great number of phytoliths
are formed by taking up silica via the plant roots. The soil carbon pool plays a vital role in the global carbon
Concurrently, carbon in those phytoliths is sequestrated cycle (Lal, 2004; Liu et al., 2014). The soil carbon storage
by a mechanism of long-term biogeochemical processes capacity is influenced by many factors, such as climate
within the plant. Phytolith occluded C (PhytOC) is very (temperature and precipitation), the nature of parent
stable and can be retained in soil for longer than a geological materials (i.e., texture and mineralogy), vegeta-
millennium. In this study, we evaluated the carbon bio- tion (crop) type, and land management practices (McCarl
sequestration within the phytoliths produced in rice seed et al., 2007). Approximately half of all soil carbon in
husks of 35 rice cultivars, with the goal of finding rice managed ecosystems has been lost into the atmosphere
cultivars with relatively higher phytolith carbon sequestra- during the past two centuries due to cultivation. Thus,
tion efficiencies. The results showed that the phytolith carbon sequestration in the agricultural ecological system
contents ranged from 71.6 mg$g‒1 to 150.1 mg$g‒1, and the is a popular subject of study (Linquist et al., 2012). Modern
PhytOC contents ranged from 6.4 mg$g‒1 to 38.4 mg$g‒1, crop production technologies are currently being adopted
suggesting that there was no direct correlation between the to reduce agricultural greenhouse gas emissions in these
PhytOC content and the content of rice seed husk rice growing areas, such as early maturing cultivars, high
phytoliths (R = 0.092, p > 0.05). Of all rice cultivars, six use of inorganic fertilizers and pesticides, and the
showed a higher carbon sequestration efficiency in expansion of irrigation facilities (Pathak et al., 2005).
phytolith seed husks. Additionally, the carbon bio- However, potential processes to effectively select and
sequestration within the rice seed husk phytoliths was breed high C occlusion efficiency cultivars have been
approximately 0.45‒3.46 kg-e-CO2$ha‒1$yr‒1. These rates disregarded thus far. Organic carbon can be occluded
indicate that rice cultivars are a potential source of carbon within phytoliths (i.e., PhytOC) during the course of plant
biosequestration which could contribute to the global growth (Parr and Sullivan, 2005). The contents of
carbon cycle and climate change. phytoliths generally constitute up to 3% of the total soil
mass (Drees et al., 1989). PhytOC has been confirmed to
Keywords carbon sequestration, seed husks, PhytOC, be an important long-term terrestrial carbon fraction in the
phytolith, rice cultivars soil (Parr and Sullivan, 2005; Gu et al., 2012), contributes
15%‒37% of the stable soil C sink (Parr and Sullivan,
2005), and represents up to 82% of the soil carbon in
topsoils of in situ decomposition after 2000 years (Wild-
ing, 1967; Wilding et al., 1967; Mulholland and Prior,
1993; Parr and Sullivan, 2005).
Phytolith, the opaline amorphous silica, is absorbed by
-

plants and is formed in the tissues of most plants (Perry

et al., 1987; Piperno, 1988; Song et al., 2012a,b), such as
Received March 6, 2015; accepted July 5, 2015 bamboo, rice, wheat, etc. After plants return to the soil, the
E-mail: [email protected] straws of these plants are decomposed by microorganisms,
684 Front. Earth Sci. 2016, 10(4): 683–690

and the phytoliths within these straws can be released 2 Methods and materials
directly into the soil. Phytoliths can tolerate extreme
environments, such as earthquakes, dust storms, floods, 2.1 Sample preparation
forest fire erosion, etc., and are often retained in the in situ
environment (Jones and Milne, 1963; Jones and Handreck, Seed samples of 35 rice cultivars, which belong to seven
1967; Wilding, 1967; Wilding and Drees, 1974; Sangster varieties, were randomly obtained from six provinces
and Parry, 1981; Pearsall, 1989; Hart and Humphreys, (Jiangxi, Jiangsu, Henan, Heilongjiang, Anhui, and
1997; Parr, 2006; Bowdery, 2007). However, during Zhejiang) in China in June 2013. These seed samples
phytolith formation in plant tissues, some elements can were derived from the conventional cultivars within the
be occluded (Jones and Milne, 1963), such as C, N, P, K, locality, as presented in Table 1. The seed samples were
Ni, Al, Fe, Ca, etc. (Wang, 1998). In this course, phytoliths weighed at 5 g and placed in respective envelopes. The
can occlude 1%‒6% organic C (Wilding et al., 1967; Parr seed samples were then dried in an oven and removed after
and Sullivan, 2005; Parr et al., 2010; Song et al., 2014a). 8 hours for hulling. The seed husk samples were preserved
The PhytOC content of plants in different crop species for extracting phytoliths and analyzing PhytOC and
ranges from 0.04% to 0.75% of the dry crop biomass (Parr PhytON.
and Sullivan, 2011; Li et al., 2013). However, the PhytOC
content also varies significantly between different cultivars 2.2 Phytolith extraction from seed husks
(Parr and Sullivan, 2011; Li et al., 2013) and tissues (Li et
al., 2013) of the same crop species. Many studies The phytolith extraction method was designed to com-
demonstrate that the PhytOC content in crops depends pletely remove all organic material from the seed husk of
on the C content of phytoliths (Parr and Sullivan, 2011; each rice species (Walkley and Black, 1934; Zuo and Lü,
Zuo and Lü, 2011) and on the phytolith content of crops 2011). In this study, a revision to the wet digestion method
(Li et al., 2013). was implemented to digest the organic matter more
Rice, which acts as a food source as well as a silica completely (Walkley and Black, 1934). The detailed
accumulator, has many phytoliths and can occlude more steps are as follows: 1) approximately 1.0 g of dry sample
organic C in its organs when compared with other plants was weighed in a centrifuge tube (to the nearest 0.1 mg); 2)
(Song et al., 2012a,b). The global rice-planting area was 5 mL of HNO3 was added to the tube and heated in a water
approximately 1.55!108 ha in 2010 (IRRI, 2011). Rice is a bath at 100°C until the reaction stopped; this was repeated
major crop in Asian countries, such as India and China. twice, and then the sample was centrifuged at 3,000 r/min
From 1950 to 2010, approximately 2.37 ! 108 Mg of CO2 for 5 min and the supernatant was decanted; 3) the sample
equivalents may have been sequestrated within rice was rinsed with 10 mL of distilled water, centrifuged at
phytoliths in China (Li et al., 2013). Assuming a maximum 3,000 r/min for 5 min and decanted, followed by two
phytolith C biosequestration flux of 0.13 Mg-e- additional rinses; 4) 10% HCl was then added, and the
CO2$ha–1$yr–1, the global annual potential rate of CO2 mixture was heated in a water bath at 80°C for 30 min,
sequestrated in rice phytoliths would be approximately centrifuged at 3,000 r/min for 5 min, and the supernatant
1.94 ! 107 Mg (Li et al., 2013), which is greater than was then decanted; 5) the sample was rinsed with 10 mL of
bamboo leaf litter (1.56!107 Mg-e-CO2 per year) (Parr distilled water, centrifuged at 3,000 r/min for 5 min and
et al., 2010), sugarcane leaf (0.72!106 Mg-e-CO2 per decanted, followed by two additional rinses; 6) 5 mL
year) (Parr et al., 2009), and millet (2.37!106 Mg-e-CO2 HNO3, was added, and the mixture was heated in a water
per year) (Zuo and Lü, 2011). bath at 100°C until the reaction stopped to ensure the
The presence of silica phytoliths within cereal crops is removal of all organic material and then centrifuged twice
well-documented (Yoshida et al., 1962; Ball et al., 1993, at 3,000 r/min for 5 min, followed by decanting of the
1996; Lux et al., 2002; Song et al., 2014a,b). According to supernatant; 7) 5 mL of H2SO4 was added, and the mixture
the correlations between phytolith content and PhytOC was heated in a water bath at 100°C for 1 h; 8) the mixture
content in crops, results show that the PhytOC content of was cooled to room temperature, and then 30% H2O2 was
plants can be increased by crop species or cultivar slowly added until the liquid cleared; and 9) the mixture
optimization (Li et al., 2013; Song et al., 2014a, b). was centrifuged four times at 3,000 r/min for 5 min,
However, the variability of phytolith accumulation within followed by decanting of the supernatant, and finally, the
the many cultivars of rice has not previously been phytoliths were dried in an oven at 70°C for 24 h.
examined. In this study, we evaluated carbon biosequestra-
tion within the phytoliths produced in the rice seed husks 2.3 PhytOC and PhytON measurements
of 35 rice cultivars. The aim of the study is to provide a
choice for farmers to plant the rice cultivars with a high Phytolith residue assemblages were mounted onto glass
efficiency of carbon sequestration in phytoliths as well as slides in a Balsam Canada mounting medium. The slides
high productivity. were viewed at 400 ! magnification using a Jiangnan XP-
 Xing SUN et al. Evaluation of the occluded carbon within husk phytoliths 685

Table 1 The 35 tested rice cultivars collected from six provinces in

China (the 35 cultivars belong to seven varieties)
Number Cultivars Varieties
1 Zhonghan 35 Indica rice
2 Xinliangyou 6 Indica two line hybrid rice
3 Tianyou 998 Indica three line hybrid rice
4 Gangyou 188
5 II you 1259
6 IIyou 501
7 Fuyou 21
8 Chuannong 1
9 Zhongyou 7
10 Yixiang 2079
Fig. 1 The phytoliths extracted by the wet Ashing Method
11 IIyou 1313 followed by Walkley and Black (1934) and Zuo and Lü (2011)
12 Yixiang 725 from the rice husk samples.
13 Tenuo 2072 Indica glutinous rice
14 Zhenzhunuo
2.4 Statistical analyses
15 Huaidao 5 Japonica rice
The mean values of all parameters were taken from the
16 Huaidao 11 measurements of three replicates, and the standard error of
17 Lianjing 11 the means was calculated. One-way ANOVA was applied
18 Yanjing 47‒12 to determine the significance of the results between
19 Wuyunjing 21
different varieties, and then Turkey’s multiple range tests
(p < 0.05) were performed. All of the statistical analyses
20 Zhengdao 18
were performed using SPSS v.13 for Windows.
21 Fengguan 16
22 Longjing 38
23 Longdao 12 3 Results
24 Longjing 21
25 Longjing 36
The results between the husk contents, phytolith contents,
and PhytOC content had considerable variation among the
26 Suidao 3
35 cultivars (Table 2). There was an even greater variation
27 Jixidao 1 in the phytolith contents in the seed husks of the cultivars,
28 Nanjing 5055 ranging from 71.6 mg$g‒1 to 150.1 mg$g‒1. Similarly,
29 Nanjing 46 substantial variations were observed in the PhytOC
(0.64 mg$g‒1 to 3.84 mg$g‒1) and PhytON (0.31 mg$g‒1
30 Xiushui 09
to 0.59 mg$g‒1) contents in the seed husks of the 35 rice
31 Zhejing 88 cultivars. Of all the rice cultivars, the highest carbon
32 Xiushui 134 sequestration within the phytoliths was observed in the
33 8you 682 Japonica three line hybrid rice Zhonghan 35 variety, followed by Tianyou 998, Tenuo
34 Sujingnuo 1 Japonica glutinous rice 2072, Huaidao 5, Yanjing 47‒12, and Longjing 38. Their
values were (3.22, 3.46, 2.84, 2.92, 2.90, and 2.57) Mg-e-
35 Zhenou 65
CO2$ha‒1$yr‒1, respectively (Table 2).
The relationships between the seven variables of the 35
rice cultivars are shown in Table 3. No correlation existed
213 microscope fitted with a polarizing filter and a 5.0 MP between the phytolith content and the PhytOC content in
color CCD camera to ensure that no organic material exists the phytoliths of the 35 rice cultivars (R = 0.092, p > 0.05).
(Fig. 1). The cross-polarized light technique was used here However, the phytolith content was correlated with the
to indicate the absence of cellulose, calcium oxalate PhytOC content in the seed husks (R = 0.380, p < 0.05)
crystals, and starch grains, which appear bright white and and with the estimated fluxes of seed husk PhytOC (R =
blue if present (Parr et al., 2010). The PhytOC and PhytON 0.344, p < 0.05). Additionally, the PhytOC content in the
were determined using an Elemental Analyzer 3000 phytoliths exhibited a strong positive correlation with the
(GmbH Company, Germany). PhytOC content in the seed husks (R = 0.865, p < 0.01)
686 Front. Earth Sci. 2016, 10(4): 683–690

Table 2 Rice cultivars, seed husk content as a percentage of rice, content of phytoliths, PhytON and PhytOC in the seed husk on a dry weight
basis, and the estimated fluxes of PhytOC per ha in Mg of CO2 equivalents (Mg-e-CO2) for rice (according to gain yields of single rice crops of
9.28 Mg$ha‒1*)
Cultivars Husk Weight Phytolith content PhytON content PhytOC content PhytOC content C/N of phytoliths Estimated fluxes of
/% /(mg$g‒1) in phytoliths in phytoliths in seed husks PhytOC in seed husks
/(mg$g‒1) /(mg$g‒1) /(mg$g‒1) /(kg-CO2$ha‒1$yr‒1)
Zhonghan 35 22.46 109.6"22.5 0.33"0.01 3.84"0.44 0.47"0.09 11.83"1.60 3.22"0.37
Xinliangyou 6 22.06 117.8"8.4 0.48"0.05 0.89"0.15 0.11"0.03 1.85"0.43 0.79"0.13
Tianyou 998 24.26 132.2"6.3 0.39"0.03 3.16"0.84 0.43"0.11 9.81"2.83 3.46"0.92
Gangyou 188 18.60 111.2"11.0 0.37"0.03 0.64"0.08 0.07"0.01 1.74"0.25 0.45"0.06
II you 1259 22.15 128.9"16.3 0.36"0.02 0.88"0.15 0.12"0.03 2.43"0.42 0.86"0.15
IIyou 501 25.16 129.3"5.1 0.45"0.02 1.72"0.83 0.22"0.11 3.81"1.71 1.91"0.92
Fuyou 21 22.53 98.8"6.5 0.49"0.03 2.32"0.35 0.23"0.02 4.73"0.51 1.76"0.27
Chuannong 1 19.67 71.6"7.6 0.50"0.09 1.10"0.14 0.08"0.01 2.22"0.19 0.53"0.07
Zhongyou 7 19.79 97.6"8.5 0.53"0.06 1.07"0.46 0.10"0.04 2.00"0.82 0.70"0.30
Yixiang 2079 22.77 118.5"4.9 0.43"0.10 1.07"0.64 0.13"0.08 2.39"1.05 0.98"0.59
IIyou 1313 20.25 110.8"3.1 0.51"0.07 1.26"0.37 0.14"0.04 2.44"0.63 0.96"0.28
Yixiang 725 22.65 132.4"16.2 0.40"0.03 0.66"0.09 0.09"0.02 1.65"0.22 0.67"0.09
Tenuo 2072 19.83 148.5"31.4 0.34"0.06 2.83"0.28 0.42"0.10 8.52"1.09 2.84"0.28
Zhenzhunuo 26.65 113.9"56.9 0.34"0.04 0.89"0.12 0.11"0.06 2.63"0.60 0.92"0.12
Huaidao 5 15.32 150.1"5.5 0.37"0.06 3.73"0.68 0.56"0.14 9.82"2.35 2.92"0.53
Huaidao 11 16.74 138.9"31.8 0.48"0.17 2.22"0.55 1.09"1.01 1.09"1.01 1.76"0.44
Lianjing 11 16.09 127.1"14.4 0.31"0.03 2.62"0.48 0.33"0.04 0.33"0.04 1.83"0.33
Yanjing 47-12 19.57 148.5"2.8 0.32"0.05 2.93"0.30 0.44"0.05 0.44"0.05 2.90"0.30
Wuyunjing 21 13.63 122.5"20.6 0.35"0.01 3.41"0.27 0.40"0.06 0.40"0.06 1.94"0.15
Zhengdao 18 16.60 131.0"55.9 0.37"0.07 0.80"0.09 0.10"0.04 0.10"0.04 0.59"0.07
Fengguan 16 17.84 79.4"6.4 0.54"0.03 3.09"0.49 0.25"0.05 0.25"0.05 1.49"0.24
Longjing 38 17.98 135.0"16.1 0.46"0.05 3.10"0.58 0.42"0.06 0.42"0.06 2.57"0.48
Longdao 12 18.14 149.9"7.9 0.43"0.02 2.01"0.15 0.30"0.03 0.30"0.03 1.86"0.14
Longjing 21 17.26 156.3"21.4 0.38"0.05 1.85"0.57 0.30"0.14 0.30"0.14 1.70"0.52
Longjing 36 16.63 119.8"10.6 0.49"0.03 2.05"0.55 0.25"0.08 0.25"0.08 1.39"0.37
Suidao 3 16.96 141.3"4.2 0.40"0.05 1.74"0.09 0.25"0.02 0.25"0.02 1.42"0.07
Jixidao 1 16.67 140.9"9.9 0.40"0.06 1.45"0.36 0.21"0.07 0.21"0.07 1.16"0.29
Nanjing 5055 15.40 136.2"9.7 0.35"0.05 1.14"0.23 0.15"0.02 0.15"0.02 0.81"0.16
Nanjing 46 14.72 105.7"4.2 0.55"0.07 2.26"0.33 0.24"0.15 0.24"0.15 1.19"0.17
Xiushui 09 15.10 125.5"1.6 0.45"0.07 1.72"0.40 0.22"0.05 0.22"0.05 1.11"0.26
Zhejing 88 15.14 146.5"10.5 0.54"0.07 1.48"0.31 0.22"0.06 0.22"0.06 1.12"0.23
Xiushui 134 17.39 138.4"4.1 0.53"0.04 1.12"0.02 0.16"0.00 0.16"0.00 0.92"0.02
8you 682 19.52 89.5"6.7 0.48"0.01 2.39"0.13 0.21"0.03 0.21"0.03 1.42"0.08
Sujingnuo 1 15.87 124.1"19.8 0.44"0.01 2.01"0.25 0.55"0.59 0.55"0.59 1.35"0.17
Zhenou 65 16.89 134.0"6.4 0.47"0.03 2.35"0.61 0.27"0.01 0.27"0.01 1.81"0.47

*Data offered by the Changshu Agroecological Experimental Station, Chinese Academy of Sciences.

and with the estimated fluxes of seed husk PhytOC (R = The husk content of Indica varieties ranged from
0.888, p < 0.01). There was a strong positive correlation (R 21.78% to 23.24%, and that of the Japonica varieties
= 0.855, p < 0.01) between the PhytOC content in the seed varied between 16.38% and 19.52%. Additionally, the
husks and the estimated fluxes of the seed husk PhytOC PhytOC content in the phytoliths of four Indica varieties
(Table 3). varied greatly between (0.89"0.15) mg$g‒1 and (3.84"
 Xing SUN et al. Evaluation of the occluded carbon within husk phytoliths 687

Table 3 Correlation coefficients between the seven variables of the 35 rice cultivars
Variables Husk weight Phytolith content PhytON content PhytOC PhytOC content in Estimated fluxes C/N of
in phytoliths content in seed husks of PhytOC in phytoliths
phytoliths seed husks
Husk weight 1
Phytolith contents – 0.237 1
PhytON content in phytoliths – 0.127 – 0.428* 1
PhytOC content in phytoliths – 0.204 0.092 – 0.180 1
PhytOC content in seed husks – 0.268 0.380* – 0.287 0.865** 1
Estimated fluxes of PhytOC in seed
husks 0.074 0.344* – 0.351* 0.888** 0.855** 1

C/N of phytoliths – 0.148 0.216 – 0.143 0.539** 0.582** 0.528** 1

* Correlation is significant at the 0.05 level (2-tailed).

** Correlation is significant at the 0.01 level (2-tailed).

0.44) mg$g‒1; significant differences in the PhytOC husk PhytOC content (R = 0.855, p < 0.05), with the
content of the phytoliths were observed among the seven estimated fluxes of the seed husk PhytOC (R = 0.930,
cultivars of the Indica varieties. However, the PhytOC p < 0.01), and with the phytolith C/N (R = 0.966,
content in the phytoliths of three Japonica varieties ranged p < 0.01). In addition, the seed husk PhytOC content
from 2.15 mg$g‒1 to 2.39 mg$g‒1 (Table 4). Although the exhibited a strong positive correlation with the estimated
husk average content of the Indica varieties was much fluxes of the seed husk PhytOC (R = 0.840, p < 0.01) and
higher than that of the Japonica varieties, the PhytOC with the phytolith C/N (R = 0.833, p < 0.05). There was a
content in the phytoliths of Indica varieties was on average strong positive correlation between the estimated fluxes of
lower than that in Japonica varieties. In summary, the seed seed husk PhytOC and the phytolith C/N (R = 0.990,
husk PhytOC content exhibited no significant differences p < 0.01) (Table 5).
between the two rice varieties.
The relationships between the phytolith content, the
phytolith PhytOC content, the seed husk PhytOC content, 4 Discussion
the phytolith C/N, and the estimated fluxes of seed husk
PhytOC were examined. The results indicated that a weak 4.1 Carbon sequestration within the phytoliths of rice seed
negative correlation existed between the phytolith content husks
and the PhytOC content in the phytoliths for seven rice
varieties (R = – 0.242, p > 0.05), whereas the phytolith The PhytOC content in bamboo, wheat, sugarcane, and
content was strongly positively correlated with the seed millet was found to have no direct correlation with the

Table 4 Rice varieties, the seed husk content as a percentage of rice, the content of phytoliths, PhytON and PhytOC in seed husk on a dry weight
basis and the estimated fluxes of PhyOC per ha in Mg of CO2 equivalents (Mg-e-CO2) for rice (according to gains in yields of single rice crops of
9.28 Mg$ha‒1*)
Varieties n Husk Weight Phytolith PhytON content PhytOC content PhytOC C/N of Estimated fluxes of
/% content in phytoliths in phytoliths content in phytoliths PhytOC in seed husks
/(mg$g‒1) /(mg$g‒1) /(mg$g‒1) seed husks /(kg-CO2 $ha‒1$yr‒1)
/(mg$g‒1)
Indica rice 1 22.46"0.00 109.6"22.5 0.33"0.01 3.84"0.44 0.47"0.09 11.63"1.60 3.22"0.37
Indica two line hybrid rice 1 22.06"0.00 117.8"8.4 0.48"0.05 0.89"0.15 0.11"0.03 1.85"0.43 0.79"0.13
Indica three line hybrid 10 21.78"2.13 113.1"19.6 0.44"0.08 1.39"0.80 0.16"0.11 3.16"1.98 1.22"0.92
rice
Indica glutinous rice 2 23.24"4.82 131.2"24.5 0.34"0.05 1.86"1.37 0.26"0.22 5.47"4.03 1.88"1.35
Japonica rice 18 16.51"1.45 132.9"18.4 0.43"0.08 2.15"0.84 0.28"0.12 5.25"2.54 1.59"0.67
Japonica three line hybrid 1 19.52"0.00 89.5"6.7 0.48"0.01 2.39"0.13 0.21"0.03 4.98"0.92 1.42"0.08
rice
Japonica glutinous rice 2 16.38"0.72 129.1"7.0 0.46"0.02 2.18"0.24 0.41"0.20 4.78"0.31 1.58"0.33

n, number of plant species samples.

*Data offered by Changshu Agroecological Experimental Station, Chinese Academy of Sciences.
688 Front. Earth Sci. 2016, 10(4): 683–690

Table 5 Correlation coefficients between the seven variables of the seven varieties
Variables Husk weight Phytolith PhytON content PhytOC content PhytOC content in Estimated fluxes of C/N of phytoliths
content in phytoliths in phytoliths seed husks PhytOC in seed husks
Husk weight 1
Phytolith content – 0.245 1
PhytON content in phy- – 0.497 – 0.261 1
toliths
PhytOC content in phy- – 0.054 – 0.242 – 0.581 1
toliths
PhytOC content in seed – 0.261 0.181 – 0.550 0.855* 1
husks
Estimated fluxes of 0.199 – 0.041 – 0.824* 0.930** 0.840* 1
PhytOC in seed husks
C/N of phytoliths 0.140 – 0.130 – 0.751 0.966** 0.833* 0.990** 1

* Correlation is significant at the 0.05 level (2-tailed).

** Correlation is significant at the 0.01 level (2-tailed).

phytolith content formed within plants (Parr et al., 2009, fluxes of rice seed husks were 0.45 to 3.46 kg-e-CO2$ha‒1
2010; Parr and Sullivan, 2011; Zuo and Lü, 2011). Our per year in terms of the PhytOC content of rice seed husk
results indicated that there was no correlation between the material on a dry weight basis (Wang et al., 2008) and the
phytolith content or the PhytOC content in the phytoliths mean annual grain production (9.3 Mg-ha‒1) of single rice
of the 35 rice cultivars studied (R = 0.092, p > 0.05). These cropping systems (Anthoni et al., 2004; Zhang et al.,
data indicate that for rice, as for wheat, bamboo, and 2010). However, Li et al. (2013) reported that the flux of
sugarcane (Parr et al., 2009, 2010; Parr and Sullivan, the rice PhytOC was 0.03‒0.13 t-e-CO2$ha‒1 per year,
2011), it is the efficiency by which carbon is encapsulated which is much higher than our results. The differences
by silica within the epidermal cell walls (phytoliths), rather between this study and Li et al.’s findings may be due to
than the actual quantity of the silica taken up by the plant, the tested cultivars and the organs (i.e., this manuscript
that is most important in determining the relative PhytOC only tested the husk, and Li et al. tested four organs). This
yield in plant materials (Parr and Sullivan, 2011). The study used an organ to estimate the PhytOC flux.
finding that the carbon was occluded within rice husk The global rice-planting area was approximately
phytoliths was consistent with observations made by the 1.55!108 ha in 2010 (IRRI, 2011). Assuming that the
aforementioned researchers. In fact, the PhytOC content largest PhytOC flux of rice seed husk was 3.46 kg-e-
depends on the efficiency of the C occluded within the CO2$ha‒1$yr‒1, 5.36 ! 105 Mg-e-CO2 would be globally
phytoliths during plant growth. Genetic and physiological occluded within rice seed husk phytoliths per year.
differences within the rice cultivars may affect the
formation and the efficiency of the C occluded within the
phytoliths (Parr et al., 2009, 2010; Parr and Sullivan, 5 Conclusions
2011). Therefore, it is possible to improve the carbon
sequestration of the phytoliths by selecting cultivars with The carbon content of phytoliths in rice seed husks
high phytolith content and high C occluded in phytoliths depends on the efficiency of the carbon occluded within
from different rice cultivars. the phytoliths during rice growth periods. The PhytOC
content in the husk of 35 rice cultivars ranged from 0.64
mg$g‒1 to 3.84 mg$g‒1. It is estimated that the flux of C
4.2 The efficiency of carbon sequestration within the occluded within the rice husk phytoliths is between 0.45
phytoliths in rice seed husks and 3.46 kg-e-CO2$ha‒1 per year. Of the 35 rice cultivars
considered, six exhibited relatively higher phytolith and
The long-term carbon sequestration of soil organic carbon PhytOC accumulation efficiencies. Therefore, enhancing
is relatively low (only 0.7% of net primary millennia) the carbon sequestration efficiency of rice cultivar
(Schlesinger, 1990). In addition, many carbon fractions phytoliths will provide us with a new approach for
involved in the carbon cycle, either directly or indirectly, increasing the soil organic carbon sink, and selecting/
are responsible for environmental issues. However, the breeding rice cultivars with high PhytOC represents critical
occlusion of carbon within phytoliths has been retained in work in this domain.
soils for more than a millennium resulting in the formation
of a long-term, terrestrial carbon fraction (Parr and Acknowledgements This work was partially supported by the National
Sullivan, 2005). In this study, we estimated that the C Natural Science Foundation of China (Grant No. 41271208), the Jiangsu
 Xing SUN et al. Evaluation of the occluded carbon within husk phytoliths 689

Planned Projects for Postdoctoral Research Funds (No. 1301061C), the China phytoliths. In: Pearsall D M, Piperno D R, eds. MASCA Research
Postdoctoral Science Foundation funded project (No. 2013M541744), and Papers in Science and Archaeology. University of Pennsylvania,
the Key Projects in the National Science & Technology Pillar Program during
Philadelphia, 21–23
the Twelfth Five-year Plan Period (2013BAD11B00). We also express our
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